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FIELD-NATURALIST

Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

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Volume 128, Number | January—March 2014

: ; ;
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patron
His Excellency the Right Honourable David Johnston, C.C., C.M.M., C.O.M., C.M.
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments
of high quality for living things.
Honorary Members

Ronald E. Bedford Michael D. Cadman Christine Hanrahan Joyce M. Reddoch
Edward L. Bousfield Paul M. Catling C. Stuart Houston Allan H. Reddoch
Charles D. Bird Francis R. Cook Theodore Mosquin Dan Strickland
Fenja Brodo Bruce Di Labio Robert W. Nero John B. Theberge
Irwin M. Brodo Anthony J. Erskine E. Franklin Pope Sheila Thomson
Daniel F. Brunton John M. Gillett

2014 Council

President: Fenja Brodo Daniel F. Brunton Don Hackett Karen McLachlan-Hamilton
1° Vice-President: vacant Carolyn Callaghan David Hobden Lynn Ovenden

2" Vice-President: Jeff Skevington Barbara Chouinard Diane Kitching Rémy Poulin

Recording Secretary: Annie Bélair Owen Clarkin Diane Lepage Henry Steger

Treasurer: Ken Young Barry Cottam Ann MacKenzie Eleanor Zurbrigg

Ian Davidson

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Cover: Melanistic Maritime Gartersnake, Thamnophis sirtalis pallidulus, from Big Tancook Island, Mahone Bay, Lunenburg

County, Nova Scotia, captured on 7 May 2012. Photographed after it shed its skin in the laboratory. Photo: R. Lloyd. See
pages 63-71 in this issue. 7

THE CANADIAN
FIELD-NATURALIST

Volume 128

2014

Volume 128
The Ottawa Field-Naturalists’ Club Transactions

Promoting the study and conservation
of northern biodiversity since 1880

THE OTTAWA FIELD-NATURALISTS’ CLUB

OTTAWA CANADA

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The Canadian Field-Naturalist

Volume 128, Number |

January—March 2014

Distribution and Abundance of Benthic Macroinvertebrates and
Zooplankton in Lakes in Kejimkujik National Park and National
Historic Site of Canada, Nova Scotia

CHRISTINA NUSSBAUMER!:?}5, NEIL M. BURGESS?, and Russ C. WEEBER4

Ecotoxicology and Wildlife Health Division, Environment Canada, 45 Alderney Drive, Dartmouth, Nova Scotia B2Y 2N6
Canada

“Current address: Water Survey of Canada, Environment Canada, Room 854, 220 — 4th Ave. S.E., Calgary, Alberta T2G 4X3
Canada

“Ecotoxicology and Wildlife Health Division, Environment Canada, 6 Bruce Street, Mount Pearl, Newfoundland and Labrador
AIN 4T3 Canada

‘Canadian Wildlife Service (Ontario region), Environment Canada, 335 River Road, Ottawa, Ontario K1A 0H3 Canada

Corresponding author: christina.nussbaumer@ec.ge.ca

Nussbaumer, Christina, Neil M. Burgess, and Russ C. Weeber. 2014. Distribution and abundance of benthic macroinvertebrates and
zooplankton in lakes in Kejimkujik National Park and National Historic Site of Canada, Nova Scotia. Canadian Field-
Naturalist 128(1): 124.

As part of the Acid Rain Biomonitoring Program at Environment Canada, we sampled aquatic biodiversity in 20 acidic lakes in
2009 and 2010 in Kejimkujik National Park and National Historic Site of Canada and vicinity in Nova Scotia. We established
an inventory of current aquatic macroinvertebrate and zooplankton species composition and abundance in each of the 20 study
lakes. A total of 197 macroinvertebrate taxa were identified; the number of taxa observed was positively correlated with pH
across the 20 lakes. Acid-tolerant taxa, such as isopods, amphipods, trichopterans, and oligochaetes, were common and abundant,
while bivalves, gastropods, and leeches were lower in abundance. The number of isopods and amphipods collected was correlated
with calcium concentration; a greater proportion of isopods than amphipods were collected from lakes with low calcium and low
pH. Taxa with hard, calcareous shells, such as bivalves and gastropods, were not present in lakes with low calcium and low pH,
with bivalves occurring only in lakes above pH 4.9. Odonates and ephemeropterans, which were low in abundance, were
associated with a wide range of acidity. Coleopteran abundance was positively correlated with concentrations of dissolved
organic carbon. A total of 26 zooplankton taxa were collected, but only cyclopoid abundance was correlated with lake pH.
Results presented here provide a summary of aquatic biodiversity in lakes in Kejimkujik National Park and National Historic
Site and vicinity and provide a baseline for future monitoring as acid deposition continues to affect this acid-sensitive region
in Atlantic Canada.

Key Words: macroinvertebrates; Kejimkujik National Park and National Historic Site of Canada; water chemistry; acidic lakes:
zooplankton; Nova Scotia

In the 1980s, Environment Canada implemented the
Acid Rain Biomonitoring Program to study aquatic
invertebrate species assemblages in acid-sensitive Bore-
al Shield lakes in Ontario (McNicol et al. 1995b; Jef-
fries et al. 2004). In 2009 and 2010, this biomonitoring
program was expanded to include Kejimkujik National

Introduction

Acid deposition remains a widespread stressor of
freshwater ecosystems across southeastern Canada de-
spite legislated reductions in emissions of acidifying
pollutants over recent decades in both Canada and the
United States (Jeffries et al. 2004; Ginn et al. 2007).

Analyses of critical loads of acid deposition in eastern
Canada have suggested regions with carbonate-poor
geology continue to be influenced by acid inputs into
the environment (Doka et al. 2003; Jeffries et al. 2003;
Dupont et al. 2005; Clair et al. 2007, 2011). The effects
of acidification on the diversity of aquatic macroin-
vertebrate species have been well studied (e.g., Dermott
1985: Peterson 1987; Schell and Kerekes 1989; Lento e/
al. 2008), and changes in the composition of the aquat-
ic food web can have an impact on higher trophic levels
that rely on these groups for food (Weeber et al. 2004).

Park and National Historic Site of Canada, which has
a long history of environmental and ecological moni-
toring (Kerekes 1975; Kerekes ef a/. 1994; Burgess and
Hobson 2006; Wyn et al. 2010; Clair et al. 2011).

In the period from 2000 to 2007, the Kejimkujik
region in southwestern Nova Scotia received an aver-
age of 8 kg « ha‘! - year! to 12 kg - ha: year of SO,”
deposition (wet and dry) (Clair et a/. 2011). This level
is relatively low compared to the rest of North America.
However, the geology of Kejimkujik National Park and
National Historic Site consists mainly of poorly weath-

i)

erable bedrock that offers little buffering capacity,
and this makes this ecosystem extremely sensitive to
additional inputs of acid from the atmosphere (Clair
et al. 2007). In addition, the landscape in Kejimkuyjik
National Park and National Historic Site and the sur-
rounding area is composed of naturally acidic habitats
due to the prevalence of bog and fen wetlands. There-
fore, even with further reductions in atmospheric acid
deposition, recovery in these aquatic ecosystems is
expected to be extremely slow (Whitfield et a/. 2006;
Clair et al. 2011).

Although information on the status of and trends
in lake chemistry in Kejimkujik National Park and
National Historic Site is well developed (Clair ef al.
2011), only limited research has been completed on the
aquatic biodiversity in these acid-sensitive lakes
(Kerekes and Freedman 1989; Schell and Kerekes
1989). The purpose of this study was: (i) to determine
the current composition and abundance of aquatic

Peskawa

Kilometers
<t

THE CANADIAN FIELD-NATURALIST

Vol. 128

invertebrate and zooplankton in 20 acid-sensitive
lakes in Kejimkujik National Park and National His-
toric Site and vicinity and (ii) to identify potential
indicator taxa with respect to biological responses to
lake acidity.

Study Area

Kejimkujik National Park and Historic Site is a pro-
tected area of 404 km? located in southwestern Nova
Scotia (Figure 1). Twenty study lakes (17 within the
Park and 3 in the vicinity) were selected to cover a
range of water chemistry parameters. Lakes were cho-
sen to cover the largest possible gradients of acidity/
alkalinity, calcium, colour, and concentration of dis-
solved organic carbon in the study area. All of the 20
lakes were accessible by road or canoe (some back-
country lakes in Kejimkujik National Park and His-
toric Site are not accessible by road, so accessibility
was also a factor). Eight of the lakes were sampled in

‘ Kejimkujk National Park
\. Land National Historic Site}

4 lakes sampled June 2010 se

@ lakes sampled June 2009 (n=8)

yew

FIGURE 1. Location of 20 acid rain biomonitoring study lakes sampled during 2009 and 2010 in Kejimkujik National Park
and National Historic Site of Canada and surrounding area, Nova Scotia.

2014

June 2009 (Beaverskin, Big Dam East, George, Graf-
ton, North Cranberry, Pebbleloggitch, Peskawa, and
Puzzle) and the remaining 12 lakes were sampled in
June 2010 (Back, Ben, Big Dam West, Big Red, Don-
nellan, Frozen Ocean, Menchan, McGinty, Peskowesk,
Snake, Turtle, and Upper Silver).

Methods
Sampling methods

As part of an Environment Canada lake monitoring
network, surface water samples were collected by hel-
icopter from the centre of each lake during the spring
and fall turnover periods each year (usually May and
October) (Clair et al. 2011). Samples were collected
at a depth of 0.5 m, kept cool, and shipped overnight
to the Environment Canada Atlantic Laboratory for
Environmental Testing (ALET) in Moncton, New
Brunswick. At every 10th lake, triplicate samples were
collected and compared to each other for quality con-
trol. All water samples were analyzed in the laboratory
for various water chemistry parameters using unfiltered
water following ALET protocols (Clair et a/. 2011;
Eaton et al. 2012).

For the collection of aquatic macroinvertebrates and
zooplankton, we followed the sampling protocols of
the Environment Canada Acid Rain Biomonitoring
Program in Ontario and Quebec (see McNicol et al.
1995b). Sampling was completed in mid-June, as this
is a time of high invertebrate biomass and richness and
it is also when local waterbirds that depend on aquatic
prey to raise their young are breeding (McNicol et al.
1996).

At each study lake we conducted 10 benthic drag
samples, 10 water column sweeps, and 10 hoop sam-
ples, and we set 6 minnow traps (McNicol et al. 1996).
All samples were taken at randomly selected sites. Ben-
thic drag samples, which targeted odonates, ephemer-
opterans, bivalves, and gastropods, were conducted in
water less than | m in depth. A D-frame dip net (0.85
mm mesh) was dragged over the substrate for a dis-
tance of 0.5 m to collect the top 1—2 cm of substrate
(total sample area of 0.14 m*) (McNicol et al. 1996).
If boulders or rocky substrates made benthic drag sam-
pling impractical, a traveling kick and sweep sample
was completed instead. For these samples, the sampler
walked backwards for a distance of | m along the
shoreline (maximum | m depth), kicking the bottom
substrate and sweeping the dislodged detritus and in-
vertebrates into the D-frame net (Rosenberg ef al.
2000).

Both the benthic drag and the travelling kick and
sweep samples were processed in the same way: detri-
tus in the net was thoroughly rinsed to remove fine
sediments and was transferred to a sample container,

where it was first preserved with 10% buffered forma-
lin for 48 hours and then transferred into 70% ethanol.
Entire benthic samples were later sorted under a dis-
secting microscope. All observed macroinvertebrates
were removed and preserved in 70% ethanol.

NUSSBAUMER BAL, « MACROINVERTEBRATES AND ZOOPLANKTON IN KEJIMKUJIK NATIONAL PARK 3

Sweep sampling targeted nektonic invertebrates ac-
tive in the water column. Sweep sampling was con-
ducted in open water less than 5 m from the shore. Sam-
pling was completed by sweeping through the water
column in 10 consecutive ares using a D-frame dip net
(0.85 mm mesh, 625 cm? capture area) over the bow of
a forward-moving canoe traveling parallel to the shore-
line. Each sweep described an are from the water sur-
face down to a maximum depth of | m and back to the
surface, and a new section of the water column was
sampled with each are. Captured invertebrates were
picked from the net using forceps and transferred to a
sample container containing 70% ethanol.

Hoop sampling targeted trichopterans and gas-
tropods. A circular hoop of coated wire (diameter of
0.64 m, area of 0.32 m?) was placed on the substrate in
water <0.5 m deep. The hoop was visually searched for
a total of 5 minutes, and all invertebrates observed on
the surface of the substrate and vegetation were re-
moved and preserved in 70% ethanol.

All benthic macroinvertebrates from hoops, sweeps,
kick and sweep samples, and benthic drags were later
identified to species (or lowest taxonomic level possi-
ble).

Minnow traps targeted large nektonic invertebrates.
Six standard Gee’s minnow traps were baited with dry
dog food (Purina Puppy Chow®) and set for a total of
24 hours in near-shore sites where water depth was
approximately | m. Specimens were preserved in 70%
ethanol.

Zooplankton sampling was conducted at 15 of the
20 study lakes (5 of the study lakes were <2 m deep
and were therefore too shallow for vertical zooplank-
ton sampling to be carried out). A single vertical haul
was completed at the deepest part of each lake, starting
from 1 m above the sediment to the water’s surface.
Samples were collected using a non-metered zooplank-
ton net (80 um mesh, 26 cm in diameter). The contents
of the net were rinsed into the bottom of the collection
jar and then poured into a sample jar containing 33%
sugared, buffered formalin. All zooplankton samples
were identified to species (or lowest possible taxonom-
ic level).

Data analysis

Counts from all benthic invertebrate sampling pro-
cedures were pooled within each lake for the statistical
analyses. The resulting data from the 20 study lakes
were summarized with respect to mean, minimum, and
maximum counts for each species, as well as the per-
centage of lakes where a given species was observed.
Rare species (n = 72 taxa) were defined as occurring
in < 10% of the study lakes, while common species
(n = 125 taxa) occurred in > 10% of the study lakes.
The abundance and percentage composition of the most
abundant taxonomic groups were determined for each
lake, and boxplots where generated to show trends for
individual taxonomic groups of interest. Taxonomic
richness was calculated as the total number of unique
taxa in each lake.

4 THE CANADIAN FIELD-NATURALIST

Associations between water chemistry parameters,
as well as between the total number of macroinverte-
brate taxonomic groups and lake acidity, were evaluat-
ed using Spearman rank correlations. This non-para-
metric method of statistical analysis was employed as
some of the data did not meet assumptions of normal-
ity required for Pearson’s correlations. All statistical
analyses were completed using SYSTAT 13 (SYSTAT
Software Inc., Chicago, Illinois).

Zooplankton data were summarized by mean den-
sity (number of individuals/m*) for each of the 15 lakes,
and the percentage of lakes a given species was ob-
served in was also calculated.

Results

Fish were present in all 20 of the study lakes (Kere-
kes 1975; Drysdale et a/. 2005). Mean water chemistry
values for each lake are presented in Table 1. Many of
the study lakes were oligotrophic and darkly coloured
(99-202 Hazen units) due to dissolved organic com-
pounds leached from nearby bogs. Mean lake pH var-
ied from 4.3 (Big Red Lake) to 6.6 (McGinty Lake)
(Table 1). pH and calcium concentrations were positive-
ly correlated in the study lakes (r, = 0.747, P < 0.001);
pH and dissolved organic carbon were negatively cor-
related (7 ——0.7 IS; P= '0.001):

A total of 26 zooplankton species were observed in
the study lakes, with many of the common taxa ob-
served across a wide gradient of acidity (Supplemen-
tary Table 1). Only the abundance of Cyclopoida was

80
Pebbleloggitch
e

70

Big Dam West
e
Peskawa
e

60

50

Number of macroinvertebrate taxa

40

30

4.5

5.0

Vol. 128

significantly correlated with lake pH (r, = 0.536, P =
0.040).

A total of 197 taxa of aquatic macroinvertebrates
were observed (149 were identified to species, 38 to
genus, and 10 to family) (Supplementary Table 2). The
total number of taxa in each lake was positively cor-
related with both lake pH (Figure 2) (r, = 0.554, P =
0.011) and calcium concentrations (r, = 0.463, P =
(0.040). Taxon richness was not significantly associat-
ed with dissolved organic carbon (r, = —0.390, P =
(0.090). Total abundance (number of individuals of all
macroinvertebrates captured in each lake) was not
correlated with any water chemistry parameter.

The most abundant benthic invertebrate groups in
the 20 study lakes were Isopoda, Amphipoda, Oligo-
chaeta, and Trichoptera (Figures 3A and 3B). Only one
species of isopod was observed (Caecidotea commu-
nis), but it constituted up to 60% of the macroinver-
tebrates collected in some lakes (e.g., Peskowesk Lake)
(Figure 3A). The abundance of isopods (Caecidotea
communis) was lower in lakes with high pH and cal-
cium levels and higher in lakes with low calcium levels
(Figure 4A) (r, =—0.614, P = 0.004). Amphipods were
also abundant, with Hyalella azteca collected in 19 of
the 20 lakes. There was a significant positive relation-
ship between amphipod abundance and calcium levels
(Figure 4B) (r, = 0.776, P < 0.001). The proportion of
isopods relative to amphipods decreased with increas-
ing lake pH and calcium, with two exceptions (Big
Dam East Lake and Turtle Lake) (Figure 4C).

Dearne

Grafton

North Cranberry

e Back

Big Dam East e
*

Upper Silver
a

5.5
Lake pH

6.0 6.5

FIGURE 2. Total number of aquatic invertebrate taxa observed in relation to pH of 20 lakes sampled in June 2009 and 2010 in
Kejimkujik National Park and National Historic Site of Canada and vicinity in Nova Scotia. Note the significant positive
trend between lake pH and the number of invertebrate taxa (P = 0.005, r. = 0.36). -

5

MACROINVERTEBRATES AND ZOOPLANKTON IN KEJIMKUJIK NATIONAL PARK

NUSSBAUMER ET AL.:

2014

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Vol. 128

6 THE CANADIAN FIELD-NATURALIS1
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Lakes arranged by pH

FIGURE 3. Percentage composition (panel A) and total abundance (panel B) of various taxonomic groups sampled in June
2009 and 2010 in 20 lakes in Kejimkujik National Park and National Historic Site of Canada and vicinity in Nova
Scotia. Total abundance is the total number of individuals collected per lake. Lakes are arranged from the most
acidic (Big Red Lake) (pH 4.3) to the least acidic (McGinty Lake) (pH 6.6).

Lakes with high pH and calcium concentrations had
a larger number of bivalves, gastropods, and leeches
(Hirudinea) (Figure 5). Bivalves were observed only
in lakes with pH greater than approximately 4.9, and
abundance was significantly correlated with lake acidity
(Figure 5A) (r, = 0.775, P < 0.001). Gastropod abun-
dance was also significantly correlated with pH (Fig-
ure SB) (r, = 0.539, P = 0.014). Similarly, Hirudinea
abundance was significantly correlated with lake pH

(Figure 5C) (r, = 0.789, P < 0.001). No leeches were
collected from lakes with pH <5.5, with the exception
of a few individuals from the Erpobdellidae family cap-
tured in Peskawa Lake (pH 4.8) and Peskowesk Lake
(pH 5.0). In contrast, abundance of coleopterans was
significantly correlated with dissolved organic car-
bon (Figure 6) (r, = 0.650; P = 0.002), but not with
pH or calcium (P > 0.05).

2014 NUSSBAUMER ET AL.: MACROINVERTEBRATES AND ZOOPLANKTON IN KEJIMKUJIK NATIONAL PARK ,

150 ,
A) Caecidotea communis

120

Abundance
WH
ro)

D
io)

BUR,

8

B) Hyalella azteca

Abundance
fon)
=|

x

chat tkas

ee Vio Vr at are ee 14
C) () Caecidotea communis
80 % Hyalella azteca

60 %

40%

20 %

Percentage composition

0%

‘ isopod Caecidotea communis (panel A), the amphipod Hyalella azteca (panel B), and the corre-
pare pepe apenas these two species (panel C) observed in 20 lakes sampled in June 2009 and 2010 in
Kejimkujik National Park and National Historic Site of Canada and vicinity in Nova Scotia. Lakes are arranged by
level of calcium from the lowest (Ben Lake) (0.18 mg/L) to the highest (McGinty Lake) (1.12 mg/L). For panels A
and B, the horizontal line indicates the median, ® indicates mean, box indicates 25th and 75th percentiles, whiskers
‘ndicate minimum and maximum data points within 1.5 « the box height from the bottom or top (respectively), and

asterisks mark outliers.

8 THE CANADIAN FIELD-NATURALIST Vol. 128

A) Bivalvia

80

B) Gastropoda

Abundance

C) Hirudinea

15

10

x SYS AO & & & nO Oo &

ER vs + So oS eS a & ae x

FSC Fe of SWS ET PF Pk XS SoS
ef eee we SoS > NS

oe ee FO

Pow Cee le

Lakes arranged by increasing pH

FiGure 5. Abundance of Bivalvia (panel A), Gastropoda (panel B) and Hirudinea (leeches) (panel C) in 20 lakes sampled in
June 2009 and 2010 in Kejimkujik National Park and National Historic Site of Canada and vicinity in Nova Scotia
Lakes are arranged from the most acidic (Big Red Lake) (pH 4.3) to the least acidic (McGinty Lake) (pH 6.6) Hori-
zontal line indicates the median, ® indicates mean, box indicates 25th and 75th percentiles, whiskers indicate mini-

mum and maximum data points within 1.5 x the box height from the bottom or top (respectively), and asterisks
mark outliers. ‘

2014 Nt

150

100

50

Abundance of Coleoptera

SBAUMER ET AL.: MACROINVERTEBRATES AND ZOOPLANKTON IN KEJIMKUJIK NATIONAL PARK 9

Lakes arranged by increasing dissolved organic carbon concentration

FIGURE 6. Abundance of Coleoptera in 20 lakes sampled in June 2009 and 2010 in Kejimkujik National Park and National
Historic Site of Canada and vicinity in Nova Scotia. Lakes are arranged by concentration of dissolved organic car-
bon from the lowest (Beaverskin Lake) (2.8mg/L) to the highest (Big Red Lake) (19.5mg/L). Horizontal line indi-
cates the median, ® indicates mean, box indicates 25th and 75th percentiles, whiskers indicate minimum and maxi-
mum data points within 1.5 x the box height from the bottom or top (respectively), and asterisks mark outliers.

Discussion

We found that both pH and calcium were signifi-
cantly correlated with the number of aquatic macroin-
vertebrate taxa observed in the study lakes. Lakes that
were less acidic and lakes with higher calcium con-
centrations tended to have greater species richness.
These findings are consistent with other studies, which
also reported fewer aquatic invertebrate taxa in more
acidic lakes (McNicol et al. 1995a; Doka et al. 1997).
However, the relationship between chemical conditions
and the abundance of macroinvertebrates was less clear.

Fish were present in all of the study lakes (Kerekes
1975; Drysdale et al. 2005), and the presence of fish
likely influenced the macroinvertebrate and zooplank-
ton species richness. The most frequently collected taxa
were isopods, amphipods, and trichopterans. Gastro-
pods, bivalves, and ephemeropterans, commonly con-
sidered to be more sensitive to acidity, were collected
less frequently during the study. Lakes with lower pH
had fewer taxa (consisting mostly of isopods, coleopter-
ans, and oligochaetes), while lakes with higher pH had

greater taxa richness.

Isopoda and amphipoda }
Only one species of isopod was collected (Caeci-

dotea communis), but this species was present in all 20

study lakes. Caecidotea communis was also the most
abundant taxon in many of the study lakes, comprising
> 30% of the macroinvertebrates collected in 11 of the
lakes. The abundance of this species was negatively
correlated with calcium concentrations, and the high-
est numbers were found in the most acidic lakes (e.g.,
Peskawa Lake, Ben Lake, Peskowesk Lake). Schell and
Kerekes (1989) also reported Isopoda in Nova Scotia
lakes with pH as low as 4.4.

Isopods are known to be acid tolerant (Merritt eg al.
2008), but their high frequency of occurrence in lakes
in this study contrasts with their relative rarity in lakes
monitored in Ontario (RCW e7¢ a/., unpublished data).
Potential explanations include differences in the species
found in the two datasets (Ontario isopods were iden-
tified only to order), regional differences in species
habitat affinities, or a relative dominance of substrate
type or other habitat conditions that encourage isopod
abundance in lakes in this study area. Because sam-
pling methods were similar in the two regions, we do
not believe sampling variation is likely to be respon-
sible for these differences.

Amphipods were also common, and their abundance
was greater in lakes with high pH and high calcium
concentrations. Two species of amphipods were col-

10 THE CANADIAN FIELD-NATURALIST

lected: Crangonyx richmondensis (collected in 55% of
study lakes) and Hyalella azteca (collected in 95% of
study lakes). In this study, 17. azteca was present across
a broad pH range (i.e., 4.3 to 6.6). Studies in Ontario
have identified this species as acid sensitive (McNicol
et al. 1995a), with a minimum pH threshold of 5.6 or
higher (Stephenson ef al. 1986; Rosenberg ef al. 1997;
Snucins 2003). In this study, however, H. azteca ap-
peared to be very acid tolerant and was observed in
lakes with pH as low as 4.3.

Peterson (1987) also observed Hyalella in lakes with
low pH (4.5—S.5) in Nova Scotia and New Brunswick,
and reported that Hyalella species in lakes in the Mar-
itimes appear to be more tolerant of acidic conditions
than other Amphipoda. However, the lakes in that par-
ticular study had higher concentrations of calcium than
the study lakes with low pH in southwestern Nova Sco-
tia or in acidified lakes in Ontario (Peterson 1987). The
lakes in this study with low pH also had low calcium
concentrations.

It may be possible that a localized population of H.
azteca has adapted to the acidic environment in the
lakes in Kejimkujik National Park and National His-
toric Site. A genetic study by Witt and Hebert (2000)
examined populations of H. azteca from various loca-
tions across North America and found a complex of at
least seven species rather than a single species as pre-
viously believed. Grapentine and Rosenberg (1992)
also suggested that populations of H. azteca may have
adapted to acidic conditions in some regions of Canada.

Interpretation of regional variation in 1. azteca habi-
tat associations and identification of their potential role
in biological monitoring of lakes in this study area
would benefit from an improved understanding of the
geographic variation in their genetic profile and the
consequences for their tolerance of acidic conditions.

When we compared the relative proportions of iso-
pods and amphipods across the 20 study lakes, we
found that isopods were dominant in lakes with low
pH and low calcium concentrations while amphipods
were dominant in lakes with high pH and high calci-
um concentrations. Both amphipods and isopods are
photosensitive and avoid bright light by moving into
crevices or under rocks, leaves, and roots (Covich and
Thorp 2001, page 791), where they are less exposed
(complex substrates provide protection from preda-
tion by fish and crayfish) (Covich and Thorp 2001,
page 791). The substrate in many of the study lakes
consists of cobbles and boulders, which may partially
explain the high abundance of these two taxa.

Bivalvia and gastropoda

Invertebrate taxa with hard, calcareous shells such as
bivalves and gastropods were generally collected only
from less acidic lakes. A total of 10 species of bivalves
and 12 species of gastropods were collected. Bivalve
abundance was correlated with lake pH: bivalves were
observed only in lakes where pH was greater than 4.9.

Vol. 128

Because many lakes in the study area are acidic and
have low calcium concentrations, low abundance of
calcium-dependent macroinvertebrate taxa was expect-
ed. Our results are consistent with a previous study of
8 acid-sensitive Nova Scotia lakes by Schell and Ker-
ekes (1989), which found that bivalves did not occur
below a pH of 5.0.

This exclusion of calcareous species in acidic lakes
has also been noted for other acid-sensitive regions of
eastern Canada (Weeber er al. 2004; Jeziorski et al.
2008). As calcium concentrations in many acidified
lakes continue to decline (Jeziorski et al. 2008), this
may further reduce the abundance and distribution of
calcium-rich taxa such as bivalves and gastropods in
lakes in the study area.

Gastropods were also generally more abundant in
lakes with high pH and high calcium concentrations;
this finding is consistent with results from Ontario
(Bendell and McNicol 1993). One exception to this is
Ferrissia fragilis, which was the only species collect-
ed in lakes in the study area with pH lower than 6.
Bendell and McNicol (1993) also reported Ferrissia as
an acid-tolerant gastropod in study lakes in Ontario,
where it was the only gastropod taxon observed in
lakes with pH below 6. That study also suggested that,
above the minimal pH thresholds, gastropod abun-
dance in small oligotrophic lakes was not limited by
acidity or calcium concentrations but rather by food re-
sources. Predation, substrate type, and macrophyte bio-
mass can also play a large role in gastropod distribu-
tions (Brown 2001, page 310). In our study lakes, the
abundance of Ferrissia fragilis also did not appear to
be associated with pH, calcium, or dissolved organic
carbon and thus is likely limited by some other con-
straint such as predation or availability of food re-
sources.

Hirudinea

A total of 12 species of leeches were collected from
the study lakes, with only 4 of those species being com-
mon (i.e., occurring in >10% of the lakes). Counts were
generally low, and abundance was correlated with lake
pH. Hirudinea were not observed in lakes with pH
<5.5, with the exception of two Mooreobhdella fervida
collected in Peskawa Lake (pH 4.8) and one Erpob-
della punctata collected in Peskowesk Lake (pH 5.0).

Bendell and MeNicol (1991) observed similar reduc-
tions in the diversity and abundance of Hirudinea in
acidic conditions below pH 5.5. However, they sug-
gested that acidity alone does not predict the distribu-
tion of leech species and that predation and availability
of suitable prey also influenced their distribution (Ben-
dell and McNicol 1991). In addition, other studies have
shown that, although leeches are sensitive to low pH,
their occurrence and abundance are also influenced by
other factors, such as lake productivity (Schalk et al.
2001). Lakes in Kejimkujik National Park and Nation-
al Historic Site are oligotrophic and generally have low

2014

productivity (especially at the lower pH range), and
lower abundance of preferred prey may therefore play
an important role in the distribution of leeches there.
Coleoptera

Although lower in abundance than other groups,
coleopterans appeared to be tolerant of acidity and were
collected in all 20 study lakes. A total of 14 taxa were
observed (8 were common and 6 were uncommon).
The abundance of this taxonomic group was correlated
with dissolved organic carbon. A study of Ontario lakes
by Lento er al. (2008) also suggested a strong correla-
tion between macroinvertebrate abundances and dis-
solved organic carbon, especially in acidic lakes. Wood
et al. (2011) reported that dissolved organic carbon can
protect against the deleterious effects of low pH on
organismal function via physiological mechanisms.
Dissolved organic carbon can alter the permeability of
cell membranes in acidic conditions and also influence
transport physiology (Wood et al. 2011).

Other studies have suggested that water chemistry is
not as important a stressor on coleopterans as predation
by fish (Bendell and MeNicol 1987; Arnott et a/. 2006).
The darkly coloured water of some lakes in the study
area (due to high concentrations of dissolved organic
carbon) may provide coleopteran taxa with some pro-
tection from predation by fish and other visual preda-
tors.

Trichoptera

Trichopterans were common and taxonomically di-
verse in the study lakes, with 23 of the 30 taxa occur-
ring in >10% of the lakes. Trichopteran species col-
lected included taxa from 10 families, with the most
common and abundant families being Hydroptilidae,
Leptoceridae, and Limnephilidae. The trichopterans
collected in the study lakes generally had a high appar-
ent tolerance to acidity, with many of the observed
species occurring across a wide gradient in lake pH.

Trichoptera abundance can be strongly influenced
by fish predation, and trichopterans generally associated
with fishless conditions, such as the leptocerid 7riaen-
odes and phryganeid Banksiola (Bendell and McNicol
1995), were rare in the study lakes. Both of these organ-
isms are quite large and thus are likely to be attractive
prey for insectivorous fish. In contrast, the leptocerid
Nectopsyche was quite abundant. They are smaller in
size and construct cases with bristling twigs or elongate
sticks attached that may make them more difficult for
fish to consume as prey (Wiggins 2004).

Ephemeroptera and odonata .
Ephemeroptera generally had low abundance in the
20 study lakes, with a total of 10 taxa collected. This is
likely due to the acidity of the lakes, as ephemeropter-
ans are recognized as being sensitive to acidity (Car-
bone et al. 1998). Seven of the ephemeropteran taxa
were common, and 3 were uncommon. The most fre-
quently collected species were Caenis diminuta and the
genus Eurylophella, which have been reported to have

NUSSBAUMER E£T AL.: MACROINVERTEBRATES AND ZOOPLANKTON IN KEJIMKUJIK NATIONAL PARK 1]

at least some tolerance to acidity (Carbone ef al. 1998).
No ephemeropterans were collected from Ben Lake,
which is low in pH (4.8) and had the lowest calcium
levels of the 20 study lakes (0.18mg/L).

Odonates were taxonomically diverse in the study
lakes, with a total of 30 species observed (22 species
were common and 8 were uncommon). However,
counts were generally low, and odonates did not make
up a large proportion of macroinvertebrates in terms
of abundance. The most abundant family of damselflies
(suborder Zygoptera) was Coenagrionidae, which was
observed across a wide gradient of acidity. Larvae in
this family are relatively small (Hilsenhoff 2001, page
671) and thus may be less visible to predators such as
fish or larger predatory odonates.

Within the suborder Anisoptera (dragonflies), the
most common families observed in the study lakes
were Cordultidae, Gomphidae, and Libellulidae, while
Aeshnidae were rare. Anisoperta taxa also occurred
across a wide gradient of acidity; for example, Cordu-
lia shurtleffi was observed in 65% of the study lakes
(pH 4.3—6.6). Bendell and McNicol (1995) also found
that abundance of this particular taxon was not related
to lake acidity in Ontario lakes.

Diptera

With the exception of chironomids, Diptera were not
abundant in the study lakes. Ceratopogonidae were
present in all 20 study lakes, and no correlation with
acidity was detected. Chironomidae were frequently
collected in all of the study lakes, but were not target-
ed in our sampling and sorting, so specimens were not
identified to species level.

Hemiptera

Very few water striders were captured in the study
lakes. The only species with high abundance was Rheu-
matobates rileyi, in particular in Upper Silver Lake.
Although the abundance of this particular species has
been shown to have a strong correlation with pH
(Bendell 1988), acidity did not appear to be the main
driver in the presence of this particular species in the
study lakes.

Zooplankton

Of the 26 zooplankton species observed in the 15
study lakes, many were common and occurred across
a wide gradient of acidity. Daphniids were the only
taxonomic group that showed a clear correlation with
acidity in the study lakes: they were not observed below
a pH of 5.5. This finding is consistent with previous
studies, which have shown daphniids to be acid sensi-
tive (Yan ef al. 2008; Korosi and Smol 2012). In addi-
tion, daphniids are sensitive to calcium levels (Jeziors-
ki et al. 2008), and this may also explain their absence
in the lakes that had low pH and low calcium concen-
trations.

With the exception of daphniids, zooplankton abun-
dance in the study lakes did not appear to be correlated
with acidity alone. Dissolved organic carbon has been

shown to affect zooplankton populations, and the high
concentrations of dissolved organic carbon in some
of the study lakes may provide some protection from
visual predators (Yan ef al. 2008). Using paleolimno-
logical methods in 3 lakes in Kejimkuyik National Park
and National Historic Site, Korosi and Smol (2012)
found that there was a more pronounced change in-
duced by acidification in the assemblage of cladocer-
ans in clearwater lakes with lower concentrations of
dissolved organic carbon over time than in assemblages
in dark water lakes with more dissolved organic carbon.
Zooplankton can also be influenced by a large vari-
ety of natural factors, such as the availability of food,
competition with other zooplankton species, the pres-
ence of parasites, and the presence of both vertebrate
and invertebrate predators (Yan et a/. 2008).

Future directions and conclusions

These results provide a summary of the aquatic mac-
roinvertebrate and zooplankton assemblages in acid-
sensitive lakes in Kejimkuyik National Park and Nation-
al Historic Site and surrounding area in southwestern
Nova Scotia. Although some of the overall trends of
macroinvertebrate species richness with respect to vary-
ing pH were similar to results reported in other regions
of eastern Canada, several differences were noted.

Some of the lakes in the study area had physical
characteristics that differed from acid-sensitive lakes
in other regions of eastern Canada, and these physical
characteristics influenced the type and abundance of
benthic macroinvertebrates that were collected. pH can
vary spatially within each lake as well as seasonally
due to runoff, with pulses of acidity in the spring and
fall (Clair et al. 2007). These pulses also coincide with
lower temperatures, and at these times of the year
organisms may be less active and therefore more toler-
ant of their acidic environment (Stephenson and Mack-
ie 1994). Although benthic microhabitats near the lake
bed can have lower acidity than the upper water column
(Grapentine and Rosenberg 1992), lakes in the study
area are shallow with a large surface area which often
allows for mixing throughout the open-water period.
Therefore, benthic organisms would likely be exposed
to high acidity throughout the active growth period in
the summer.

All aquatic sampling methods have inherent biases
in their sampling efficiencies for different invertebrate
taxa. We employed multiple sampling methods in order
to collect a wide range of taxa, but there was likely to
have been variation in efficiency among the sampling
methods with respect to particular taxa. Because the
same suite of methods was used in all lakes, we assume
the effects of this variation were consistent across the
20 study lakes, and we emphasize comparisons of in-
vertebrate taxa patterns between lakes, rather than with-
in lakes.

Our sampling methods, which were initially devel-
oped to collect benthic invertebrates from thick organic
sediments in small Boreal Shield lakes in Ontario

THE CANADIAN FIELD-NATURALIST

Vol. 128

(MeNicol e¢ al. 1995b), may not be have been as suit-
able for lakes with rocky substrates. Although regional
variation in species’ habitat affinities may have con-
tributed to particular differences between the findings
from this study and reports from other regions (€.g.,
isopods, H. azteca distributions), substrate or other dif-
ferences in the habitat also may have been a factor.
Hoop sampling (visual searches in a confined area
along the shoreline) worked particularly well in our
lakes for sampling species of Trichoptera. Future stud-
ies should incorporate traditional benthic drag sampling
with other methods such as kick and sweep, rock pick-
ing, or artificial substrates.

Carbone et al. (1998) successfully sampled macro-
invertebrates in shallow, rocky littoral habitats using
substrate cages filled with native rocks to match the
rocky littoral substrate of sample lakes. This method
might work well in Kejimkuyik National Park and Na-
tional Historic Site, where the littoral zone of many
lakes is extremely shallow and consists of cobble and
boulders. Many species collected in the study were rare
(occurring in only one or two of the lakes) and had low
counts. Increasing sampling effort, especially in the
large lakes with varying substrate types, would reduce
the likelihood of missed taxa.

Another interesting difference between the lakes in
Kejimkujik National Park and National Historic Site
and the lakes in the Boreal Shield in Ontario is the high
concentration of dissolved organic carbon due to natu-
rally occurring bogs and wetlands in the watersheds.
The extremely dark waters of some lakes in the study
area may benefit particular invertebrate species through
physiology, protection from visual predators, or other
reasons.

The data presented here establish a baseline for
future monitoring in Kejimkujik National Park and
National Historic Site as acid deposition continues to
affect this region. Because the lakes are naturally acidic
and are extremely vulnerable to additional acid inputs,
recovery is slower than in other regions in eastern
Canada affected by acid deposition (Clair et a/. 2011).
Additional effort may be required to reduce the impacts
of acidification on the aquatic organisms that live in
these ecosystems.

Acknowledgements

This study was funded by the Clean Air Regulatory
Agenda (CARA) program of Environment Canada.
Collections were made under scientific permits from
Parks Canada and Fisheries and Oceans Canada. All
animal handling followed procedures approved by the
Environment Canada Animal Care Committee. We
would like to thank Bohdan Bilyj (BIOTAX), who
completed taxonomic identifications of macroinver-
tebrates, and the Sudbury Freshwater C 0-op Unit at
Laurentian University, where zooplankton identifica-
tions were completed. In addition, we thank Tom Clair
and colleagues in the Water Science and Technology

2014

Directorate, Environment Canada for providing water
chemistry data. We thank Adam Martens and Miriam
Morgan for field and lab assistance. Logistical support
was provided by staff at Kejimkujik National Park and
National Historic Site, Parks Canada.

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Received 30 August 2012
Accepted 20 February 2013

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An Illustrated Key to the Mandibles of Small Mammals of Eastern
Canada

peeled i ; 7 5 .

DOMINIQUE FAUTEUX!4, GILLES LUPIEN2, FRANCOIS FABIANEK?, JONATHAN GAGNON!, MARION SEGuy!,
Louis IMBEAU!
Vy Bea ey 3 . . P ~ .
Chaire industrielle G RSNG-UQAT-UQAM* en amenagement forestier durable and Centre d’étude de la forét, Université du
wn, 4 Quebec en Abitibi-Témiscamingue, Rouyn-Noranda, Québec JOX SE4 Canada
“Ministere du Développement durable, de l'Environnement, de la Faune et des Parcs, Direction régionale du Saguenay-Lac

Saint-Jean, Jonquiére, Québec G7X 8L6 Canada
3 37 ~ an P a . , . A P, ° ae - -
ps entre d etude de la forét, Faculté de foresterie, géographie et geomatique, Université Laval, Québec, Québec G1 V 0A6 Canada
Corresponding author: dominique.fauteux. | @ulaval.ca

Fauteux, Dominique, Gilles Lupien, Francois Fabianek, Jonathan Gagnon, Marion Séguy, and Louis Imbeau. 2014. An illustrated
key to the mandibles of small mammals of eastern Canada. Canadian Field-Naturalist 128(1): 25-37.

Skulls are often used to identify small mammals, and most identification keys to small mammals have been developed on the
assumption that whole skulls will be available. However, the skulls of small mammals are seldom found intact in predator
pellets or nests, and the bones of several individuals are often scattered and mixed, making counting impossible without the
use of a specific cranial part. In addition, only a few keys include all the species found in the eastern provinces of Canada.

Mandibles readily resist degradation by the gastric acids of both avian and mammalian predators and are often found intact
in food caches of mustelids and in bat hibernacula. We therefore designed an illustrated dichotomous key to small mammals
(mean mass <5 kg) of eastern Canada based on diagnostic mandible characters (including the teeth and one dentary bone).
We identified and confirmed diagnostic characters to distinguish 55 species from the orders Lagomorpha, Rodentia, Sorico-
morpha, Carnivora, and Chiroptera. These diagnostic characters are based on a review of the literature and were confirmed
by measurements performed on both museum and trapped specimens. In order to facilitate identification, photographic illus-
trations are provided for each couplet of the key.

The ability to identify small mammals using their mandibles will reduce the number of skull components needed and has proven
to be a useful tool in the study of the diet of predators. This key may also be helpful in identifying bats in the genera Myotis,
Perimyotis, and Eptesicus, which are presently affected by the spread of white-nose syndrome (caused by Pseudogymnoascus
destructans) throughout the eastern part of Canada.

Key Words: Lagomorpha; Rodentia; Soricomorpha; Carnivora; Chiroptera; shrews; moles; voles; lemmings; mice; bats; hares;
weasels; lower jaw; skull; dentary; eastern Canada

Introduction

Small mammals consumed by predators are partic-
ularly difficult to identify because their skulls are often
physically damaged or they have been degraded by
gastric acids (Mayhew 1977). Cranial bones that resist
degradation often disassociate from the larger com-
ponent they were affixed to and are often found scat-
tered in predator scats, pellets, or nests (Buidin et al.
2007; Khalafalla and Iudica 2010). They may also be
found as concentrations of loose bones near caves or
other shelters used by predators (Buden 1974). Preda-
tors such as mustelids have “food caches” in which
they store carcasses for later consumption (Oksanen ef
al. 1985). As a result, prey remains may be disassoci-
ated and may accumulate.

Several published keys to small mammal skulls are
based on the assumption that the whole skull is avail-
able (van Zyll de Jong 1983; Glass and Thies 1997;
Lupien 2001, 2002; Nagorsen 2002; Chapman ef al.
2007), but this is rarely the case with prey remains
(Mollhagen et al. 1972; Buden 1974; Balciauskiene ef

al. 2002). Furthermore, loose bones of different indi-
vidual prey items are often mixed. The minimum num-
ber of individuals is a derived unit of abundance often
used in paleozoology (Lyman 2008). By using a single
skull component, this method avoids overestimating
species abundance in bone aggregations. The mandible
has been proposed as a useful cranial component for
identifying groups of mammals (Roest 1991; Balci-
auskiene ef a/. 2002), but it has rarely been used to
identify mammals to the species level, except for
shrews (Repenning 1967; Carraway 1995).

The mandible, or lower jaw, is composed of teeth
and a pair of dentary bones (Figure 1). The teeth of the
mandible are often referred to as the lower dentition,
and each tooth is identified with a lower case letter
(1.e., p3 for the third premolar). For the present arti-
cle, we focused on the mandible and thus omitted the
term “lower”. Because the left and right dentary often
separate as a result of degradation, it is imperative that
the same dentary bone (1.e., left or right, but not both)
be used for counting purposes.

26 THE CANADIAN FIELD-NATURALIST

Several diagnostic characters make the mandible an
ideal tool for identifying most mammalian species that
have very few but sturdy bones. The size, the dental
formulae, and the occlusal patterns of the molar enamel
are key characteristics that are often used in keys to
skulls (Repenning 1967; Glass and Thies 1997; Lupien
2002; Nagorsen 2002). Furthermore, diagnostic char-
acters of the dentary bones are found on both the ante-
rior and the posterior parts. The size and shape of the
lower edge of the ramus and the position of the mental
and dental foramina, as well as the size and shape of the
condylar, coronoid, and angular processes, are useful
characters requiring only a few metric measurements
(Roest 1991; Carraway 1995).

We present an identification key to the mandibles of
all established small mammals (mean mass of <5 kg)
of eastern Canada to assist in the identification of prey
remains and other types of loose bones when skulls are
incomplete or damaged. Each criterion mentioned in
the couplets of the key is illustrated by a picture as a
visual support. A glossary and the general nomencla-
ture are also provided.

Methods

According to Merritt (2010), mammals may be cat-
egorized as small when the average mass of the species
is less than 5 kg. Based on this criterion, we selected
all the small mammals established in the provinces of
Ontario, Quebec, Newfoundland and Labrador, Prince
Edward Island, New Brunswick, and Nova Scotia
(Peterson 1966; Banfield 1974; Dobbyn 1994; Des-
rosiers et al. 2002; Naughton 2012). The general tax-
onomy used in the key is listed in Table 1.

This key summarizes all diagnostic mandible char-
acters that we have found in the literature for the orders
Lagomorpha (Roest 1991), Rodentia (Klingener 1963;
Phillips and Oxberry 1972; Grayson ef al. 1990; Roest
1991; Lupien 2002; Chapman ef al. 2007), Sorico-
morpha (Hallet 1978; Yates and Schmidly 1978; van
Zyll de Jong 1983; Carraway 1995; Glass and Thies
1997; Lupien 2001), Carnivora (Roest 1991; Glass and
Thies 1997), and Chiroptera (Gaudin et al. 2011). Cer-

Vol. 128

tain species were very difficult to distinguish using the
morphologic features of the mandible alone. Therefore,
we included morphometric measurements such as the
length of the mandible, the length of the mandibular
tooth row, and the height of the coronoid process when
two species or groups of species could be distinguished
only by size.

We validated the mandible characteristics presented
in this key by studying specimens from Ontario, Que-
bec, Newfoundland and Labrador, Prince Edward Is-
land, New Brunswick, and Nova Scotia preserved in
the Canadian Museum of Nature and Université Laval.
Morphometric measurements were validated on 10
specimens of each species when possible. Otherwise,
all specimens available were used. We further extract-
ed a sample of reference mandibles from complete
frozen specimens, in collaboration with the Ministere
du Développement durable, de l’Environnement, de la
Faune et des Parcs du Québec and the Université du
Québec a Rimouski, and from specimens trapped dur-
ing a related study (Fauteux et al. 2012). The relevance
of the diagnostic characters in identifying prey remains
was validated by Séguy (2010) using nest remains to
quantify the diet of Northern Saw-whet Owls (Aegolius
acadicus).

Results and Discussion

We found that 55 of the 60 small mammal species
of eastern Canada could be identified from their man-
dibles. The White-footed Mouse (Peromyscus leuco-
pus) and the Deer Mouse (P. maniculatus) could not be
identified to the species level, because their mandibles
are identical. Although both Peromyscus species may
be differentiated using several skull measurements,
biochemical and genetic markers are probably the only
reliable methods to date (Aquadro and Patton 1980;
Rich et al. 1996). Similarly, three species of lagomorphs
(i.e., Lepus arcticus, L. europaeus, and L. townsendii)
could not be distinguished using the mandibles alone.

Consulting species’ distribution may facilitate iden-
tification of small mammals (Banfield 1974: Desrosiers
et al. 2002; Naughton 2012). For example, Sciurus

FIGURE I. (A. Labial view; B. Occlusal view) The mandibles of carnivores (Martes americana) (A) and rodents (Ondatra
zibethicus) (B). Labels refer to the incisor (i), canine (c), premolar (p), molar (m), mandibular tooth row (mt), coronoid
process (cor), condyle (con), angular process (ang), vertical ramus (vra), and horizontal ramus (Ara).

2014 Faureaux ET AL.: ILLUSTRATED KEY TO THE MANDIBLES OF SMALL MAMMALS OF EASTERN CANADA 27

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2014 FAUTEAUX ET AL.: ILLUSTRATED KEY TO THE M

niger are found only in extreme southern Ontario, and
the distribution of Sorex maritimensis is restricted to
New Brunswick and Nova Scotia.

The mandible is highly polymorphic between and
within orders. The order Soricomorpha can be distin-
guished from other orders because the canine is simi-
lar in size to the premolars and the angular process is
long and slender (Figure 2B) (key section D). In Lago-
morpha, the large angular process and the very small
coronoid process are probably the most distinctive char-
acters (Figure 3A) (key section B). In contrast, species
of the order Rodentia have a well-developed coronoid
process, often with complex occlusal patterns on the
molars (Figures 3B) (key section C). Carnivores have
large canines and a coronoid process that is dispropor-
tionately larger than the condyle and the angular pro-
cess (Figure 4B) (key section E). Species from the order

=

Be >.

ANDIBLES OF SMALL MAMMALS OF EASTERN CANADA 29

Chiroptera are mainly characterized by the relatively
small vertical ramus and the conspicuous bump on the
lower edge of the horizontal ramus beneath the canine
(Figure 5B) (key section F),

In some cases, mandibles may be broken and/or
teeth may be missing. To address this problem, we pro-
vide two or more criteria. However, we struggled to
find more than one mandibular characteristic in cer-
tain groups of species. In the orders Lagomorpha and
Carnivora, only the length of the mandibular tooth row
and the height of the coronoid process may be used
effectively to distinguish the hares (Lepus spp.) and the
weasels (Mustela spp.). Voles and lemmings may be
more effectively differentiated with dental criteria, and
identifications may become difficult when the teeth
are missing (Banfield 1974; Lupien 2002). Although
identifications using heavily degraded mandibles (e.g.,

% b

;
4
x

FiGuURE 2. (/abial view) Dentary bone of rodents with a large diastema (dia) (Glaucomys volans) (A), and soricomorphs
(Blarina brevicauda) (B).

FiGurE 3. (labial view) Coronoid process (cor) and condyle (con) of lagomorphs (Lepus arcticus) (A) and rodents (Marmota

monax) (B).

ang

F g 4. (labial view) Size of the angular process as well as the size of the canine compared to the adjacent premolar in sori-
IGURE 4. (Ic gul J amet ss
comorphs (Parascalops breweri) (A) and carnivores (Neovison vison) (B).

30 THE CANADIAN FIELD-NATURALIST

FiGurE 5. (labial view) Dentary bones of carnivores (Mustela erminea) (A) and chiropterans (Perimyotis subflavus) (B )
with the height of the condyle (/,,,), height of the coronoid process (/,,,,.), and the conspicuous mandibular bump of
chiropterans.

complete absence of teeth on specimens of Cricetidae)
may be generalized, the resistance of mandibles to
degradation and the number of criteria we included in
the key should prove useful in identifying lightly de-
graded mandibles to the species level.

Sex and age are important factors that may mean
that certain mandible criteria may not be useful (be-
cause of sexual dimorphism and growth). We acknowl-
edge that this may be a limitation to a key based on
osteometry. Identifications conducted on bones of juve-
niles that are mixed with bones of adult prey may have
a lower resolution (i.e., identifications stop at the genus
level) than when only adults are present. As a solution,
we included in the vast majority of couplets one or
more known morphologic characters that are persistent
through age and that do not differ between males and
females, such as the morphology of the ramus. Using
the mandible is also a useful tool for the counting of
individual remains and do not necessitate lengthy and
costly methods that often require advanced laboratory
skills (e.g., identifications using DNA).

This is a new tool for identifying and monitoring
all of the small mammals of eastern Canada. To our
knowledge, this is the first comprehensive key designed
in North America that uses the mandible exclusively.
Use of the mandible enables degraded specimens of
most small mammals to be identified down to the spe-
cies level and it facilitates counting activities. More-
over, bats of the genera Myotis, Perimyotis, and Eptesi-
cus have declined dramatically in the past few years as
a result of the spread of white-nose syndrome (caused
by Pseudogymnoascus destructans) in the eastern part
of the United States and Canada (Blehert et al. 2009).
Identifying mandibles on the floor of caves and in
other hibernacula might be useful for monitoring car-
casses.

Acknowledgements

Financial support was provided by the Natural Sci-
ences and Engineering Research Council of Canada
(NSERC) and the Fonds de recherche du Québec —
Nature et technologies (FRQNT) of Quebec. We thank
the Centre for Forest Research and the Conseil de re-

cherches en sciences naturelles et en génie du Canada
— Université du Québec en Abitibi-Témiscamingue—
Université du Québec a Montréal (CRSNG-UQAT-
UQAM) Industrial Chair in Sustainable Forest Mana-
gement for technical support. We thank the Canadian
Museum of Nature and Michel Gosselin and Kamal
Khidas (both of the Canadian Museum of Nature) for
their help with the skull collection and for providing
us with a digital camera attached to a binocular. We
thank David W. Nagorsen for providing constructive
comments on earlier versions of the manuscript. We
thank Amélie Drolet for linguistic revision of the text.
We also thank all field assistants who participated in
the collection of bones and the small mammal trapping
campaigns.

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Supplementary material available at:
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Received 9 January 2013
Accepted 8 April 2013

Glossary of terms

Alveolus
Angular process

Anteroconid
Anteromedian fold
Anteroposterior length
Brachydont tooth
Condyle/condylar process

Coronoid process
Closed triangle (of enamel)
Mandibular foramen

Dentary bone

Diastema (plural: diastemata)
Enamel

Horizontal ramus

Hypoconid

Hypsodont tooth

Interdenticular space
Labial
Labiolingual width

Length of the mandibular tooth row

Lingual
Mandible
Mandibular tooth row

Mental foramen
Metaconid

Occlusal

Paraconid

Pigmentation
Postmandibular foramen
Premetaconid fold
Protoconid

Re-entrant angles

Temporal fossa
Vertical ramus

THE CANADIAN FIELD-NATURALIST Vol. 128

Socket in which the roots of a tooth are set (Figures 24, 25, and 28) (aly).
Posterior and ventral-most bony projection of the mandible; the angular
process is posterior to the coronoid process (Figures | and 24) (ang).
Anterior-most cusp on the m1 of jumping mice (Figure 19) (antc).
Concave fold created by the anteroconid on the anterior part of m1 (ant/).
Length in the direction of the mandibular tooth row.

Closed-rooted tooth with determinate growth (Figures 22 and 23).
Bony projection located on the ramus between the coronoid and the
angular process (Figures | and 24) (con).

Posterior and dorsal-most bony projection of the mandible; the coronoid
process is anterior to the angular process (Figures | and 24) (cor).

In rodents, the external layer of molars that forms occlusal triangular
shapes (Figures 16 and 21) (c/).

Small hole located below the temporal fossa and serves as a canal for
the dental nerve.

One side (half) of the mandible.

Space between two adjacent teeth (Figure 2) (dia).

The hard external layer of the tooth.

The anterior part of the dentary that supports the teeth (Figure 1B) (Ara).
The most posterior cusp (Figure 24).

Continually growing tooth. The enamel typically covers most of the
tooth. Teeth are rootless (Figures 22 and 23).

Space between the cusps present on the incisors of shrews (Figure 30).
Next to the lips.

Length of teeth in the direction perpendicular to the mandibular tooth
row.

Length of the lower tooth row (cl—m3) (Figures 1, 6, 33, 37, and 42).
Next to the tongue; the interior of the mouth.

Both dentary bones, often referred as the lower jaw (ma).

All contiguous teeth of one dentary bone (mz). In Carnivora, Chiroptera,
and Soricimorpha, all teeth form the toothrow. In Rodentia and Lago-
morpha, premolars and molars form the toothrow.

Small hole located on the labial face of the horizontal ramus (Figure 24).
Cusp posterior to the anteroconid on the lingual side of m1 in jumping
mice.

The side of the teeth which meets with the opposing teeth.
Anterior-most cusp on molars in lateral view (Figure 24).

Coloration of the teeth (pg). It is often dark in shrews.

Small hole next to the mandibular foramen that connects with the tem-
poral fossa (Figure 30).

Small depression, resembling a trench, separating the anteroconid from
the metaconid on the molars of jumping mice (Figure 19) (prm/).
Middle cusp on the molars of shrews in lateral view (Figures 19 and
24).

Inward pointing angle defined by the margin of the prismatic molars in
voles (Figures 16 and 21) (ra).

Large opening on the lingual side of the vertical ramus.

The posterior part of the dentary, composed of the coronoid, condylar,
and angular processes (Figure 1B) (vra).

2014 FAUTEAUX ET AL.: ILLUSTRATED KEY TO THE MANDIBLES OF SMALL MAMMALS OF EASTERN CANADA 33

Key to the mandibles of small mammals of eastern Canada

(full key illustrated with pictures provided in Supplementary material available at: http://www.canadianfield
naturalist.ca)

A. General key to small mammals
la. Wide diastema between the incisor and molars (Figure 2A)

5 RAPE TES IRIAN, sein gd Behl Update 2
lb. No diastema between the incisor and molars (Figure 2B) ........0.0 0000 ece eu eeveeeeues 3
Di. an recat = itemcue Rae : ' Hee :
2a. Two premolars and three molars; coronoid process and condylar process not differentiated or
coronoid process munute (Figure SA) ccc odes. ancacmecmsme ances aman Lagomorpha (section B) 5
2b. One premolar or none and three molars; coronoid process clearly differentiated from the condylar
DIO ee RE TINS VE hoe oo sh id elastin wan ahs a Gm mie ame etes «EO ae's sf oN Rodentia (section C) 7
3a. Canines and premolars similar in size; well-developed angular process that is often the most
posterior part of the dentary bone (Figure 4A) ................00005. Soricomorpha (section D) 31
3b. Canines two to three times the size of the adjacent premolar; small but robust angular process
nhs Bs eae RCT aRE Pee eT AUN, MM eae Te, Caceres AT wal See 1 OL wt RES, (Figure 4B) 4

4a. The most posterior molar often much smaller than the most anterior molar; lower edge of ramus
without a bump under the canine; height of the coronoid process much higher than the height of the
COMIC PROCESS (RAOICS SAN) 2 oe a oe eth w tw oe RS Uk ewe a Re a Oa Carnivora (section E) 42
4b. Three W-shaped molars of similar size; lower edge of ramus with a bump under the canine; height
of the coronoid process similar in size to or slightly higher than the height of the condylar process
Ai Re CSN tae rete, Ue Merete hn cates ee aS ahh sm eat caairte tne wi whe MIS Nee x OS Chiroptera (section F) 50

B. Lagomorpha (Leporidae)
5a. Height of coronoid process >40 mm; length of mandibular tooth row >16 mm (Figure 6A)

GP SR a eae 9h denis ih Oe Aw hans Hash Ra vere deamR dor igitime dock Lepus arcticus, L. townsendii, L. europaeus
5b. Height of coronoid process <40 mm; length of mandibular tooth row <16 mm (Figure 6B) .... 6

6a. Mental foramen easily visible from the occlusal view (Figure 7A)
re ere ce ee eee eta eee ELSE ee eee ee Sylvilagus floridanus

ace phn le BETA Cd RUA gx Le COSI NS JEL AE POOR AES HOE LT 4 SO Lepus americanus

C. Rodentia (Cricetidae, Dipodidae, Erethizontidae, Muridae, and Sciuridae)
7a. Lower edge of horizontal ramus with sharp angle under p1 (Figure 8A); angular process clearly
smaller than the coronoid process; cheek teeth with closed circular patterns of enamel (Figure 8B)
I i a aL i et at Rl: 8 ea He ib Rida gi dsr i ra Erethizon dorsatum
7b. Lower edge of horizontal ramus smooth; the coronoid process and the angular process are similar
in size or the angular process is larger than the coronoid process; cheek teeth with triangular patterns
of enamel or without clearly defined patterns .. 0.26. c cence eect eee eee e eee e es 8

8a. Angular process clearly the most exterior part of the mandible (Figure 9A); angular process about
twice as wide labially as the condylar process (Figure 9A); anterior edge of the coronoid process
that connects with the angular process creates a bump pointing outwards at the level of pl-m1 in the
occlusal view (Figure 9B); 2-2... cece cree eee e reer een e ncn n es eseneesen aces Poe Marmota monax
8b. The condylar process or the coronoid process is the most exterior part of the mandible (occlusal
view); no bumps created by the edge of the coronoid and angular processes next to pl-m1; angular

process about the same labial thickness or less than the condylar process ........-+-++eeseeeee 9
9a. Tip of the angular process clearly higher than the teeth igure LUA). as.s cone vee Ondatra zibethicus
9b. Tip of the angular process below or even with the teeth (Figure 10B) .......5....05.050ee. 10
10a. One premolar (Figure 11 A); angular process extends slightly behind the coronoid process (Figure
GC i aah ROS Re ee A oho Perce tet afore anny Nee Ae Rei <ebee baba 11
10b. No premolar (Figure 11B); angular process extends well behind the coronoid process (Figure

: 18

Oy Ge Sane Pe NT AOR PUD. LE TP ry Goat TL RNAI PRT PHRMA OS 1 0.

34 THE CANADIAN FIELD-NATURALIST Vol. 128

lla. Coronoid process long; size of the notch between the coronoid and condylar processes similar
in size to the notch between the condylar and angular processes (Figures 12A and 12B) .....---.
11b. Coronoid process relatively short; size of the notch between the coronoid and condylar processes
clearly smaller than the notch between the condylar and angular processes (Figures 12C and 12D) ..

12a. T-shaped condylar process; angular and condylar processes equally posterior (Figure 12A)
icy each ebeelad fg eo 00 44 gale Re’ dere ate aa S SE Oe eer aes eae) ae Ae Poliocitellus franklinii

12b. A-shaped condylar process; condylar process clearly the most posterior component of the ramus

CFigure. IAB ots keesioheits deg «vetavaaitig Ce Ghar aaah me arin oes Te ag ee ree 13
(3a, Léengthiofithe mandibular tooth now <5 Sime e aie eee es tie sas oie Tamias minimus
[3b; Lenethotithemandibular toothrow =S-5.00m ..4¢. se ond sos seeeenae as aoreeee Tamias striatus
14a. Height of the coronoid process >17 mm; length of the mandibular tooth row >35mm ....... 15
14b. Height of the coronoid process <17 mm; length of the mandibular tooth row <35mm ....... 16

15a. Coronoid process short; notch created by the coronoid process and the condylar process appears

wide open; lower tip of the angular process appears squared (Figure 13A) ................. Sciurus niger
15b. Coronoid process longer; coronoid notch narrow; lower tip of the angular process appears rounded
(igure ISB)" slsied ad aude yall dba eee Bee ae eee Sciurus carolinensis
16a. Uppermost edge of the condylar process relatively flat (Figure 14A) ........ Tamiasciurus hudsonicus
16b. Uppermost edge of the condylar process concave (Figure 14B) ................-.2.000-- 17
17a. Posterior tip of the angular process above the notch on the lower edge of the horizontal ramus
(Chee Sv) i en a ee Glaucomys volans
17b. Posterior tip of the angular process below or at the same level as the notch on the lower edge of the
hotizontalkrannasy @ig@une USB) ree: Samal 6 i ccscscna sve Fup io buenas ree sf Se ol dees Be Glaucomys sabrinus
18a. Molars without re-entrant angles or closed triangles (Figure 16A) ....................--. 19
18b. Molars with well-defined lingual and labial re-entrant angles (Figure 16B), often with closed triangles
Oenainel(WSMREIGC): oss. ade bv oe eRe SOE eee ee oe eee 23

19a. Condylar process clearly the most posterior part of the dentary bone; coronoid process small, at
about the same height as the condylar process (Figure 17A)

MeSH E fats) 3f 9 «: «6. eR oe Seige egy ee eee ee Peromyscus leucopus or P. maniculatus
19b. Condylar process slightly posterior to the angular process or about equally posterior; coronoid

process relatively long and higher than the condylar process (Figure 17B) .................... 20
20a. Molars with complex patterns of enamel loops (Figure 18A) ...............-cccceeeeeee 21
20b. Molars with simple patterns of enamel loops (Figures 18B and 18C) .................... 22

21a. Anteromedian fold present on m1; anteroconid of m1 clearly separated from the protoconid by
the preprotoconid and premetaconid folds (Figure 19A) ........... ccc cc ecceeccccece. Zapus hudsonius
21b. Anteromedian fold absent on m1; anteroconid of m1 not separated or sli ghtly separated from the
protoconid by the premetaconidifold (igure 19B) cc... ee. . couscc.ss see. cauean Napaeozapus insignis

22a. Molars with simple patterns of enamel (Figure 20A) ..........0. ccc eecccccccuu. Mus musculus
seh coh Sir Rattus norvegicus

23a. Re-entrant angles of molars much deeper on lingual side than on labial side (Figures 21A, 21B,
ANG2NC)s pata hveRds,«: 4 «x a's nn oy nied Ae Re ea Oe re pete when Soke ied ei 24

ree ee eee er ee Pk Peery ee re eI E E G 26
24a. Brachydont teeth (molars closed-rooted) (Figures 22A, 22B, 22C, 23A, and 23B); several small

closed triangles on the labial side of molars (Figure 21A) ........................ Phenacomys ungava
24b. Hypsodont teeth (molars open-rooted) (Figures 22D and 23C): one closed triangle or none on the
labial side of each molar (Figures 21B and 21C) 2.1.2... cece cee cece cee ccc eccececee.. 25

2014 FAUTEAUX 27 4L.: ILLUSTRATED KEY TO THE MANDIBLES OF SMALL MAMMALS OF EASTERN CANADA 35

ser A single closed triangle on the labial side of each molar (Figure 21B) ...........: Synaptomys cooperi
25b. No closed triangle on the labial side of molars (Fi WUNS- SG) |; s9We lh es ooe aa Synaptomys borealis

26a. Brachydont teeth (molars closed-rooted) (Figures 22A, 22B, 22C, 23A, and 23B); occlusal triangles

of molars rounded and “enclosed” by the enamel borders (Figures 21D and 21E) ............... yi)
26b. Hypsodont teeth (molars open-rooted) (Figures 22D and 23C); occlusal closed triangles with sharp
CpGeote eT 2G Sean IID: nc ows danauc cles PRRp os ee Oe eee oe 28
27a. Occlusal triangular shapes of enamel of m1 and m2 often connected by wide bridges; shape of the
anterior triangle of m3 is typically similar to the posterior triangles (Figure 21D) ......... Myodes gapperi
27b. Occlusal triangles on m1 and m2 often connected by narrow bridges; shape of the anterior triangle
of m3 often different from the other triangles (Figure 21E) .............cceeeeeeeeeee Myodes glareolus

28a. Presence of a small fold of enamel on the anterior and lingual side of m2 (Figure 21F)

Ra as ke EE ee teeter terete tenes e sees ss Dicrostonyx hudsonius
28b. Absence of a small fold of enamel on the anterior and lingual side of m2 (Figures 21G, 21H,

SERED A RIM Ree MRSA yen scl Hee aotearoa ti vate Blea RMR CLIN ra hohe SS OT RO 29
29a: Three Glosed trianpleson mh (Pigure 2G). wed. eee ies Sed. ee eee ds oat Microtus pinetorum
25>) Pive elosed tiangles on ml (Ficures 2Uhiand 201)" Phys Pe eG oe 30
SWee Two closed triangles on m2 (Figure 21H)» 222. Soot ee Fee Microtus chrotorrhinus

30b. Four closed triangles on m2 (Figure 211)

D. Soricomorpha (Soricidae and Talpidae)
31a. Teeth all white; incisors without a posterior cusp; alveolus of incisors does not extend under pre-

Hlolaes Gr tiniel aes Ci retin AN eS See a ce aids eA ate airs We a ae EA Op tae NO are alee 32
31b. Tip of teeth often with red and/or brown pigments; incisors with a posterior cusp; alveolus of
incisors extends beneath the first premolar or posteriorly (Figures 24 and 25B) ..............-. 34

32a. Two incisors, no canine, and three premolars; presence of a short diastema between the second

incisor and the first premolar (Figure! 264) Yen. eee Ee I 8 Scalopus aquaticus
32b. Three incisors, one canine, and four premolars; presence of several short diastemata between the
premolars (Figure 26B) or complete absence of diastemata (Figure 26C) .........-.--+++++55- 33

33a. Canine and the first three premolars separated by short diastemata; angular process long and
slender; condylar process about the same height as the coronoid process or higher; coronoid process

clearly smaller than the condylar process (Figure 26B) .......--- 00 esee eee eee ees Condylura cristata
33b. Canine and first three premolars not separated by diastemata; coronoid process higher than the
condylar process; coronoid larger than the condylar process ( PIgMneQ6E)i te: oie Mewes Parascalops breweri

34a. Alveolus of incisor extends slightly or substantially beneath m1; alveolus extends at the level of
the m1 paraconid or posteriorly (Figure 27A) ........ 0. eee e eect eee eee ene een eee eens 35
34b. Alveolus of incisor does not extend beneath the m] paraconid (Figure 27B). 2.5.6 400. es. 36

35b. Three small cusps on the occlusal surface of the incisor; angular process long and very slender

CRicutie Zou tre Oe EE es PSE a oth es LN MURINE EOS oe Sorex hoyi
35a. One or two small cusps on the occlusal surface of the incisor; angular process relatively short and
sObUst (RigmRO SEN) ©. NNT AUIS VaR Ree PAN Ta PN ee leek Ae As 94 Blarina brevicauda
36a. Mental foramen located beneath the m1 paraconid; the space between both cusps on the molars
is relatively large (Figure 29A) ......--+ +++ sere reer teres peer tenet tenes eres Sorex dispar
36b. Mental foramen located beneath the m1 protoconid or posteriorely; the space between both cusps
on the molars is relatively small (Figure 29B) ....cicisevevseoveverer tes eessub ener newsece 37

37a. Postmandibular foramen present (Figure 30A); deep interdenticular spaces on il (Figure 30B) ... 38
37b. Postmandibular foramen absent; shallow interdenticular spaces on il (Figure 30C) ............ 39

36 THE CANADIAN FIELD-NATURALIST Vol. 128

Sorex maritimensis

38a. Height of coronoid process <4.5 mm (Figure 24) 00... 6. sce ise see wens
Sorex arcticus

38b: Height of coronoid process >4.5 mm (Fipiiie 24) 2. ied eee needed waa

39a. Height of coronoid process +4.5 mm (Figuré 24) ...0.... 0. case ested steed ena see Sorex palustris
39b, Height of coronoid process!=4.5 mm (Tisure 24) ovakt. Gait aes yas oes -Oe eters 40
40a. Heleht of coronoid process<3. 75 mm (Pismme 24) 2.2... 22. a hanes ee eee ee Sorex cinereus
A0b, Height of eoronoid process =3, 75 mm (ricure 24)" 0s... sors es ae asv eyes sae ers Sorex fumeus
E. Carnivora (Canidae, Mephitidae, and Mustelidae)
Ala. Two molars; very small to no diastema between cl and p1; anterior part of the ramus under the
Canimnennieke(Gisure STA)... . ova. case ena oeh le |e SRE eh ete Somer eie ERR ae 42
41b. Three molars; diastema between cl and p! about equal in size to p! or larger; anterior part of the .
ramus under the canine Slender (Hirotnc Sie tee tr. eile cs settee? ee cient gee eet 48
42a, Four premolars) (Pieureis Ay) te ahs. trwuitl Puee. seen. oe OG aes bo aoe fae eo ee 43
Hw) Whe er PLEUNOVANS: (ELOUGE S21), <o.crys o>. aeveuarancue anesSinde sie acanks tee mata eae ee ea ee oo
43a. Length of the mandibular tooth row <38 mm; posterior mental foramen located beneath the
hypoconid of p3; coronoid process relatively sharp (Figure 33A) ..............-.-.-. Martes americana
43b. Length of the mandibular tooth row >38 mm; posterior mental foramen often located beneath
the protoconid of ps; coronoid process rounded (Figure 338) 7:2." o)4-s ee eee Martes pennanti
44a. Bump present on the anterior part of the horizontal ramus approximately beneath p1 (Figure 34A);
pil’cleanhy smaller than m2 (FistreS4C)" 2... .. +. s+ «> s+ «aoe Beene eee nee Mephitis mephitis
44b. Absence of a bump on the anterior part of the horizontal ramus (Figure 34B); p1 larger than m2 or
aboutsmmilar in size (guns SAD)? .. 06 aqnewee ctu one erie aseirees suse. Ge eee or 45
45a. p2 often with a well-developed paraconid (Figure 35A); posterior edge of the vertical ramus with
a distinct convex notch between the coronoid process and the condylar process (Figure 36A)

eves 9: 9, EATERY Zee AR ep eC SokeY PORE EAL CEES Set ey See eee, eae ICES OR pee IA Se Se Neovison vison
4Sb. p2 often without a small anterior cusp (Figure 35B); posterior edge of the vertical ramus straight
between the coronoid process and the condylar process (Figure 36B) ..................--00-. 46

46a. Height of coronoid process <7.1 mm; length of mandibular tooth row <10 mm (Figure 37)
Jet yas ae BUOY senate gel ese. Web ANt, SHEE wet Petar: Were eee ee Be Mustela nivalis

47a. Height of the coronoid process generally <10.5 mm; length of mandibular tooth row never >16 mm
(Figure 37); posterior edge of the vertical ramus between the coronoid process and the condylar process
FOlLAUVElY Llav(PISMMOOA) oe. Sear aey tere oth cas cee eine ene chives piace. kes Mustela erminea
47b. Height of the coronoid process generally >10.5 mm; length of mandibular tooth row often >16 mm
(Figure 37); posterior edge of the vertical ramus between the coronoid process and the condylar process
With aconvex curve (Miche 3S), sink anee eee! Sop oe ObOkr ie nes oe es Mustela frenata

48a. Presence of a clearly defined step on the lower edge of the horizontal ramus anterior to the
angular process; diastemata between cl and p1, between pl and p2, and between p2 and p3 (Figure
COUN MRI AT Pn. Roe aie doa p98 Pte eee ees A Sinai. bak Ms Wan hiidh Uh ak WO | Urocyon cinereoargenteus
48b. Lower edge of the horizontal ramus anterior to the angular process smooth, without a step; only
one diastema between cl and pl (Figtte 39) fir4..c.....0s--.ccvecsss.s5.. 49

49a. Anteroposterior length of the diastema between cl and p! smaller than p1 (Figure 40A) ... Vulpes lagopus
49b. Anteroposterior length of the diastema between cl and p1 about equal to pl or larger (Figure 40B) j

Levu Oh eee F 1 eS Dey REDE TE PAS eR HELE OOGME GE RTE hor Ae See een er ae Vulpes vulpes
F. Chiroptera (Vespertilionidae)

30a. Three premolars (Migere Ani {7 5), fa ncaa eared aril ike, ices Andee he an 51
50b. Two premolars (Figure 41B) :

2014 Faureaux ET AL.: ILLUSTRATED KEY TO THE MANDIBLES OF SMALL MAMMALS OF EASTERN CANADA 37

Sia. Mandibular length >11.5 mm (Figure 42); hypoconid of p3 with lingual crest directed medially

creating a distinct himgual bales (Mgure ASA) osanio.sccacavecdscuasesecrss Lasionycteris noctivagans
Sb. Mandibular length <11.5 mm (Figure 42); hypoconid of p3 without a distinct lingual bulge
COSTS Dea ee ee ee eC es ae a ar

or Svea ar « or ati parent aha . . ini 1 ‘ 1

52a. p3 rectangular, anteroposterior length greater than labiolingual width (Figure 44A); mandibular
lenbty pemerallycx Linon (Migured2). cscsas ccogsbdltlandnereee ia ieee fiw new date Myotis septentrionalis
52b. p3 squared, anteroposterior length approximately equal to labiolingual width (Figure 44B);
fandibularilength<llinmn(Pisure 42)" te.wss < Weed 146.4) ORMdome tines) oheuwls eel | 53

53a. Mandibular length generally >10 mm; length of the mandibular tooth row generally >5.5 mm

(Figure 42) ERs ORT dee ais Wikia ee MRE Or he eee cn Mane etait ce 8 here, ae Myotis lucifugus
53b. Mandibular length generally <10 mm; length of the mandibular tooth row generally <5.5 mm

USGA? AAR Ieo teh wey oak Quer gare deen te St ek? Deel oom, ahh Dee | Myotis leibii
ere DMIOLOM LENO rl 2a MO (INS AS) dey cs ces sass ee mest ss tees Shey yeh On ee be a)
SEPANG E Goh oral erty Lea eso so ining d dp feat Ss 2 oa ae Pear erg eraser gr Pe 56

55a. Mandibular length >14 mm (Figure 42); rounded coronoid process much taller than cl (Figure
45A); p2 squared, with labiolingual width approximately equal to anteroposterior length (Figure 46A)
ES Ca ee. ee Eee epee, ee Oe ag Cee MT, aOR ee ee a een er een Pee Eptesicus fuscus
55b. Mandibular length <14 mm (Figure 42); sharp coronoid process approximately same height as cl
(Figure 45B); p2 rectangular with labiolingual width greater than anteroposterior length (Figure 46B)
alan We ANON Ad av eind <ocaned arenas Seat el hues oh aie? Sea ed SIRS Sein) & pCR Spa Kiebrahop hrcelienis: Wyte: epene Me Heer gees) ay Gi 5 Lasiurus cinereus

56a. Length of the mandibular tooth row <5 mm (Figure 42); small diastemata separate i2 from 13, 13
from cl, and p1 from p2; cl approximately same height as p2 (Figure 47A) ......... Perimyotis subflavus
56b. Length of the mandibular tooth row >5 mm (Figure 42); no diastema between 12 and 13, i3 and cl,
or pl.and.p2;:elitallenthan p2 (igure 47 Beda. < . .cRe & loom aye eiele saiereiely ere eee Lasiurus borealis

Characteristics of Barred Owl (Strix varia) Nest Sites in Manitoba,

Canada

Topp M. WHIKLO!:2:4 and JAMES R. DUNCAN?

'Department of Biological Sciences, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada
*Current address: 122 Northlands Pointe NE, Medicine Hat, Alberta, Canada
3Wildlife Branch, Manitoba Conservation, Box 24, 200 Saulteaux Crescent, Winnipeg, Manitoba R3J 3W3 Canada

‘Corresponding author: twhiklo@shaw.ca

Whiklo, Todd M., and James R. Duncan. 2014. Characteristics of Barred Owl (Strix varia) nest sites in Manitoba, Canada.

Canadian Field-Naturalist 128(1): 38—43.

During 2009 and 2010, nine Barred Owl (Strix varia) nest sites were located in Manitoba, Canada, and data on nest trees, nest
structure, and nest site habitat were collected. Nests were located in a variety of tree species, including Balsam Poplar (Populus
balsamifera), Paper Birch (Betula papyrifera), Trembling Aspen (Populus tremuloides) and Burr Oak (Quercus macrocarpa).
All nests were in tree cavities, and the majority of nests were in dead trees (67%) and had lateral openings (67%). Habitat
surrounding nest trees and estimated canopy cover were highly variable. Diameter at breast height of nest trees, cavity width,
and cavity depth were consistent and were determined to be the most reliable indicators of nest suitability for breeding Barred
Owls. We conclude that the distribution of nesting Barred Owls is influenced more by availability of suitable nest sites than by

nest tree species or nest site habitat.

Key Words: Barred Owl; Strix varia; habitat; nesting; raptor; Manitoba

Introduction

Barred Owls (Strix varia) nest in a variety of natu-
ral and anthropogenic structures (Robertson 1959;
Shackleford 1996; Houston 1999), but are considered
to be primarily cavity nesters (Mazur et al. 1997a,
1997b). They use tree cavities created by other birds,
disease, rot, and/or tree damage (Mazur ef al. 1997a;
Vaillancourt et al. 2009). Because of its reliance on large
diameter trees for nesting, the Barred Owl is considered
an indicator species of forest health (McGarigal and
Fraser 1984). The availability of nest sites limits its dis-
tribution, population size, and density (Robertson and
Rendell 1990).

As a highly adaptable species (Robertson 1959:
Shackleford 1996), the Barred Owl persists in some
habitats that have been altered by human activity (Kelly
et al. 2003; Houston 1999). However, nesting require-
ments must be met in order for avian populations to be
maintained (Robertson and Rendell 1990). The Barred
Owl’s nesting requirements are poorly documented
throughout most of its range (North America) (Mazur
et al. 1997a), and specifically in Manitoba (Holland e¢
al. 2003). io

Across its range, the Barred Owl uses forest types
along a gradient from hardwood to mixedwood to soft-
wood forests (Nicholls and Warner 1972). Hardwood
forests are rare throughout a large portion of its north-
ern range, leaving only mixedwood and mostly boreal
forests (Duncan and Kearns 1997). The link between
large cavity-nesting species and mature stands of mixed-
wood forests is known (McGarigal and Fraser 1984;
Potvin ef al. 2000; Hodson 2003; Payer and Harrison
2003).

Barred Owl management and conservation by the
government in Manitoba and elsewhere will be more

effective if we understand which factors create suitable
Barred Owl habitat within various mature mixedwood
stands. Our objectives were to locate Barred Ow! nest
sites in Manitoba, Canada, and collect data on nests,
nest trees, and nest site habitat. Describing these factors
will contribute to hypotheses about nest and habitat
selection in this species and the limits to their distri-
bution in Manitoba and elsewhere.

Study Area

Research was conducted from February 2009 to
September 2010 within the southern portion of Mani-
toba, Canada (49°0.0'N to 53°52.7'N and 95°9.2'W to
101°44.2'W). This area consists of prairie pothole, bore-
al hardwood transition, boreal taiga plain, and boreal
softwood shield regions (Zoladeski ef al. 1995). Pre-
dominant tree species in the study area were White
Spruce (Picea glauca), Black Spruce (Picea marinana),
Tamarack (Larix laricina), Jack Pine (Pinus banksiana),
Trembling Aspen (Populus tremuloides), Balsam Poplar
(P. balsamifera), and Paper Birch (Betula papyrifera).
Southern Manitoba lacks major topographic changes;
however, small shifts in elevation, along with the abun-
dance of wetlands and waterways, create a highly vari-
able habitat (Zoladeski et al. 1995).

Methods

Barred Owl nest sites were located using nocturnal
audio surveying and diurnal audio playback with pas-
sive observation during the breeding season (February
— June in 2009 and 2010) (Frith er al. 1997; Whiklo
2011). Survey transects were laid out in areas based on
Barred Owl detection data obtained from the Manitoba
Nocturnal Owl Survey (JRD, unpublished data: op. cit.
Duncan and Kearns 1997) and historical accounts, and

2014

transects were also laid out in suitable habitat adja-
cent to known areas of Barred Owl activity. In total
approximately 1321 km of transect lines were surveyed
in 2009 and 2010. Survey locations were situated 1.6 km
apart along survey transects where playback of Barred
Owl vocalizations were used to elicit responses (Whik-
lo 2011). Areas where Barred Owls were detected dur-
ing nocturnal surveys were searched during daylight
for active nests.

Nest trees were categorized as live or dead, and tree
species, the height of the nest above ground, and diam-
eter at breast height (DBH) were recorded. Diameter
at breast height was calculated by measuring the cir-
cumference of the tree and then calculating the diam-
eter: D= C /x. Cavity height (distance from the lowest
point inside the nest to the highest point inside the nest),
cavity width (distance from the furthest right-hand point
inside the cavity to the furthest left-hand point inside
the cavity), and cavity depth (distance inside the cavity
perpendicular to cavity width) were measured; nest type
(cavity, stick, other) and cavity orientation (lateral or
apical) were also recorded.

Habitat within a 30 m circular plot surrounding the
nest trees was categorized using Manitoba Forest In-
ventory classifications (Zoladeski et al. 1995), and the
percentage canopy cover was estimated (Whiklo 2011).
All measurements are reported as mean and standard
deviation (SD).

Results

A total of nine Barred Owl nests were located in
2009 and 2010 within a 25 000 km? rectangle in south-
eastern Manitoba. All nests were cavity type structures;
six were lateral cavities and three were apical cavities
(Table 1). Six nest trees were dead and three were liv-
ing (Table 1). Five nests were found in Balsam Poplar,
two in Paper Birch, one in Trembling Aspen, and one in
Burr Oak (Table 1). The mean nest height above ground
was 7.7 m (SD 2.6). The mean diameter at breast height
of nest trees was 49.2 cm (SD 18.9). The mean cavity
height was 71.8 cm (SD 46.9), the mean cavity depth
was 42.1 cm (SD 33.0), and the mean cavity width was
27.3 cm (SD 5.4) (Table 2).

Four nest trees were located in Balsam Poplar mixed-
wood (V1) stands, two in Black Ash (Fraxinus nigra)
hardwood (V2) stands, one in a White Spruce/Balsam
Fir (Abies balsamea) (V21) stand, one ina Miscella-
neous Hardwood (V3) stand, and one in an area that
could not be classified due to a lack of living trees (a
pond created by an American Beaver, Castor canaden-
sis) (Table 1). The mean estimated canopy cover was

42.8% (SD 27.2) (Table 2).

Discussion .
There are a considerable number of studies that ex-

amine one or more aspects of the nest site structure, the
nest tree species, and/or the habitat associated with the
nest sites of the Barred Owl (Nicholls and Warner 1972;

TABLE |. Data for nine Barred Owl (Strix varia) nest sites in Manitoba, Canada (2009 to 2010).

Manitoba forest classification!

Nest tree status Nest type

Nest tree species

Owl nest site

V1: Balsam poplar hardwood and mixedwood

Dead Lateral cavity

Balsam Poplar

Cow Moose Lake (Barred Owl 4)

Watson P. Davidson Wildlife

V1: Balsam poplar hardwood and mixedwood
V1: Balsam poplar hardwood and mixedwood

Apical cavity
N/A*

Dead

Paper Birch

5)

(=

Management Area (Barred Owl
Stead (Barred Owl 11)

Dead Lateral cavity

Balsam Poplar

Dead Lateral cavity

Balsam Poplar
Balsam Poplar

Otter Falls (Barred Owl 20)

V2: Black ash (White elm) hardwood
V21: White spruce/Balsam fir shrub

Lateral cavity

Live
Dead

Nutimik Lake (Barred Owl 27)

Apical cavity

Balsam Poplar
Paper Birch

West of Woodridge (Barred Owl 31)
East of Piney (Barred Owl 36)

WHIKLO AND DUNCAN: BARRED OWL NEST SITES IN MANITOBA

V1: Balsam poplar hardwood and mixedwood
V2: Black ash (White elm) hardwood

V3: Miscellaneous hardwoods

Apical cavity

Dead

J

Contour area (Barred Owl

Live Lateral cavity

Trembling Aspen
Burr Oak

Lateral cavity

Live

Dencross (Barred Owl 56)

*Habitat was determined to be unclassifiable according to Manitoba forest classifications due to lack of living trees.

' Zolandeski et al. (1995)

THE CANADIAN FIELD-NATURALIST

Vol. 128

TABLE 2. Further data for nine Barred Owl (Strix varia) nest sites in Manitoba, Canada (2009 to 2010).
eee — ese

Nest tree Height of the

diameter at nest above Cavity Cavity Cavity Canopy

breast height the ground height depth width cover
Owl nest site (cm) (m) (cm) (cm) (cm) (%)
Cow Moose Lake (Barred Owl 4) 43.1 4.5 68.8 26.4 Ni fere' 30
Watson P. Davidson Wildlife

Management Area (Barred Owl 5) 42.9 7.4 24.9 B29) 29.4 39

Stead (Barred Owl 11) 50 10.2 156.0 127.0 24.5 60
Otter Falls (Barred Owl 20) 39.7 6.4 AI 22.8 26.1 0
Nutimik Lake (Barred Owl 27) 56.2 2A 67.9 35.9 35.9 70
West of Woodridge (Barred Owl 31) B34) 5.8 11.4 YS) 2S 75
East of Piney (Barred Owl 36) 33},3) 5.4 42.0 29.8 ZAR? 5
Contour area (Barred Owl 55) 48.7 7.6 102.2 30.8 35,1 50
Dencross (Barred Owl 56) 95.5 9.9 Sy) 51.0 24.0 60
Mean (SD) 49.2 (18.9) 7.7 (2.6) 71.8 (46.9) 42:1(33.0) 27.354) 42.8 (27.2)

Haney 1997; Mazur et al. 1997a, 1997b; Postupalsky
et al. 1997; Winton and Leslie 2004; Olsen ef a/. 2006;
Grossman ef a/. 2008; Singleton ef a/. 2010) and gen-
eral Barred Owl habitat associations (McGarigal and
Fraser 1984; Booth and Harrison 1997; Mazur et al.
1998; Hamer et al. 2007; Russell 2008). These studies
vary considerably, as described in more detail below,
in the way study areas were selected, in the size and
habitat fragmentation of the study areas, and in the size
and measurement of nest habitat plots. However, there
is less variation in the way nest trees and nest sites were
measured.

This variation in methodology limited our ability to
compare results; nevertheless, some Barred Owl nest
site characteristics were consistent across studies.

Nest type

In contrast to other studies (Mazur et al. 1997a;: Pos-
tupalsky et al. 1997; Olsen et al. 2006), all nests (n = 9)
located in the study were in tree cavities (Table 1).
Mazur et al. (1997a) reported that 5 of 15 Barred Owl
nests (33%) in the study in the boreal forest of Sas-
katchewan were in structures other than tree cavities; in
witch’s broom (the dense branching caused by Arceu-
thobium spp. in a White Spruce tree), in Red Squirrel
(Tamiasciurus hudsonicus) nests, or in stick nests. In
a study in the boreal mixedwood forest in Alberta
(Olsen et al. 2006), 9 of 10 nest sites (90%) were in
tree cavities (one Barred Owl nested in a stick nest).
In Michigan, in hardwood (deciduous) and mixed for-
est habitat, Postupalsky er a/. (1997) described 13 of
57 nests (23%) as being open sites, including hawk
(Red-shouldered Hawk (Buteo lineatus) or Broad-
winged Hawk (Buteo platypterus) and Northern Gos-
hawk (Accipiter gentilis)) stick nests, a ground nest,
a flat area in the fork of a Yellow Birch (Betula alle-
ghaniensis), and a nest platform intended for Great
Horned Owls (Bubo virginianus); the remainder were
in tree cavities (n = 26) or nest boxes (n = 18).

The likelihood of finding an open Barred Owl nest
in Manitoba would presumably increase with increased
effort and sample size. However, it is noteworthy that,
even though Barred Owls are known to use artificial
open nests (Olsen ef a/. 2006), none were found nesting
on a cumulative total of 2527 natural and/or artificial
open stick platform nests in a variety of habitats checked
for raptors over a 27-year period (1984-2010) in the
same 25 000 km? study area in southeastern Manitoba
(Duncan 1992; JRD, unpublished data).

The aforementioned studies (Mazur et al. (1997a),
Postupalsky et al. (1997), and Olsen et al. (2006)) varied
considerably in the way the study areas were selected or
described, in the size of the study areas, in the methods
used to find nests, in the forest habitat composition/
fragmentation, and in other quantified ways (i.e., prey
density, human disturbance). For example, this study
was larger (~25 000 km?) with varied habitat, the study
described in Mazur et al. (1997a) was conducted with-
in a 3 874 km* national park, the study described in
Olsen et al. (2006) was a 800 km? predetermined
area, and two study areas (28 km? and an undefined
larger area) were studied in Postupalsky er al. (1997).

Smaller fragmented study areas or isolated protect-
ed areas (1.¢., national parks) may vary in terms of the
availability of cavity nests, the prey density, the forest
habitat, and/or intra and interspecific competition, re-
sulting in the variation observed in the proportion of
nest type use by breeding Barred Owls. How these fac-
tors affect the availability of suitable cavity nest sites
and the proportion of Barred Owls using open nest sites
is unknown. However, the propensity of Barred Owls
for cavity nests likely results from natural selection:
Barred Owls nesting in cavities experience greater
reproductive success than those that use open nests
(Postupalsky et al. 1997).

Nest cavity characteristics

Given the importance of nest cavities to Barred Owl
reproduction, we recorded a series of measurements.
Cavity height and depth ranged widely (height ranged

2014

from 11.4 to 156.0 cm and depth ranged from 22.3 to
127.0 cm) with high standard deviations, whereas cav-
ity width was remarkably consistent (21.2 to 35.9 em)
(Table 2), despite the variation in nest tree species and
Status (live or dead) (Table 1). Cavity depth varied the
most, perhaps as a result of the variable and sometimes
advanced stages of tree decay, e.g., the nest site near
Stead (Table 2). Postupalsky ef al. (1997) recorded a
similar mean cavity width (26.9 cm, range 18-44,
n= 25), but did not report cavity depth measurements
(as defined in this study) or standard deviations. Nest
cavity measurements were not reported in other studies.

Nest tree diameter at breast height

Mean diameter at breast height of nest trees in this
study (49.2 cm, SD 18.9) was consistent with that re-
ported in other studies. Mazur et al. (1997a) recorded
an average diameter at breast height of 47.4 cm (SD
12.8, n = 15), despite recording considerably higher
values for the height of nests from the ground (13.3 m,
SD 4.1) than this study (7.7 m, SD 2.6) (Table 2). Olsen
et al. (2006) recorded an average diameter at breast
height of 51.6 cm (SE 4.3), along with a relatively inter-
mediate nest height above ground (10.4 m, SE 2.1).

There were relatively large differences in many nest
tree variables among these studies (e.g., nest tree height,
nest height, proportion of cavity nest structures, and
nest tree species); therefore, the similarities in the diam-
eter at breast height of nest trees suggest it is a valid
and practical indicator of Barred Owl nest tree suit-
ability.
Nest tree species, percentage canopy cover, and forest
stand composition

Barred Owls nested in four hardwood tree species in
this study (Table 1), and this variation was similar to
that found in other studies. Mazur ef al. (1997a) re-
ported Barred Owl nests in both softwood (coniferous)
and hardwood tree species, including White Spruce
(n = 5), Trembling Aspen (n = 5), Balsam Poplar (n=
A), and Paper Birch (n = 1). Olsen et al. (2006) docu-
mented Barred Owl nests in fewer tree species in a
smaller study area: Balsam Poplar (n = 8) and Trem-
bling Aspen (” = 2). Barred Owls use a variety of nest
tree species, live or dead, and they readily breed in arti-
ficial nest boxes placed in a variety of trees (Postupal-
sky et al. 1997). It is therefore unlikely that Barred
Owls choose a nest site based on tree species per se.

High percentage forest canopy cover has been cit-
ed as a determining factor in Barred Owl selection of
breeding habitat, possibly because it provides solar
insulation (Nicholls and Warner 1972; Haney 1997;
Winton and Leslie 2004; Grossman ef al. 2008), but
the influence of forest canopy may depend on the size
of the area that was measured. In this study, canopy
cover was measured within a 30 m circular plot cen-
tered on the nest tree, and it did not appear to influ-
ence Barred Owl nest tree habitat use: more than half
the sample had a canopy cover of <50% (Table 2).

WHIKLO AND DUNCAN: BARRED OWL NEST SITES IN MANITOBA 4]

Mazur et al. (1997b) used a similar small-scale plot
(11.3 m radius) with the nest tree at the centre, and
reported a somewhat higher mean percentage cover of
57% (SD 17); this was not significantly different from
random plots. Other studies reported yet higher per-
centage canopy cover within larger Barred Owl home
ranges: 96% (SE 1.1) (Haney 1997), 62.8% (Winton
and Leslie 2004), utilized “dense” cover disproportion-
ately (no values given) (Nicholls and Warner 1972),
>66% (Grossman et a/. 2008), and >56% (Singleton ef
al, 2010).

Forest stands within the 30 m circular plots (centered
on nest trees) were classified as one of three types of
stands: hardwood and mixedwood, softwood shrub, or
unclassified (American Beaver pond) (Table 1). This
variation in nest habitat use is reflective of the great
variety of forested areas over the considerable North
and Central American range of the Barred Owl, from
swamps and riparian areas to upland regions (Mazur
and James 2000). This variation of forest stand nesting
habitat use suggests that the Barred Owl is a forest habi-
tat generalist.

Management of forests for Barred Owls

Strong selective pressure on Barred Owls appears
to have resulted in their propensity for nest cavities in
trees. Observed higher reproductive success in cavity
nests implies nest site selection for cavities by this
species (Postupalsky ef al. 1997). This conclusion is
supported both by our results and by those of others,
in which the most consistent nest characteristics and
nest habitat characteristics reported are the width of the
nesting cavity and the diameter at breast height of the
nest tree. Other Barred Owl nest habitat characteristics
discussed herein vary considerably across the range of
the Barred Owl. Apart from its effective dependence on
suitable nest tree cavities, the Barred Owl is otherwise
generally considered a forest habitat generalist (Mazur
and James 2000).

The persistence of Barred Owl populations depends
on the maintenance of forests with trees with a mini-
mum diameter at breast height capable of producing
cavities large enough for this large cavity-nesting
species (Haney 1997). Knowledge of ecological factors
and processes that promote the formation of suitable
nest tree cavities is also critical to the maintenance of
Barred Owls in a managed forest environment.

Barred Owls are associated with water (Mazur ef
al. 1997b; Hamer et a/. 2007), mature or “old-growth”
forest stands (McGarigal and Fraser 1984; Mazur ef
al. 1998), and mixedwood or hardwood stands (Booth
and Harrison 1997; Mazur et al. 1997b; Russell 2008).

The role and importance of heart rot in hardwood
species in the formation of nest cavities, as well as the
role of snags in an ecosystem, are well documented
(Thomas et al. 1979; Witt 2010). Barred Owl nest cav-
ities found in this study were natural and had resulted
from damage to and decay of the tree. These cavities

42 THE CANADIAN FIELD-NATURALIST

were not readily attributable to excavation by primary
cavity nesters.

Cavities not created by primary cavity nesters are
often created by tree decay and rot (Bunnell ef al.
2002). Fungal rot is prevalent in older and/or larger
stands of trees (Witt 2010) and has positive effects for
both primary and secondary cavity nesters (Bunnell e¢
al. 2002). Higher levels of moisture and humidity, fac-
tors found at sites within close proximity to water, in-
crease the rate of decay in trees (Jackson and Jackson
2004). In Manitoba, hardwood species decay at a high-
er rate than most softwood species: annual losses of
hardwood species to decay are double that of softwood
species (Brandt 1995).

Barred Owl conservation would benefit from the
development and use of a standard methodology to
characterize nest sites and nesting habitat. Standard
methodology would allow the results from studies
across this species’ range or through time to be com-
pared. We also recommend that tree species composi-
tion, diameter at breast height, and ecological forest
decay indicators be developed and used to identify pri-
ority Barred Owl habitat conservation areas where for-
est habitat loss affects the viability of local Barred Owl
populations.

Acknowledgements

This research was supported by Manitoba Conser-
vation, Manitoba Hydro, the University of Manitoba,
the Manitoba Model Forest Inc., the Raptor Research
Foundation, and the Winnipeg Foundation. Thanks to
Robert Nero and Terry Galloway for reviewing drafts
of this manuscript and providing comments. We also
thank the two anonymous reviewers for their helpful
comments.

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Received 15 March 2013
Accepted 29 May 2013

Yellow Warblers (Setophaga petechia) Rear Second Broods in Some
Years at Delta Marsh, Manitoba

SPENCER G. SEALY

Department of Biological Sciences, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada; email: sgsealy@ec.uman
itoba.ca

Sealy, Spencer G. 2014. Yellow Warblers (Setophaga petechia) rear second broods in some years at Delta Marsh, Manitoba.
Canadian Field-Naturalist 128(1): 44-49.

Twenty cases of double brooding by colour-marked Yellow Warblers (Setophaga petechia) were recorded in 5 of 11 years
(1975-1986, no data collected in 1977) during studies of breeding ecology in the dune-ridge forest at Delta Marsh, Manitoba
(1 pair in 1975, 3 pairs in 1976, 3 pairs in 1984, 9 pairs in 1985, and 4 pairs in 1986). At least one member of each of the 20 pairs
was marked. Eleven pairs re-used their first nest for the second attempt, whereas 9 females built a new nest, in 5 cases because
the original nests had disintegrated. Four of the second nests (3 in 1985 and | in 1986) were parasitized by Brown-headed Cow-
birds (Molothrus ater). All 20 first nests produced at least one young, a condition for double brooding, and 13 second nests,
including 3 that were parasitized, were successful. Failure of about 60% of annual nesting attempts at Delta Marsh may contribute
to the low number of pairs with double broods recorded in some years and the absence of double brooding in years of comparable
phenology. This is the first published evidence of double brooding in the Yellow Warbler.

Key Words: double broods; Yellow Warbler; Setophaga petechia; Delta Marsh; Manitoba

Introduction but only in some years. Other workers have suspected
Knowledge of the number and success of broods double brooding by Yellow Warblers, but their obser-
attempted by birds in each breeding season is important vations were based on one or only a few nests attended
for an understanding of the dynamics of avian popula- _ by unmarked birds (e.g., Stoner 1932; Salt 1973; Ban-
tions and life history evolution (Martin 1987). One of — croft 1979). Confirmation of double brooding by Yel-
the components of the life cycle is the number of broods low Warblers at the northern latitude of Delta Marsh
individuals typically rear in each breeding season, suggests that double brooding may be a regular occur-
whether one, two or even more in certain years when _ rence in this species’ breeding biology.
conditions are suitable. This contrasts with the replace-
ment of clutches and broods that may be depredated Study Area
or otherwise fail, possibly due to inclement weather. Data from first and second broods of the Yellow
An irrefutable determination that birds have raised Warbler were recorded from nests discovered in the
one brood and attempted to raise another requires care- dune-ridge forest that separates Lake Manitoba and
ful monitoring of colour-marked individuals or radio- | Delta Marsh, Manitoba, 50°11'N, 98°19"W (Goossen
tracking throughout the entire breeding season. In some and Sealy 1982; MacKenzie 1982: MacKenzie et al.
species, researchers have recorded double brooding in- 1982; Pohajdak 1988). Nests were monitored over 11
frequently, often as isolated cases (e.g., in the Northern _ years (1975-1986, except 1977) by J. P. Goossen (1975
Yellow-throat, Geothlypis trichas (Stewart 1953); in and 1976) and by me in the other years, although some
Kirtland’s Warbler, Setophaga kirtlandii (Radabaugh __ nests in the latter years were located by co-workers con-
1971); in the Bobolink, Dolichonyx oryzivorus (Gavin ducting other studies. The number of Yellow Warbler
1984); and in the Dickcissel, Spiza americana (Bolinger nests, which included replacement nests, monitored in
and Maddox 2000)), whereas others have recorded — each year of the study was: 119 in 1975, 148 in 1976,
double brooding by 648% of pairs, with successful 64 in 1978, 59 in 1979, 44 in 1980, 126 in 1981. 157 in
second broods increasing an individual’s annual pro- 1982, 237 in 1983, 260 nests in 1984, 225 in 1985. and
ductivity (e.g., in the Black-throated Blue Warbler, 241 in 1986.
Setophaga caerulescens (Holmes et al. 1992); in the Yellow Warblers investigated in this study nested in
Hooded Warbler, Setophaga citrina (Evans Ogden and —_a 7-km portion of the dune-ridge forest (~80 m wide;
Stutchbury 1996); and in the Louisiana Waterthrush, 56 ha), abutted by the lake on the north side and an
Parkesia motacilla (Mulvihill et al. 2009)). extensive marsh of cattail (Typha latifolia) and reed
During an Il-year study of breeding and feeding (Phragmites communis) on the other (see map in Sealy
ecology of Yellow Warblers (Setophaga petechia) ina 1980a). This portion of the ridge forest stretched from
riparian habitat at Delta Marsh, Manitoba, from 1975 the west at Cram Creek eastward along the property
to 1986 (no data were collected in 1977), co-workers of the Portage Country Club to the eastern edge of the
and I recorded irrefutable evidence of the rearing of — property of the Delta Marsh Field Station (University
second broods after young had fledged from first nests, | of Manitoba). Yellow Warblers nested in the ridge for-

44

2014 SEALY: DOUBLE BROODING BY YELLOW WARBLERS AT DELTA MARSH, MANITOBA 45

est predominantly in Sandbar Willow (Salix interior),
Manitoba Maple (Acer negundo), Red-berried Elder
(Sambucus pubens), Pin Cherry (Prunus pensylvanica),
Cc hoke Cherry (P. virginiana), and Red-osier Dogwood
(Cornus stolinifera). Peach-leaved Willow (S. amyg-
daloides), Green Ash (Fraxinux pennsylvanica), and
Eastern Cottonwood (Populus deltoides) also contri-
buted to the overstory vegetation in this riparian
habitat, but few Yellows Warblers nested in them
(MacKenzie et al. 1982).

Methods

Adults of this sexually dichromatic, seasonally mon-
ogamous species (Reid and Sealy 1986) were mist-
netted and colour-marked as after-hatch-year (AHY)
males or females, following an annual banding pro-
tocol established late in 1974 and continued in each
subsequent year of the study (e.g., Sealy 1980b; Sealy
and Biermann 1983). Thus the first spring in which
marked individuals were present in the study area was
in 1975.

In addition, males and females were opportunistical-
ly captured and colour-banded near their nests (Cosens
and Sealy 1986; Hobson and Sealy 1989). Colour-
marking of nestlings and hatch-year (HY) individuals
mist-netted prior to fall migration also began in 1975,
and birds marked in that and subsequent years resulted
in individuals of known age present in the study area in
successive years. These individuals originally received
a single coloured band, which, in combination with an
aluminum band, denoted the year of hatch, so these
birds became recognizable in their first spring as second-
year (SY) individuals, and older in succeeding years.
Upon recapture, year-marked individuals received two
additional coloured bands that uniquely identified them.
This marking program resulted in a sample of nests each
year that were attended by at least one marked male
or female (in several cases both adults were marked).

Yellow Warblers nested at densities up to 29 pairs/ha
in the ridge forest in those years (Goossen and Sealy
1982: also see Sealy 1995). This density allowed dozens
of nests to be discovered in most years before clutches
were initiated, enabling day-to-day inspections as eggs
were laid and inspections every one to four days through
fledging of first and second broods. The extremely high
nesting densities (Goossen and Sealy 1982), however,
meant that only a small undetermined number of indi-
viduals could be marked, thus precluding the determi-
nation of the proportion of pairs that attempted second
broods in each of the years in which double brooding
was recorded.

| am confident that second broods were not attempt-
ed by any pairs in the 6 years in which none were re-
corded because I monitored a sample of nests, both
those attended by marked birds and those attended by
unmarked birds, equally diligently throughout all breed-
ing seasons of the study (see Sealy 1995; Guigueno
and Sealy 2010; Mazarolle e/ al. 2011).

Results

Double brooding was recorded in 20 pairs of Yellow
Warblers of known age in 5 of the 11 years of the study
(Table 1): 1975 (n = 1 pair), 1976 (n = 3 pairs), 1984
(n = 3 pairs), 1985 (n = 9 pairs), and 1986 (n = 4 pairs).
At least one member of each pair was marked. Males
(at 8 nests), females (at 5 nests), or both males and fe-
males (at 7 nests) were marked, thus permitting irre-
futable confirmation of double brooding (Table 1).

These data suggest that all pairs remained together
during the second nesting attempt, although polygyny,
albeit infrequent in this population (Sealy 1984; Reid
and Sealy 1986), could have resulted in a change of
mate at nests where only one of the adults was marked.
This would not change the fact, however, that double
brooding occurred.

Among the 20 pairs with double broods were 4
whose second clutch was parasitized by a Brown-
headed Cowbird (Molothrus ater) (3 in 1985 and | in
1986). None of the first nests of these or any of the
other pairs that went on to attempt to raise a second
brood were parasitized (Table 1). The frequency of par-
asitism by Brown-headed Cowbirds in this population
of Yellow Warblers (also see Sealy 1995) was 26.1%
of 119 nests monitored in 1975, 23.0% of 148 nests in
1976, 18.1% of 260 nests in 1984, 18.7% of 225 nests
in 1985, and 17.8% of 241 nests in 1986.

Each pair’s first nesting attempt successfully fledged
at least one young, a necessary condition for double
brooding. At one of the first nests in 1985, however,
incubation was delayed about 3 days while the female
replaced 3 of 4 eggs that had gone missing from the
original nest (footnote 3 of Table 1). Clutches at 18 of
the 20 first nests (90%) were initiated by 2 June (Table
1) in years where first eggs in the population were ini-
tiated on or up to 7 days before this date (also see table
III in Sealy 1995).

All 7 marked pairs remained together for their sec-
ond nesting attempt, lending support to the assump-
tion that the members of the other pairs also did not
change (although in these cases it was confirmed only
that the same marked individuals (8 males and 6 fe-
males) remained for the second attempt). Eleven pairs,
including the 4 pairs whose second clutches were para-
sitized, used their first nest for the second attempt. One
female whose nest had been parasitized built a new
nest over top of the one Brown-headed Cowbird and
the two Yellow Warbler eggs that had been laid in the
original nest. Nine females built a new nest for the
second attempt (Table 1), in 5 cases because the orig-
inal nests had disintegrated. The other 4 pairs built a
new nest in another location nearby. None of the indi-
viduals with double broods was among the pairs that
double brooded in previous or subsequent years, and
no young produced in first or second nests, although
banded, were subsequently captured in the study area.

Mean dates (nearest day and standard deviation) of
initiation of the 20 first and second clutches were 30

Vol. 128

THE CANADIAN FIELD-NATURALIST

46

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2014

May (SD 2.9 days) and 30 June (SD 3.2 days), respec-
tively, and the mean number of days between initia-
tion of first and second clutches was 30.2 days (SD
2.3 days). Mean initiation of first clutches by females
with double broods followed the earliest dates that
clutches were initiated in the population within | day
(1975), 3-9 days (1976), 0-2 days (1984), 0-3 days
(1985), and 1-3 days (1986). Initiation of second
clutches preceded the last dates of the season for fe-
males still attempting to lay clutches by 6 days (1975),
0-7 days (1976), 3—7 days (1984), 7-11 days (1985),
and 1-14 days (1986). The mean number of days be-
tween dates of fledging of the last nestling from first
broods and dates of clutch initiation at unparasitized
second nests (7 = 16 nests) was 3.6 days (SD 1.5 days):
2 days (n = 2 nests), 3 days (7 nests), 4 days (4 nests),
5 days (1 nests), 6 days (1 nest), and 8 days (1 nest).

Parasitized nests were excluded from this analysis
because of possible interference with initiation of the
Yellow Warblers’ clutches; indeed, a delay of 3 days in
the completion of the clutch occurred at one nest par-
asitized in 1986 (Table 1), in which the Yellow Warbler
buried the eggs. This nest eventually failed (see Clark
and Robertson (1981) and Guigueno and Sealy (2010)
for details of the chronology of laying by Yellow War-
blers parasitized by Brown-headed Cowbirds).

Discussion
Documentation of double brooding

Data presented in Table | provide the first published
evidence that Yellow Warblers attempt second broods
in some years. Double brooding by Yellow Warblers
has not been confirmed elsewhere in North America
(but see below), but it has been confirmed in a non-
migratory population on the Galapagos Islands, Ecua-
dor (Snow 1966), located at the equator. Snow record-
ed a single marked pair that attempted to rear two
broods in each of two consecutive years, but in both
years the second attempt failed. He added (Snow 1966,
page 46) that the “season is amply long enough for
[rearing second broods], and two broods are probably
common.” The fact that double brooding has not been
recorded at other sites, tropical or temperate, probably
reflects the lack of observations of uniquely marked
individuals throughout the breeding season.

These data support an earlier observation of an un-
marked pair of Yellow Warblers that suggested dou-
ble brooding in Manitoba, along the west shore of Lake
Winnipeg, in 1978 (Bancroft 1979; also see Gollop
1979). Dates of initiation of the first and second clutch-
es were in line with those presented for Delta Marsh
and Lake Manitoba in Table |. This pair’s first nest
was under construction on 26 May when it was first
observed, and by 2 June it contained two Yellow War-
bler eggs. An undetermined number of young had left
the nest by 23 June, but by 7 July, the (same?) female
was incubating 3 more eggs in the same nest from

SEALY: DOUBLE BROODING BY YELLOW WARBLERS AT DELTA MARSH, MANITOBA 47

which the first brood had fledged. Again, an undeter-
mined number of young, in the second brood, fledged
by 27 July, as noted by feces on foliage near the empty
nest and audible vocalizations of fledglings and adults
in a nearby hedge (Bancroft 1979), Both adults fed
young from the first brood, but it was not reported
whether both cared for young in the second brood.

Post-hoc examination of the nest by H. W. R. Cop-
land of the Manitoba Museum in Winnipeg after the
second brood had fledged revealed the nest had been
parasitized by a Brown-headed Cowbird during the
laying of the second clutch, whereupon the Yellow War-
bler buried the Brown-headed Cowbird egg plus two
of her own eggs, one of which was broken, under a
new nest, and started again. The time spent burying the
Brown-headed Cowbird egg and her own eggs, recon-
structing the nest, and replacing the initial eggs of the
second clutch suggests the final clutch was initiated on
28 or 29 June, consistent with the dates of initiation of
second clutches at Delta Marsh, including the nest par-
asitized in 1986 in which the Yellow Warbler buried the
Brown-headed Cowbird and her first two eggs (Table
1).

Additional irrefutable records of double brooding by
Yellow Warblers came to light during the review of this
manuscript. One is in a population nesting at the north-
ern limit of the species’ range, near Inuvik, Northwest
Territories (68°21'N, 133°45'W), and the other is near
Revelstoke, British Columbia (50°57'N, 118°10'W)
(Anna Drake, personal communication). Near Revel-
stoke, four cases of double brooding were recorded
over eight years at a latitude similar to that of Delta
Marsh. In 2004, a second-year female paired with an
after-hatch-year male initiated first and second clutch-
es on 25 May and 23 June, respectively. Three young
fledged from each clutch. In 2005, two cases involved
after-second-year females (one the female with the
double brood in 2004 paired with the same male),
which initiated first clutches on 27 and 30 May (fledg-
ing 5 and 3 young, respectively), and second clutches
on 24 and 28 June, respectively (fledging 3 young from
each nest). In 2011, an after-second-year female pro-
duced a maximum of three fledglings in the first clutch
(initiated 30 May) and she produced a second clutch
(date of initiation unknown). Dates of initiation of first
and second clutches in this population were a few days
earlier than those recorded at Delta Marsh (Table 1).

At Inuvik, Drake recorded a marked pair of after-
hatch-year individuals that reared two broods in 2010,
although the male was not observed while the female
tended the second brood. First and second clutches
were initiated on | June and | or 2 July, respectively,
and the young fledged on 26 or 27 June and 23 or 24
July, respectively. These dates of initiation were re-
markably similar to those recorded for first and sec-
ond clutches at Delta Marsh (Table 1). Of the two cases
at Delta Marsh in which first clutches also were initi-

48 THE CANADIAN FIELD-NATURALIST

ated on | June (both in 1984) (Table 1), the second
clutches were initiated on 30 June, one or two days ear-
lier than those at Inuvik.

Initiation of the earliest clutches by Yellow Warblers
in other populations nesting at the northern limit of the
range—Churchill, Manitoba (Briskie 1995), Yukon
(Sinclair e¢ al. 2003), and Alaska (Kessel 1989)—has
not been recorded before the middle of June. Briskie
(1995, page 539), commenting on the length of the
breeding season of the Yellow Warbler at Churchill,
observed some clutches that were replaced after fail-
ure early in the season, but found “... no evidence of
double-brooding and [stated that] it is unlikely to occur.
In the average year, even the earliest nests do not fledge
young before late July and any second brood would
require birds to remain in the area well past the end of
the short subarctic summer.” The dynamics of breed-
ing of this and other species at the northern limit of
their range require additional study, especially when
temperatures are increasing.

Implications of double brooding

The proportion of pairs with double broods recorded
in the population in suitable years at Delta Marsh is
not known but its determination was likely influenced,
in the first instance, by the availability of marked pairs
and, more importantly, by the number of marked indi-
viduals whose first nests managed to survive through
fledging, where ~60% of all nesting attempts by Yellow
Warblers are depredated or fail due to inclement weath-
er (Goossen and Sealy 1982; Guigueno and Sealy
2010). Probably no more than 5—10% of pairs attempt-
ed to rear second broods in the 5 years in which they
were recorded, but double brooding also may have
occurred in 4 additional years of the study (1978, 1980—
82, see table III in Sealy 1995) that experienced similar
nesting phenology. Nests tended by the marked pairs
in those years may have failed before second clutches
could be initiated, although if double brooding occurred,
I should have recorded re-use of some first nests by
unmarked pairs. Spring temperatures in 1979 and 1983
were lower and first clutches were not laid until 9 and
12 June, respectively (table III in Sealy 1995), proba-
bly too late for double brooding.

Within the liniits of the small sample size of Yellow
Warbler nests parasitized by Brown-headed Cowbirds
(all of them second nests) (Table 1), it is noteworthy
that parasitism was rejected at one of the four nests.
Females tend to bury Brown-headed Cowbird eggs in
the latter portion of the clutch-initiation season, but
almost all females accept Brown-headed Cowbird eggs
towards the end of the breeding season, when they are
running out of time (Clark and Robertson 1981; Sealy
1995; Guigueno and Sealy 2010). This single instance
of burial was unusual because it was initiated only two
or three days before the last Yellow Warbler eggs of the
season were laid at Delta Marsh in 1986 (table III in
Sealy 1995).

Vol.

Short-lived individuals (Klimkiewicz ef al. 1983)
might be expected to raise more than one brood in
years when conditions are favourable (Evans Ogden
and Stutchbury 1996), but they would have to be res-
ponsive to early spring temperatures. In a study of
arrival and clutch initiation by Yellow Warblers over
30 years at Delta Marsh, Mazerolle er al. (2011) report-
ed that individuals exhibited considerable plasticity
in the dates of clutch initiation in response to mean May
temperatures. This plasticity was observed only at the
beginning of the breeding season, as Yellow Warblers
molt and migrate early (Morton 1976) and arrive on the
wintering grounds from Mexico south to Peru where
they compete for feeding territories, as early as late
August (Neudorf and Tarof 1998). Rigid scheduling
of molt and migration may preclude attempts to rear
second broods, except in early seasons, but this may
become more frequent with increasing temperatures.

Acknowledgements

I thank J. Briskie, D. Busby, H. den Haan, J. Goos-
sen, K. Hobson, D. MacKenzie, R. Olenick, G. (Bier-
mann) Pohajdak, J. Porter, and G. Sutherland for assis-
tance in the field during the early years of the research
on songbirds at Delta Marsh, Manitoba, which includ-
ed considerable assistance with banding and colour-
marking. I thank an anonymous reviewer and Anna
Drake for constructive comments offered during the
review of the manuscript. In addition, I would like to
thank Anna for allowing me to include the record of
double brooding by a pair of Yellow Warblers at an
even more northerly latitude and D. J. Green, M. Hepp,
and S. P. Quinlan for providing data from a site near
Revelstoke, British Columbia. This work was funded
chiefly by the Natural Sciences and Engineering Re-
search Council of Canada, augmented by substantial
in-kind support provided by the Delta Marsh Field
Station (University of Manitoba).

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Received 6 June 2013
Accepted 8 July 2013

Asynchronous Breeding and Variable Embryonic Development
Period in the Threatened Northern Leopard Frog (Lithobates pipiens)
in the Cypress Hills, Alberta, Canada: Conservation and Management
Implications

. Z
LEA A. RANDALL!:3, LYNNE D. CHALMERS!:2, AXEL MOEHRENSCHLAGER!2, and ANTHONY P. RUSSELL

‘Centre for Conservation Research, Calgary Zoological Society, 1300 Zoo Road NE, Calgary, Alberta T2E 7V6 Canada
Department of Biological Sciences, University of Calgary, 2500 University Drive NW, Calgary, Alberta T2N 1N4 Canada
3Corresponding author: lear@calgaryzoo.com

Randall, Lea A., Lynne D. Chalmers, Axel Moehrenschlager, and Anthony P. Russell. 2014. Asynchronous breeding and

variable embryonic development period in the threatened Northern Leopard Frog (Lithobates pipiens) in the Cypress

Hills, Alberta, Canada: conservation and management implications. Canadian Field-Naturalist 128(1): 50—S6.
Understanding breeding phenology is critical for establishing monitoring strategies, comprehending population dynamics,
and developing conservation actions for at-risk species, such as the Northern Leopard Frog (Lithobates pipiens). The timing
of spawning and hatching in the Northern Leopard Frog may be highly variable depending on regional environmental condi-
tions, which can make establishing the timing of surveys difficult. In spring 2006, eggs were laid over 30 days (24 April to
23 May) and hatching occurred over 2 weeks (14-28 May) at three neighbouring ponds in Cypress Hills, Alberta, Canada.
Although spawning occurred over a month, all eggs hatched within a 2-week period, indicating variable embryo development
rate. Among 26 egg masses, eggs laid later in the season developed approximately four times faster than those laid earlier,
and Akaike information criterion-ranked models suggested that both Julian date and water temperature were important predictors
of embryo development rate: later spawning date and warmer water were associated with faster rates. Some egg masses survived
colder temperatures than previously reported for this species. Asynchronous breeding and variable development rates reveal
the need to conduct multiple surveys over the breeding season, even within a small geographic area, to document reliably the
presence of egg masses and identify breeding habitat. Identification of key breeding habitat is necessary to mitigate human-caused
disturbances of such regionally imperiled species.

Key Words: Northern Leopard Frog; Lithobates pipiens; amphibian; breeding; egg; conservation; embryo development: spawn-
ing period; phenology; Cypress Hills; Alberta

Introduction its range in the 1970s and 80s (Roberts 1981*; Leonard

Breeding phenology and the time between spawning et a/. 1999), perhaps due to such factors as disease,
and hatching may be highly variable within and among drought, competition by invasive species, and habitat
populations of amphibians (Thumm and Mahony 2002; _ loss and fragmentation (COSEWIC 2009*). As a result,
Ryan and Plague 2004) as a result of an assortment of _ the western boreal—prairie populations of Northern
exogenous and endogenous factors (Reading 1998; Leopard Frogs are designated of “special concern” by
Oseen and Wassersug 2002; Grant et al. 2009). This | the Committee on the Status of Endangered Wildlife in
variability can make timing of population surveys chal- Canada (COSEWIC 2009*) and “threatened” under
lenging. Alberta’s Wildlife Act (AESRD 2012*).

Industrial development continues to increase in Al- We investigated predictors of spawning time and
berta, particularly in association with oil and gas extrac- | embryo development rate of Northern Leopard Frogs,
tion, and may negatively affect wildlife unless proper _ such as temperature and time of year, as part of a larger
mitigation measures can be implemented. Permits for study of tadpole microhabitat selection and juvenile dis-
industrial exploration and extraction often stipulate a _ persal behaviour. Natural history observations of this
requirement for amphibian surveys before land devel- type may be useful for improving the probability of
opment if at-risk amphibians are predicted to inhabit the detection by providing information regarding the timing
area (Dr. David Prescott, Species at Risk Biologist, of the breeding period and the frequency of surveys
Alberta Environment and Sustainable Resource Devel- required to identify and protect important habitat.
opment, personal communication, 29 March 2012).

However, for these surveys to be effective, they must Study Area
be carried out at a time and in a manner appropriate for Our study was conducted in and near Cypress Hills
the species of interest. Interprovincial Park (49°39'N, 110°01'W) which strad-

Once widely distributed and abundant in North dles the border between Alberta and Saskatchewan and
America, the Northern Leopard Frog (Lithobates pip- is just north of the Sweetgrass Hills of Montana (Fig-
iens) disappeared from much of the western portion of ure 1). The Alberta portion of the Cypress Hills is char-

50

2014

Legend

@ Breeding Pond
—— Road

Cypress Hills Interprovincial Park

RANDALL ET AL.: BREEDING AND EMBRYONIC DEVELOPMENT IN NORTHERN LEOPARD FroGs 5]

FiGure |. Locations of four Northern Leopard Frog (Lithobates pipiens) breeding ponds surveyed in 2006 in and near Cypress
Hills Interprovincial Park, Alberta, Canada (49°39'N, 110°01'W).

acterized by grasslands and boreal forest. Elevation
ranges from 1370 m to 1465 m, and average annual
temperature is lower than that of the surrounding grass-
land plains (Greenlee 1981*). These environmental
conditions likely present challenges to the reproduc-
tive success of Northern Leopard Frogs.

In 2006, we drove along roads and flew over the
study area in a fixed-wing aircraft to identify potential
Northern Leopard Frog breeding ponds within a 25-km
radius of our main study pond (Pond 1, Figure 1). Al-
though other breeding ponds may have been present,
these four represented all known breeding ponds at the
time.

Pond | was 0.1 ha in surface area and was surround-
ed by Populus spp. woodlands on the east, west, and
south sides; the north side had a relatively steeper slope
of mixed grass prairie (Fraser 2007). Ponds 2 and 3 had
surface areas of 0.17 ha and 0.04 ha, respectively, and
were also surrounded by mixed grass prairie and aspen
woodland. Pond 4 had a surface area of 0.24 ha and
was surrounded by mixed forest, mainly white spruce
(Picea glauca) on the west and north sides and dead-
fall on the south and east sides.

Methods

Study organism . .
Adult Northern Leopard Frogs are medium-sized
frogs, 5-10 cm long from snout to vent (Hine et al.

1981*: Russell and Bauer 2000). Within a population,
breeding period ranges from a few days to a few weeks
(Wells 1977). In Alberta, spawning occurs over a short
interval between late April and early June (Russell and
Bauer 2000; Kendell 2002) at temperatures of 10—25°C,
although spawning may be prolonged if the temperature
drops below this range (Hine e¢ al. 1981*; Gilbert ez
al. 1994; Kendell 2002*). Females deposit 600-7000
eggs in a single egg mass, which they attach to sub-
merged vegetation. Preferred water bodies are ephemer-
al or fishless permanent ponds or slow-moving back-
waters of streams and rivers in shallow water (AESRD
2012*). The period from spawning to hatching may
last from 5 days to 3 weeks (Russell and Bauer 2000;
Werner et al. 2004) and metamorphosis typically occurs
between July and August in Alberta (Kendell 2002*).

Observation techniques

Beginning approximately 30 minutes after sunset,
we listened for breeding calls of adult male Northern
Leopard Frogs for up to 20 minutes (AESRD 2013*).
If calls were detected, we returned the following day to
confirm spawning.

We conducted call and shoreline surveys from 25
April to 8 June 2006 and searched the shoreline of each
pond at least once every 2 days for new egg masses to
determine the duration of the breeding season and the
embryo development period (Merrell 1977*; Dorcas et

Nn
i)

al. 2010; Paton and Harris 2010). Individual egg mass-
es were identified by differences in either their size or
shape or the stage of development of the embryos (Mer-
rell 1977*). Their location along the shoreline was
marked with a flag to prevent counting an egg mass
twice (Gilbert et al. 1994; Dougherty ef a/. 2005).
During each survey, egg masses were identified as
hatched or unhatched (egg mass intact). Hatching was
confirmed when newly hatched tadpoles were seen ag-
gregating around the egg mass, feeding on the remain-
ing jelly.

For each egg mass, we measured its depth below the
water surface, distance to adjacent masses, and distance
from the shoreline using a metre stick. During shoreline
surveys, we measured the temperature of the water
within 10 cm of each egg mass using a Hanna pH pen
thermometer. We considered the breeding season com-
plete once male calling had ceased and no new egg
masses had been observed for at least 2 weeks.

Statistical analysis

We ran a one-way ANOVA to determine whether
ponds differed with respect to water temperature when
egg masses were laid, followed by Tukey’s honest sig-
nificant difference (HSD) test. We assessed whether
residuals were normally distributed using a Shapiro—
Wilk goodness-of-fit test (W > 0.93). We used Julian
date (JD) in our models to account for seasonal vari-

150
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S SANQOSE WSN
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110

THE CANADIAN FIELD-NATURALIST

Vol. 128

ation of environmental factors, such as photoperiod.
To evaluate predictive factors related to embryo
development, we first performed an exploratory regres-
sion analysis of embryo development rate versus JD
and water temperature (WT). To further investigate the
relation of JD and WT to embryo development rate we
used restricted maximum likelihood mixed-model
regression (JMP, version 7, SAS Institute Inc., Cary,
North Carolina, USA). Pond was a random factor; all
other factors were fixed. We formulated a set of can-
didate models that all included pond alone, pond with
JD or WT, or pond with both JD and WT in an additive
model and with the interaction of JD and WT. Because
collinearity of predictor variables can yield unstable
parameter estimates and inflated standard errors (Quinn
and Keough 2002), we verified that JD and WT were
not highly correlated (77 < 0.6) before we included them
together in a model (Royston and Sauerbrei 2008). We
compared models using small-sample-size Akaike
information criterion (AIC_) to select the “best” model
given a candidate set of models and considered models
to have equivalent support if AAIC. was < 2 (Burnham
and Anderson 2004). We assessed the goodness-of-fit of
the global model using a Shapiro—Wilk test (W > 0.90).

Results
Spawning occurred over 30 days at ponds 1-3,
beginning on 24 April and ending 23 May (Figure 2).

May 29

May 22

May 15

May 08

May 01

Apr 24

Pond

euRE oD” : J eye are een ey Seat. : Ri:
FIGURE 2. Development period for Northern Leopard Frog (Lithobates pipiens) eggs at ponds 1, 2, and 3

in Cypress Hills

Interprovincial Park, Alberta (see Figure 1) in April and May 2006. Each column of black circles represents days

from spawning to hatching for one of the 26 egg m

asses observed. The shaded area represents the period during

which egg masses were present at all three ponds (10-17 May).

2014 RANDALL ETAL.: BREEDING AND EMBRYONIC DEVELOPMENT IN NORTHERN LEOPARD FROGS 53

Although calling and spawning were confirmed at pond
4, we were unable to monitor egg masses because
pollen blanketed more than half of the water surface,
affecting our ability to observe them. Hatching occurred
at the three remaining ponds over 2 weeks, beginning
14 May with the last eggs hatching on 28 May. We ob-
served a total of 26 egg masses: 14 in pond 1, 7 in
pond 2, and 5 in pond 3. Egg masses were laid within
2-10 cm of the water surface, typically grouped togeth-
er in shallow areas within 2—3 m of the shoreline. All
monitored egg masses survived to hatching, but we
were unable to monitor the proportion of eggs that
hatched successfully.

The onset of spawning differed by only 3 days be-
tween ponds | and 2, but occurred approximately 2
weeks later at pond 3. The WT near each egg mass on
the day of spawning was significantly higher at pond 3
(17.4°C + 1.9 [mean + standard error], = 0.40, F, ,. =
7.62, P = 0.0029) than at ponds | and 2, but did not
differ between ponds | and 2 (9.1°C + 1.6 and 9.1°C +

Apr 24 May 01
25 ‘ '

20

15

10

Embryonic development period (days)

115 120 125

May 08

1.1, respectively). At ponds | and 2, egg laying first
occurred on 24 and 27 April, respectively. Up to five
egg masses were laid on the same day in ponds | and 2,
and the spawning period lasted for almost a month.
Egg masses were detected simultaneously at all three
ponds during only | week, 10-17 May.

At the time eggs were laid WT ranged from 5.7°C to
25.8°C. At pond |, WT dipped as low as 3°C during
egg development (27 April). Time from spawning to
hatching ranged from 5 to 20 days and decreased with
JD at laying (Figure 3). Eggs deposited at the beginning
of the breeding season (24 April) took about four times
as long to hatch as the last eggs laid (23 May).

Although we found a strong, negative relation be-
tween embryonic development period and JD, the rela-
tion between that period and spawning temperature
was not as strong (Figure 4). The top model, with 93%
of the AIC, weight, was an additive model that in-
cluded WT and JD (Table 1). No other models were
<2 AAIC, of the top model.

May 15
'

140

130 135 145

Julian date

FIGURE 3. Relation between dev
(Lithobates pipiens) egg Masses
The numbers above eac

elopment period and Julian date (24 April-28 May 2006) for 26 Northern Leopard Frog
at ponds 1, 2, and 3 in Cypress Hills Interprovincial Park, Alberta (see Figure 1).
h symbol represent the number of egg masses laid on that date. Symbols for ponds at which

egg masses were laid on the same date have been slightly offset (~ = 0.91).

54 THE CANADIAN FIELD-NATURALIST Vol. 128

25

A Pond 1

CO Pond 2
@ Pond3

20

15

10

Embryonic development period (days)

8 10 12 14 16 18

Water temperature (°C)

FIGURE 4. Relation between development period (24 April—28 May 2006) and spawning temperature for 26 Northern Leop-
ard Frog (Lithobates pipiens) egg masses at ponds 1, 2, and 3 in Cypress Hills Interprovincial Park, Alberta (see
Figure 1). Numbers above each symbol represent the number of egg masses laid at that water temperature. Symbols
for ponds at which egg masses were laid at the same water temperature have been slightly offset (7? = 0.32).

TABLE 1. Variation in development period of Northern Leopard Frog (Lithobates pipiens) eggs with Julian date (JD) and water

temperature (WT). The top model was selected using Akaike information criterion adjusted for small sample size (AIC).
3.08 aVvYa—Naa@a@oQqoQqwo@maomamaSq_q_—_ aa —“—“_O—————————————————————————

Model —2LL K AIC, AAIC, AIC, weight
Embryo development period = Pond + JD + WT ODESY 5) 115.59 0.00 0.93
Embryo development period = Pond + JD 111.24 d 121.14 5:55 0.06
Embryo development period = Pond + JD +WT + JD* WT 107.59 6 124.01 8.42 0.01
Embryo development period = Pond + WT 158.07 4 167.97 52.38 0.00
Embryo development period = Pond 164.85 3 171-95 56.36 0.00

Note: -2LL is —-2*model log-likelihood, K is the number of parameters in the model. AAIC. is the difference between the
AIC, of each model and the top model.

Discussion Werner ef al. 2004). It is interesting to note that, where-
The onset of spawning was not synchronous in our —_as spawning occurred over the course of a month. all
study area and varied by over 3 weeks among our ponds. _ eggs hatched within a 2-week period with the first eggs
However, spawning was often synchronous within a deposited taking almost four times as long to develop as
pond, with several egg masses laid ona single day. The __ the last eggs laid.
spawning period was also protracted, lasting up to 30 Eggs were laid at WT ranging from 5.7°C to 25.8°C.
days at a single breeding pond (pond 1). The duration which is consistent with known egg temperature tol-
of the breeding season and the embryonic development __erances for this species (Moore 1939, 1949). However.
period was consistent with other published reports our minimum spawning temperature was more than
(Merrell 1968; Wells 1977; Russell and Bauer 2000; 2°C colder than that recorded for Northern Leopard

2014 RANDALL £7 4L.: BREEDING AND EMBRYONIC DEVELOPMENT IN NORTHERN LEOPARD FROGS

Erogs in Quebec, which do not spawn at WT below
8°C (Gilbert et al. 1994). In addition, WT at one of our
ponds dropped to 3°C during the development period,
which is 2°C below the reported threshold for normal
embryonic development in this species (Moore 1949).
However, the temperature at the centre of an egg mass
can be up to 2°C warmer than the surrounding water
(Hassinger 1970). Although the four affected egg mass-
es survived to hatching, we were unable to evaluate
whether the embryos had developed normally.

The top model for embryonic development period
included both JD and WT as predictive variables, sug-
gesting that temperature alone is not sufficient to
explain differences in development period, which might
also be affected by seasonal differences in the region.
However, there was no evidence that the effect of tem-
perature on development changed over time (the inter-
active model was > 8 AAIC, from the top model). Al-
though it has long been known that the development
of Northern Leopard Frogs is temperature dependant
(Atlas 1935: Moore 1939), clearly this is not the only
factor affecting embryo development period.

Conservation and Management Implications
Industrial development permits issued by regulators
often stipulate that developers determine whether at-
risk species, such as the Northern Leopard Frog, are
present and establish appropriate mitigation strategies
to reduce or eliminate negative impacts of their activ-
ities. Often only single surveys are conducted to deter-
mine the presence or absence of at-risk amphibians
(Kendell 2003*). Our results suggest that inappropri-
ately timed breeding surveys may fail to detect North-
ern Leopard Frogs and could, thus, limit the ability to
develop appropriate strategies to conserve this species.
Even within our small study area, egg masses would
have been observed in all three ponds during only |
week — a narrow timeframe for the completion of ef-
fective and comprehensive egg-mass surveys. Because
breeding phenology is likely to vary annually, region-
ally, and locally, we recommend that researchers adjust
the timing of their surveys so that they are relevant to
each specific site. To identify breeding ponds, multiple
breeding surveys separated in time should be conduct-
ed. The need to adjust the timing and number of sur-
veys necessary to identify breeding sites is not unique
to Northern Leopard Frogs (AESRD 2013*). As such,
we recommend that breeding phenology be considered
when developing monitoring strategies or industrial
mitigation procedures for amphibians elsewhere.

Acknowledgements . . . .
All research was conducted in compliance with Uni-

versity of Calgary Animal Care Protocol BI 2005-39
and the Calgary Zoo BRRC 2005-05 approval number.
We are grateful for funding provided by Husky Energy
and Cenovus Energy Ltd. We thank Typhenn Brichieri-
Colombi for providing the map of the study area.

Nn
Nn

Documents Cited (marked * in text)

AESRD (Alberta Environment and Sustainable Resource
Development). 2012. Alberta Northern Leopard Frog
recovery plan, 2010-2015. AESRD, Edmonton, Alberta.
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AESRD (Alberta Environment and Sustainable Resource
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lines. AESRD, Edmonton, Alberta. Page 128.

COSEWIC (Committee on the Status of Endangered
Wildlife in Canada). 2009. COSEWIC assessment and
update status report on the Northern Leopard Frog Litho-
bates pipiens, Rocky Mountain population, Western Bore-
al/Prairie populations, Eastern populations in Canada.
COSEWIC, Ottawa, Ontario. 69 pages.

Greenlee, G. M. 1981. Soil survey of Cypress Hills, Alberta,
and interpretation for recreational use. Alberta Research
Council, Edmonton, Alberta. 77 pages.

Hine, R. L., B. L. Les, and B. F. Hellmich. 1981. Leopard
Frog populations and mortality in Wisconsin. Technical
Bulletin 122. Wisconsin Department of Natural Resources,
Madison, Wisconsin. 39 pages.

Kendell, K. 2002. Survey protocol for the Northern Leopard
Frog. Alberta Species at Risk Report. Alberta Sustainable
Resource Development, Fish and Wildlife, Edmonton,
Alberta. 30 pages.

Kendell, K. 2003. Status of the Northern Leopard Frog (Rana
pipiens) in Alberta: update 2003. Alberta Sustainable Re-
source Development, Fish and Wildlife and Alberta Conser-
vation Association, Edmonton, Alberta. Page 61.

Merrell, D. J. 1977. Life history of the Leopard Frog, Rana
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Minneapolis, Minnesota. 22 pages.

Roberts, W. 1981. What happened to the leopard frogs?
Alberta Naturalist 11: 1-4.

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Grant, R. A., E. A. Chadwick, and T. Halliday. 2009. The
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opment in the eggs of amphibia. Ecology 20: 459-478.
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ters in Rana pipiens Schreber. Evolution 3: 1-24.

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Received 16 September 2013
Accepted 18 November 2013

Diet of the Pacific Sand Lance (Ammodytes hexapterus) in the Salish
Sea, British Columbia, in the 1960s

J. MARK HIPENER!3 and MorrA GALBRAITH?

cs bh Research Division, Environment Canada, R.R. 1, 5421 Robertson Road, Delta, British Columbia V4K 3N2 Canada
nstitute of Ocean Sciences, Ocean Sciences Division, 9860 West Saanich Road, Sidney, British Columbia V8L 4B2 Canada
*Corresponding author: mark.hipfner@ec.ge.ca

Hipfner, J. Mark, and Moira Galbraith. 2014. Diet of the Pacific Sand Lance (Ammodytes hexapterus) in the Salish Sea, British
Columbia, in the 1960s. Canadian Field-Naturalist 128(1): 57-62. '
The diet of the Pacific Sand Lance (Ammodytes hexapterus) was quantified from the stomach contents of 115 Pacific Sand
Lance caught in the Strait of Georgia and Saanich Inlet (Vancouver Island) in the Salish Sea, British Columbia, in the spring
and summer of 1966, in the Strait of Georgia in the spring and summer of 1967, and in the Strait of Georgia and Saanich Inlet
in the spring and summer of 1968. There were 12 major taxa of prey in diets, 8 of which were Crustacea. Based on an index of
relative importance, copepods were the dominant prey in 1966 and 1968, but not in 1967, when cladocerans, larvaceans, and
teleosts also were common. The copepods Pseudocalanus spp. and Calanus marshallae were the only taxa to appear in diets in
all three years. Pseudocalanus dominated the copepod component of diets in 1966, when sampling occurred in July; unspecified
copepod nauplii (an early larval stage) were dominant in 1967 and 1968, when sampling occurred earlier (April to June). With
the profound changes that have occurred in the Salish Sea over recent decades, these data can serve as a baseline for comparison.
Key Words: Pacific Sand Lance; Ammodytes hexapterus; diet; interannual variation; decadal variation; Strait of Georgia; Salish
Sea; British Columbia

Introduction 1967 in the Strait of Georgia, and 81 were collected be-
In marine systems of the temperate northern hemi- _ tween 23 April and 7 June 1968 in the Strait of Georgia
sphere, the six species of sand lances (Ammodytes spp.) _ and Saanich Inlet. The gastrointestinal tracts of the 115
often act as important trophic links between zooplank- _ Pacific Sand Lance were excised with scalpel and for-
ton and the broad suite of predators that feed on small __ceps and were placed in vials filled with 15% buffered
forage fishes (Wanless et al. 2005). The energy densi- formalin.
ties, growth rates, and fecundity of sand lances all vary In the laboratory, the excised tracts were washed
spatially and temporally (Robards ef al. 2002; Wanless — over a (0.063 mm mesh to remove the formalin, which
et al. 2005), and this suggests that feeding conditions —_ was captured for neutralization treatment and disposal.
for sand lances can be highly variable. Each stomach was then separated out, transferred to a
The Pacific Sand Lance (Ammodytes hexapterus) is 1 mm gridded Petrie dish, and slit longitudinally under
a vital component of northeastern Pacific Ocean food =a Wild M4 dissecting microscope with ocular microm-
webs (Beacham 1986; Borstad et al. 2011). Despite its __ eter. The contents were removed using water from a
ecological importance, however, information on the diet squeeze bottle and then the items were counted. Their
of this species is extremely limited. A recent study quan- life stage was determined and they were identified to
tified the diet of the Pacific Sand Lance at several sites _ the lowest taxonomic level possible. Some stomach
along British Columbia’s outer coast in the summers of contents were fresh and in a condition that allowed the
2009, 2010, and 2011 (Hipfner and Galbraith 2013). prey to be identified to species, but most contents had
Here. we have collated and analyzed data collected _ been at least partially digested. As a result, the repre-
in the protected inner waters of the Salish Sea in the _ sentation of soft-bodied prey types such as larvaceans
spring and summer of 1966, 1967, and 1968 (Barra- _ in diets could have been underestimated (Barraclough
clough and Fulton 1968a*, 1968b*; Barraclough et al. and Fulton 1968a*, 1968b*; Barraclough et al. 1968*:
1968*: Robinson et al. 1968*; Robinson 1969*). Be- Robinson ef al. 1968*; Robinson 1969*).

cause the Salish Sea ecosystem has undergone dramatic For this report, dry weight estimates were assigned
changes since that time (Therriault et al. 2009), these to each species and stage based on weight vs. length re-
data can serve as a baseline for comparison. gressions from Fulton (1968) and McCauley (1984), as

well as direct lab measurements. Total length was used
Methods for most prey items, the exceptions being fork length

Pacific Sand Lance were collected in surface trawls _ in the case of larval fish and prosome length in the case
in waters of the Salish Sea off the coast of British Col- of copepods.
umbia: 8 were collected between 4 and 8 July 1966 in To facilitate direct comparison with information col-
the Strait of Georgia and Saanich Inlet (southern Van- lected elsewhere in British Columbia, we analyzed the
couver Island), 26 were collected between 5 and9 June data as described in Hipfner and Galbraith (2013). An

a

58 THE CANADIAN FIELD-NATURALIST

index of relative importance (Pinkas e¢ al. 1971) was
calculated to quantify the importance of each prey type
in Pacific Sand Lance diets in each year. The formula is:

(1) /,, = the proportion of all non-empty stom-

achs that contained prey type (7) * (the propor-

tion by number of prey type (7) item relative to
all prey types + the proportion by dry mass of
prey type (/) relative to all prey types).

Following Santic et al. (2012), we then standard-
ized this index for each prey type to a scale of 0-100
using the formula:

Cao WOO en (ene):

When a predator consumes exclusively small prey,
there tend to be strong correlations among the three
component measures of /,,, (percentage frequency of
occurrence, percentage by number of individual prey
items, and percentage mass of prey), and these corre-
lations can exaggerate differences in the importance
of prey types (Macdonald and Green 1983). There-
fore, we also report each of the three component
measures for all prey types that constituted 10% or
more of the %/,, in at least one of the three years.

Results

The fork lengths of the collected Pacific Sand Lance
varied from 10 to 115 mm, with the mean fork length
being longer the later in the season that sampling oc-

125
100
»
=> 75
I
oD)
S
ao)
rer ag
—
oO
ie
95
0

4-8 Jul 1966

5-9 Jun 1967

Vol. 128

curred (Figure 1). Based on Figure 5 in Blackburn and
Anderson (1997*) and Figure 7 in Robards ef al.
(2002), we assume that all Pacific Sand Lance sampled
in 1966 and 1968 were young-of-year fish (age 0), as
were the vast majority in 1967 (those <100 mm in fork
length). Food was found in the stomachs of all but | of
the 115 Pacific Sand Lance, the lone exception occur-
ring in 1967.

Across the three years, there were 12 major taxa of
prey in Pacific Sand Lance stomachs, including 8 groups
of crustaceans, as well as trematodes, molluscs, larva-
ceans (plankton), and teleosts (fish). However, prey
from only 2 (in 1968), 4 (1966), and 10 (1967) of the
12 major taxa were present in any one year (Table 1).
Of the 12 major taxa, copepods were by far the domi-
nant group in 1966 (%/,, = 97.0) and 1968 (99.9), but
not in 1967 (21.5), when cladocerans (33.1), larvaceans
(18.0), and teleosts (25.7) also were common (Figure 2)
(Table 2).

The copepods Pseudocalanus spp. and Calanus paci-
ficus were the only prey types recorded in all three
years (Table 1). In considering just the copepod com-
ponent of the diet of the Pacific Sand Lance, Pseudo-
calanus dominated in 1966 (%/,, = 72.8), whereas
unspecified copepod nauplii dominated in both 1967
(Yol,, = 72.0) and 1968 (%/,, = 64.1) (Table 3) (Fig-
ure 3). Three large calanoids—Calanus pacificus,

23 Apr-7 Jun 1968

Dates of collection

FIGURE 1. Fork lengths of Pacific Sand Lance (Ammodytes hexapterus) collected in trawl surveys in the Salish Sea, British
Cc olumbia, in relation to the date of sampling. Box plots show means, 25th and 75th percentiles, Sth and 95th percentiles
(whiskers), and outliers. Sampling occurred from 4 to 8 July 1966 in the Strait of Georgia and Saanich Inlet, from 5
to 9 June 1967 in the Strait of Georgia, and from 23 April to 7 June 1968 in the Strait of Georgia and Saanich Inlet

2014

TABLE 1. Prey taxa and life sta
Salish Sea, British Columbia, in 1966-1968.

HIPFNER AND GALBRAITH: SAND LANCE DIET IN THE SALISH SEA 59

ges recorded in the stomach contents of Pacific Sand Lance (Ammodytes hexapterus) in the

SSSO0°50—OoOo<“Vea2=2>0— ee

Major taxa

Sub-type/species 1966 (n= 8) 1967 (n = 26) 1968 (n= 81)
Trematodes () Yes ()
Molluses Pelecypoda () Yes ()
Crustaceans
Cirripeds Barnacle 0 Yes (nauplii) 0
; Balanus 0 Yes (eggs) 0
Cladocerans Podon Yes Yes 0
Ostracods Alacia minor 0 Yes 0
Copepods Copepod 0 Yes (eggs, nauplii, Yes (eggs, nauplii,
copepodites) copepodites)
Acartia longiremis Yes 0 0)
Calanus pacificus Yes Yes Yes
Microcalanus 0 0 Yes
Pseudocalanus Yes Yes Yes
Cyclopoid 0 Yes 0
Neocalanus plumchrus 0 Wes 0
Scolecithricella minor Yes 0 0
Paraeuchaeta elongata 0 0 ives
Hyperiids Themisto pacifica 0 Yes 0
Mysids — Yes 0 0
Euphaustids Euphausiid 0 0 Yes (eggs)
Decapods Brachyuran Yes (zoea) Yes (megalops) 0
Larvaceans Oikopleura 0 Yes 0
Teleostei — 0 Yes 0

Neocalanus plumchrus, and Paraeuchaeta elongata—
were the dominant copepods in terms of mass in 1966,
1967, and 1968, respectively, despite being present
in small numbers and in few stomachs—especially P.
elongata (Figure 3) Although Paraeuchaeta elonga-
ta constituted 44.97% of the copepod component of

100 Gl 1966
=>) 1967,
90 a 1968
=
8 80
Cc
oO
6
Qa 40
i=
g
o 30
oO
==
15)
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o
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10
0 ” ” n n ” ” ny ” ” &
& 3 so] (= Be] ne] ne] Ao] ne] UD [= 7]
0 7) ® © 9 2 = a a 9 ee ers
fe) = Q 5 13) a o oi 3 a @
eo. 1S obec wom (ee oe Sata
ESS 6 ~~ 6 boo t& ie a eS
® oO ie} Oo = a is
E S) < ) By -
Crustaceans
Taxon

FIGURE 2. Indices of relative importance (%lp,) of the 12
major taxa of prey consumed by Pacific Sand Lance
(Ammodytes hexapterus) in the Salish Sea, British
Columbia, in each of three years (1966, 1967, and

1968).

diets in terms of mass in 1968, it was present in very
small numbers (2.50%) in very few stomachs (2.75%)
and thus had %/,, of <2%. Scolecithricella minor (in
1966) and unspecified copepodites (in 1968) were the
only other prey that constituted >10% of copepod %/,.,
in at least one of the three years (Table 3) (Figure 3).

100
80
60
40

20

Index of relative importance (%)

(0) =

wo x 2] 7) = no
= g g g 2 +S) s es} 2 [ow 3
& 3 = = = ¢ s 2 s 2) AT
= ee a ae a Gee ay
Se BO ee. ae hn
= Q ° fe) se Ms > a
KS} g tay ES) Q Oo & Oo ie)

= =

SY = 3s a er ry =
t © rif = Ss =
8 3 s o's
hal s 8 8

; eg

<= a

Copepods

FIGURE 3. Indices of relative importance (“%/,,) of the 11 taxa
and life stages of unspecified copepod prey consumed
by Pacific Sand Lance (Ammodytes hexapterus) in the
Salish Sea, British Columbia, in each of three years
(1966, 1967, and 1968).

128

Vol.

THE CANADIAN FIELD-NATURALIST

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Discussion

The Pacific Sand Lance is a vital component of
northeastern Pacific Ocean food webs (Beacham 1986:
Gjerdrum ef a/. 2003), so there is need for a thorough
understanding of its trophic relations. In the protected
inner waters of the Salish Sea, 12 major prey taxa were
found in its diet: 8 crustacean groups, plus trematodes,
molluscs, larvaceans, and teleosts. This suite of prey
taxa is similar to that reported for other species of sand
lances (Scott 1973; Sekiguchi 1977; Richards 1982)
and for the Pacific Sand Lance elsewhere (Blackburn
and Anderson 1997*), including at four sites along
British Columbia’s outer coast (Hipfner and Galbraith
2013). Further, with one exception, in which mysids
were the main prey type (O’Connell and Fives 1995),
calanoid copepods were the primary prey of sand lances
in all studies, so it was interesting that a more balanced
array of prey, also including cladocerans, larvaceans,
and teleosts, was taken in one of the three years of this
study (1967).

The diet of the Pacific Sand Lance along British
Columbia’s outer coast included a broader array of
copepod taxa and different life stages (Hipfner and
Galbraith 2013) than in the Salish Sea (this study).
Pseudocalanus (especially in the north) and Calanus
marshallae (especially in the south) were the primary
prey along the outer coast. Pseudocalanus was the pri-
mary prey in only one of the three years in the Salish
Sea, and C. marshallae was not found at all. Copepod
nauplii, the main prey of Pacific Sand Lance in two
of the three years in the Salish Sea, occurred in trace
amounts in just 2 of 12 site-years along the outer coast.

The differences between the two sets of studies could
reflect any or all of four factors: (1) four decades sepa-
rate them; (2) the age and size structure of the Pacific
Sand Lance differed, with the outer coast study includ-
ing only Pacific Sand Lance >100 mm fork length;
(3) the timing of sampling differed (April to early
July in the Salish Sea and July to early August along
the outer coast); and (4) the copepod communities of
the Salish Sea and the outer coast differ.

We cannot assess the role that changes in the marine
environment over time may have played, but there is
reason to believe that all three of the remaining fac-
tors were involved. First, sand lance diets vary with age
and size (O’Connell and Fives 1995); for example, the
amount of the larger C. marshallae consumed by Pacif-
ic Sand Lance increased with fork length, while the
amount of the smaller Pseudocalanus did not (Hipfn-
er and Galbraith 2013). Second, the fact that copepod
nauplii (an early larval stage) were very common in
diets in the Salish Sea in both years when sampling
occurred in spring but not in any year in the Salish Sea
or along the outer coast when sampling occurred in
summer implies that seasonal timing 1s a factor. And
lastly, the summertime copepod community of the Sal-
ish Sea is dominated by small copepods, including
Pseudocalanus (Sastri and Dower 2009), which was

HIPFNER AND GALBRAITH: SAND LANCE DIET IN THE SALISH SEA 6]

the primary prey of Pacific Sand Lance in July 1966,
whereas Pseudocalanus and Calanus marshallae, the
dominant prey along the outer coast in all three years
of study (2009, 2010, and 2011) (Hipfner and Galbraith
2013), are two of the three most abundant zooplankters
in continental shelf waters from Oregon to the Bering
Sea (Mackas and Coyle 2005).

Acknowledgements

We thank W. Barraclough, J. Fulton, and D. Robin-
son, all of the Fisheries Research Board of Canada, for
conducting the trawl surveys (which were also funded
by the Fisheries Research Board of Canada) in the
1960s, and we thank the students from the South
Hampton Work Experience Program for transcribing
their data. Thanks also to T. Gaston for commenting
on a previous draft of the paper.

Documents Cited (marked * in text)

Barraclough, W. E., and J. D. Fulton. 1968a. Number, size
composition and food of larval and juvenile fish caught
with a two-boat surface trawl in the Strait of Georgia July
4-8, 1966. Fisheries Research Board of Canada Manuscript
Report Series No. 940.

Barraclough, W. E., and J. D. Fulton. 1968b. Food of larval
and juvenile fish caught with a surface trawl in Saanich
Inlet during June and July 1966. Fisheries Research Board
of Canada Manuscript Report Series No. 1003.

Barraclough, W. E., D. G. Robinson, and J. D. Fulton.
1968. Number, size composition, weight, and food of lar-
val and juvenile fish caught with a two-boat surface traw]
in Saanich Inlet April 23—July 21, 1968. Fisheries Research
Board of Canada Manuscript Report Series No. 1004.

Blackburn, J. E., and P. J. Anderson. 1997. Pacific Sand
Lance growth, seasonal availability, movements, catch vari-
ability, and food in the Kodiak-Cook Inlet area of Alaska.
University of Alaska Sea Grant College Program Report
No. 97-01.

Robinson, D. G. 1969. Number, size composition, weight and
food of larval and juvenile fish caught with a two-boat sur-
face trawl in the Strait of Georgia April 24-25, 1968. Fish-
eries Research Board of Canada Manuscript Report Series
No. 1067.

Robinson, D. G., W. E. Barraclough, and J. D. Fulton.
1968. Number, size composition, weight and food of lar-
val and juvenile fish caught with a two-boat surface trawl
in the Strait of Georgia June 5-9, 1967. Fisheries Research
Board of Canada Manuscript Report Series No. 972.

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Received 24 September 2013
Accepted 8 November 2013

Melanistic Diversity in the Maritime Gartersnake, Thamnophis sirtalis

pallidulus, in Nova Scotia, Canada

JOHN GILHEN!:2 and Frep W. Scott!

Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada

“Corresponding author: gilhenja@gov.ns.ca

Gilhen, John, and Fred W. Scott. 2014. Melanistic diversity in the Maritime Gartersnake, Thamnophis sirtalis pallidulus, in Nova
Scotia, Canada. Canadian Field-Naturalist 128(1): 63-71.

We describe an unusual diversity of melanistic conditions in populations of the Maritime Gartersnake, Thamnophis sirtalis
pallidulus, in Nova Scotia: pure melanism, melanism, nigrismus (predominantly melanistic), abundismus (mosaic melanism).
This variability is widespread in Nova Scotia, and it includes coastal islands.

Key Words: Maritime Gartersnake; Thamnophis sirtalis pallidulus; pure melanism; melanism; nigrismus; predominantly melanistic;
abundismus; mosaic melanism; coastal islands; Nova Scotia

Introduction

Rowell (2012) explained that melanism is a Mendel-
ian recessive trait that is rare or absent in most Common
Gartersnake (Thamnophis sirtalis) populations. Peters
(1964) and Klauber (1972) provide definitions of mela-
nism, nigrismus, and abundismus. Melanism is an un-
usual increase in the normal amount of black pigment
within a particular individual as contrasted with other
members of his species. Occasionally used to character-
ize a species in which all individuals are equally black.
Nigrismus is a type of melanism in which the specimen
is not completely black and shows a change in the ele-
ments of the basic pattern as a consequence of an in-
crease in the size of the black portion of the pattern, mak-
ing the specimen predominantly black. Abundismus is
a mosaic type of melanism in which the specimen is not
completely black but shows a change in the elements of
the basic pattern as a consequence of an increase in the
number of black spots or blotches in the non-black areas
of the pattern.

We propose a fourth type of melanism: pure mela-
nism. The pure melanistic condition is uniform black on
the back and does not show a spotted pattern when the
skin is spread. This is the main character which separates
the pure melanistic condition from the melanistic con-
dition.

All four melanistic conditions are variable in the
Maritime Gartersnake, Thamnophis sirtalis pallidulus,
and are widespread in specimens found on the main-
land, on Cape Breton Island, and on two coastal islands
in Nova Scotia (Figure 1). In general terms, these mel-
anistic conditions in the Maritime Gartersnake in Nova
Scotia can be defined as follows.

Pure melanism

Individuals are a uniform satiny black to bluish-black
on the back (Figure 2), rarely with purplish gleam. The
back will be lustrous in some lights when the snake is

63

freshly shed. The top of the head, the underside of the
trunk, and the tail are glossy black to bluish-black, (like
the melanistic condition, see the photograph on the coy-
er). The infralabials, chin shields, and gular scales, are
enamel white, rarely bright pinkish-white. The white
shades change abruptly to black or bluish-black on the
first few anterior ventral scales, and the black or bluish-
black extends to the tip of the tail. The skin between the
scales is dull dark grey to bluish-grey throughout. Only
4 specimens of the Maritime Gartersnake are known to
manifest the pure melanistic condition (Table 1). They
do not show a Maritime Gartersnake spotted pattern
when the skin is spread.

Melanism

Individuals are usually satiny black to bluish—black
on the back (Figure 3 before shedding and the same
snake, see cover, two days after shedding), rarely with
a purplish gleam. The back is lustrous in some lights,
particularly when the snake is freshly shed. The top of
the head and the underside of the trunk and tail are glossy
black to bluish-black. The infralabials, chin shields, and
gular scales can be enamel white, bluish-white, yellow-
ish-cream, light orange (on snakes from Georges Island,
Halifax County), and rarely pinkish-white. In addition,
some individuals have black spots or streaks on the side
of the jaw, the chin, and the throat. These light colours
change abruptly to the darker ventral colour on the first
few anterior ventral scales. The skin between the scales
can be bluish-white, light grey, or light yellowish-brown.
When the skin is spread, large ellipsoid-shaped spots or
smaller alternating spots, and dorsal stripe, if present,
(characteristic of the Maritime Gartersnake spotted pat-
tern), are revealed. This pattern, when the skin is spread,
is the most distinguishing character which separates it
from the pure melanistic morph.

Counting the 3 young from Wellington, there are 27
records of the melanistic condition (Table 2) in Nova

64 THE CANADIAN FIELD-NATURALIST

New 2
Brunswick

Prince Edward Island

Vol. 128

Pure Melanistic

Melanistic

Negrismus

Abundismus

More than one condition

FiGurE 1. Distribution of the four melanistic conditions in the Maritime Gartersnake, Thamnophis sirtalis pallidulus, in

Nova Scotia.

TABLE |. Four pure melanistic Maritime Gartersnakes, (Thamnophis sirtalis pallidulus) from Nova Scotia, Canada, in

chronological order.

Age Sex Locality

Adult Female | Canso, Guysborough County

Adult Female Lake Rossignal,Queens County

Adult Female — Londonderry,Colchester County

Adult Female _ Big Tancook Island, Lunenburg County 18 August 2013

Date collected Remarks

5 October 1905 Nova Scotia Museum, Piers no. 2937

21 October 1970 Nova Scotia Museum970-Z-400-1
(Figure 2)

Sight record

Anomaly. Brown labials, black ventral
scales, and grey subcaudals (Figure 7)

24 May 1980

Sao 0» 0—=o®oqlewloeleloeleo—s

Scotia; 11 of these reported observations were made
from a distance and do not include a description of
the underside of the body or tail.

Nigrismus or predominantly black

This is a type of melanism in which the individual
looks uniform black from a distance and up close it is
predominantly black. The spotted pattern, and dorsal
stripe, when present, of a Maritime Gartersnake are
black and the remaining ground colour is a mosaic of
blackish clove-brown with scattered white scales
(Gilhen 2000) (Figure 4). This morph is dark steel grey

“plumbeous” to grey-black on the underside of the
trunk and the tail (Figure 5). The underside of the tail
of one individual was blackish clove-brown. The infra-
labials, chin shields, and gular scales may be bluish-
white, or yellowish-cream to orange-cream, and may be
freckled with grey. There are only four observations of
the nigrismus or predominantly black condition in Nova
Scotia (Table 3).
Abundismus or mosaic

In these Individuals, the normal spotted pattern is
invaded by dark grey-black to black spots and/or blotch-

2? 7 ] ] UF a . / .
2014 GILHEN AND Scott: MELANISTIC DIVERSITY IN THE MARITIME GARTERSNAKE. IN NOVA SCOTIA 65

FIGURE 2. Pure melanistic Maritime Gartersnake, Thamnophis sirtalis pallidulus, from Lake Rossignol, Queens County, Nova
Scotia, captured on 21 October 1970, by James Harding. Photo: R. Merrick.

J

=
a,
<a

i J. i
he < ee AS

eS ae | ’ \4

ype? 3

3. Melanistic Maritime Gartersnake Thamnophis sirtalis pallidulus, trom Big lancook Island, Mahone Bay, Lunenburg
Si, elanis Mle c c ; £

FIGURE - ;
County, Nova Scotia, captured on 7 May 2012 (before it shed its skin). Photo: H. Dione

: T 9)
66 THE CANADIAN FIELD-NATURALIST Vol. 128

TABLE 2. Twenty- seven melanistic Maritime Gartersnakes (Thamnophis sirtalis pallidulus) from Nova Scotia, Canada

chronological order.

De eee... en

Age Sex Locality Date collected Remarks

Unknown Unknown MeNabs Island, Halifax County 13 May 1929 Nova Scotia Museum no. 6519. “from its
colour it would be mistaken for a Black
Snake (Coluber constrictor)”

Adult Female Fletcher ’s Lake, Halifax County 15 July 1962 Sight record

Adult Unknown Jacques Landing, Queens County 16 August 1971 National Museum of Canada 13674. Park
highway road kill.

Adult Female George ’s Island, Halifax County 4 July 1991 Sight record

Adult Female George ’s Island, Halifax County 23 July 1992 Sight record no. |

Adult Female George ’s Island, Halifax County 23 July 1992 Sight record no. 2. Dark brownish-black
individual

Adult Male George ’s Island, Halifax County 10 August 1992 Anomaly. Individual had Yellow dorsal stripe
anteriorly (Figure 8)

3 Young Unknown Wellington, Yarmouth County 27 June 1995 F. R. Cook field numbers 18758, 18762 and
stillborn 18771

Unknown Unknown Mull River, Inverness County 16 May 1999 Nova Scotia Herpetological Atlas no. 3679

Unknown Unknown Upper Margaree, Inverness County 10 June 1999 Nova Scotia Herpetological Atlas no. 10072

Unknown Unknown Margaree Forks, Inverness County 10 June 1999 Nova Scotia Herpetological Atlas no. 11900

Unknown Unknown Margaree Forks, Inverness County 11 June 1999 Nova Scotia Herpetological Atlas no. 10073

Unknown Unknown Clark Lake, Colchester County 25 July 2000 Nova Scotia Herpetological Atlas no. 4352

Unknown Unknown Ponds, Pictou County 15 August 2000 Nova Scotia Herpetological Atlas no. 4883

Adult Female Dilligent River, Cumberland County 3 July 2002 Nova Scotia Herpetological Atlas no. 9196

Unknown Unknown Timberlea, Halifax County August 2002 Remains of house cat kill

Unknown Unknown Sampsonville, Richmond County 12 October 2002 Nova Scotia Herpetological Atlas no. 10000

Adult Female — Fairmont Road, Antigonish County 9 June 2003 Nova Scotia Herpetological Atlas no. 10451

Adult Female Grafton Lake, Queens County 10 July 2003 Sight record. Individual had bluish stripes

Unknown Unknown Gamble Lake, Colchester County 6 July 2004 Sight record

Adult Female —_ Dilligent River, Cumberland County 12 August 2004 = Nova Scotia Museum catalogue no. 55514

Adult Female Argyle Sound, Yarmouth County Summer 2004 Nova Scotia Museum catalogue no. 55593

Adult Female Garden of Eden, Pictou County 4 July 2008 Sight record. Individual near shedding,
blue eyes

Adult Female Big Tancook Island, Lunenburg County 7 May 2012 Individual has bluish stripes

Adult Female Lake Doucette, Digby County 30 September 2012 Nova Scotia Museum catalogue no. 55587

FIGURE 4. Dorsal view of nigrismus Maritime Gartersnake, Thamnophis sirtalis pallidulus, ftom McCabe Lake. Halifax County
Nova Scotia, captured on 24 June 1969. See also Figure 5. Photo: J. Gilhen. 7

20) : a
2014 GILHEN AND Scott: MELANISTIC Drv RSITY IN THE MARITIME GARTERSNAKE, IN NOVA SCOTIA — 67

FiGuRE 5. Ventral view of the nigrismus Maritime Gartersnake, Thamnophis sirtalis pallidulus, shown in Figure 4. Photo: J.
Gilhen.

es (Figure 6). The spots may be small, and involve only __itime Gartersnake in Nova Scotia. There are 9 records
a few scales on one side, may be large and invasive of _ of the abundismus condition (Table 4). Abundismus
the dorsal spotted pattern and stripe, or may involve _ individuals are often referred to in Nova Scotia as part-
just the underside of the trunk and tail. The remain- _ melanistic.

ing ground colour or pattern is as variable as any Mar-

TABLE 3. Four nigrismus Maritime Gartersnakes (Thamnophis sirtalis pallidulus) from Nova Scotia, Canada, in chronologi-

cal order.

ee
Age Sex Locality Date collected Remarks

Adult Unknown Paddy Lonis Brook, Halifax County 29 April 1918 = Nova Scotia Museum catalogue no. 4604
Adult Female Withrod Lake, Halifax County 6 May 1934 Nova Scotia Museum catalogue no. 7783
Unknown Unknown Mill Brook, Colchester County 7 August 1935 Nova Scotia Museum Catalogue no. 8224
Adult Female McCabe Lake, Halifax County 24 June 1969 Canadian Museum of Nature Amphibians

and Reptiles no.12153 (Figures 4 and 5)

ees oouowom-ees>s><c—0— Os

TABLE 4. Nine abundismus Maritime Gartersnakes (Thamnophis sirtalis pallidulus) from Nova Scotia, Canada, in chrono-

logical order.

SSS Oa e—mom=—a—asaausr mer’rs OTA
Remarks

Age Sex Locality Date collected

Adult Male Tomahawk Lake, Halifax County 5 July 1970 Site record

Unknown Unknown _ Big Pond, Cape Breton County 17 August 1977 National Museums of Canada
NMCAR25368

Adult Male George ’s Island, Halifax County 26 May 1989 Sight record no. | (see Figure 6)

Adult Male Georges Island, Halifax County 26 May 1989 Sight record no. 2

Adult Male George’s Island, Halifax County 14 April 1992 Sight record

Adult Male George’s Island, Halifax County 11 May 1992 Sight record

Adult Male South Ingonish beach, Victoria County 5 September 1996 Nova Scotia Museum 55056

Adult Male Big Tancook Island, Lunenburg County 24 May 2003 : Sight record vies

Adult Male Mack Lake, Kings County 23 August 2003 Nova Scotia Museum 55592

SSS ren

68 THE CANADIAN FIELD-NATURALIST

Vol. 128

cas

FIGURE 6. Abundismus Maritime Gartersnake, Thamnophis sirtalis pallidulus, from Georges Island, Halifax Harbour, Halifax
County, Nova Scotia, captured on 26 May 1989. Photo: J. Gilhen.

Discussion

For almost a century and a half, there have been pub-
lished and oral reports of black snakes from Nova Sco-
tia, New Brunswick, and Prince Edward Island (Bleak-
ney 1958). John Matthew Jones (1865) published the
first list of amphibians and reptiles of Nova Scotia and
he included the Black Snake, Coluber constrictor (i.e.,
North American Racer). Gilpin (1875) also gives an
account of the Black Snake. These reports of the Black
Snake, C. constrictor, are believed to be in error and
actually can be attributed to the melanistic morph of the
Maritime Gartersnake, as indicated by Harry Piers
(Gilhen 2000) and mentioned by Bleakney (1958),
Cook (1967), and Gilhen (1984). The North American
Racer and North American gartersnakes (Thamnophis
spp.) are easily distinguished from one another by the
scales on their back. The North American Racer has
smooth dorsal scales and North American gartersnakes
have strongly keeled scales.

The 4 melanistic conditions described above are un-
common in Nova Scotia but widespread. The ratio of
melanistic individuals to four other recognized col-
our patterns in wild populations in Nova Scotia is not
known. Only Georges Island, Halifax County, was sur-

veyed specifically for Maritime Gartersnakes (Barnes
et al. 2006). Between May and October 1993, a total of
391 Maritime Gartersnakes were marked on this 5-ha
drumlin in Halifax Harbour. Barnes ef al. (2006) state,
“Rarely were individuals melanistic (solid black) or
partially melanistic (five individuals).”

Over 100 Maritime Gartersnakes have been observed
on Big Tancook Island by JG but only 3 individuals
— one pure melanistic anomaly, one melanistic, and
one abundismus were observed and photographed. We
have kept 6 melanistic females from various localities
until they gave birth, but they did not produce mela-
nistic young. We have observed more than 20 non-mel-
anistic females from various localities in Nova Scotia
give birth, and none of their young were melanistic.
Also, we have not found melanistic young in the field.
However, Francis R. Cook collected two non-melanistic
pregnant females from Wellington, Yarmouth C ounty,
one on 27 June 1995 and the second on 2 July 1995.
Both females gave birth in captivity. The female from
27 June, gave birth to 16 young (3 melanistic and 13
non-melanistic). The female from 2 July gave birth to
13 normal young. During annual visits from 1963 to
1995 to the former Frank Crosby farm at Wellington,

012 al : ‘ - = — etre . \ \
2014 GILHEN AND Scott: MELANISTIC DIVERSITY IN THE MARITIME GARTERSNAKE, IN NOVA SCOTIA 69

Francis R. Cook and Joyce Cook never saw a melanis-
tic Maritime Gartersnake. Also, Joyce Cook lived there
from 1948 to 1958 and did not see a melanistic snake,
nor were any noted by her parents or two brothers.
Melanism is more common in the Eastern Garter-
snake, Thamnophis sirtalis sirtalis. Rowell (2012)

provides a table (Table 24.1) listing the incidence of

melanism reported at locations in southern Ontario, par-
ticularly on islands of western Lake Erie and at nearby
mainland locations. The incidence varied from 6.4%
at Point Pelee to 12%—59% at Point Pelee Island, East
Sister Island, and Middle Island, and 24%-—S51.2 % at
Long Point.

Pure melanistic

The first recorded observation of pure melanism in
a Maritime Gartersnake is by Harry Piers (Accession
Book No. 1, Nova Scotia Museum no. 2937). He stat-
ed, “Melanistic variety of Common Garter Snake, All
Saints Rectory grounds, Canso, Guysborough County,
by Rev. R. M. Leigh’s man on 5 October 1905”. Under
Remarks he stated “Colour. — Back black, with no sign
whatever, in any light, of blotches or stripes, etc.; under-
side of head white; belly slate-coloured, the under plates

(belly & tail) narrowly margined with lighter. Ventral
plates with a black spot on outer margin, near begin-
ning of dorsal scales, thus being almost concealed by
the next ventral plate.”

A large female from Londonderry was most unusual
in that it was black with a distinct purplish lustre. The
infralabial scales, chin shields, gular scales, and first
few ventral scales were glossy pinkish-white and the
remaining underside was grey-black with a pinkish
lustre. The large female from Big Tancook Island was
different and unusual in that the supra-labials and neck
were brown fading to yellowish-cream on the infra-
labials, chin shields, gular scales, and first few ventral
scales. This snake had a light lateral stripe on both sides
anteriorly. The belly was black nearly to the anal plate
and then changed abruptly to grey to the tip of the tail
(Figure 7). It resembled mostly the pure melanistic con-
dition. When the skin was spread, a Maritime Garter-
snake spotted pattern was not revealed.

Melanistic

The first recorded observation of melanism in a Mar-
itime Gartersnake is by Harry Piers (Accession Book
No. 3, Nova Scotia Museum no. 6519). He recorded a

F re 7.Adult male melanistic Maritime Gartersnake, Thamnophis sirtalis pallidulus, anomaly with light yellow anterior dorsal
: : : J. Gilhe
bas stripe from Georges Island, Halifax County, Nova Scotia, captured on 10 August 1992. Photo: J. Gilhen.

70 THE CANADIAN FIELD-NATURALIST

“Striped Garter Snake ABNORMAL MELANISTIC
INDIVIDUAL!!” from McNab’s Island, Halifax Har-
bour, Halifax County, captured on 13 May 1929 by
Joseph Perrin. “From its colour it would be mistaken
for a Black Snake (Coluber constrictor)” (i.e., North
American Racer). This may be the same variation of
melanism exhibited by a female from Grafton Lake,
Queens County, on 10 July 2003 and a female from Big
Tancook Island, Lunenburg County, on 3 October 2011.
Both females had blue-black dorsal and lateral stripes.

Of the 27 Maritime Gartersnakes reported here as
melanistic, 9 were reported in the Nova Scotia Herpeto-
logical Atlas (NSHA). Since a description and/or image
was not given, the Nova Scotia Herpetological Atlas
number for each one is provided in Table 2.

The underside of the snake from Garden of Eden was
not recorded and its blue eyes indicate it was about to
shed. Two rare variations and one anomaly were found
in the melanistic condition on Georges Island. One indi-
vidual was dark brownish-black with orange at the cor-
ner of the mouth fading to yellowish-white on the chin
and extending onto the first few ventral scales. Another
snake from Georges Island was similar to a large pure
melanistic female from Londonderry in that it was black
with a distinct purplish gleam (Barnes ef al. 2006.). The
chin and first few ventral scales were glossy pinkish-
white and the remaining underside was dark pinkish-
grey. One snake from Timberlea, Halifax County, was

Vol. 128

dark blueberry-blue on the back. The underside of the
trunk was bluish-black. The sides of the head were blue
fading to bluish-white on the chin shields. One adult
male anomaly had a light yellow dorsal stripe anteriorly
(Figure 8).
Nigrismus

The first recorded observation of nigrismus in a
Maritime Gartersnake is also by Harry Piers (Accession
Book No. 2, Nova Scotia Museum no. 4604). He re-
corded a striped garter snake, with melanistic tendency
from Paddy Lonis Brook, one mile SSE of Enfield, Hal-
ifax County, on 29 April 1918. His detailed account of
the colour is as follows: “Colour from fresh specimen.
General colour very dark; stripes moderately plain (on
anterior parts; spots very obscure owing to dark colour
of back. Top of head and back blackish, Clove-Brown.
Upper labials and lower part of snout plate (rostral)
Buff. Iris, brown. Dorsal stripe (on dorsal row of scales
and part of row on each side of it), dirty white; plainest
for about 5 inches back of snout, and becoming lost
about 12 % inches behind snout (that is, a little more
than half-way of length). The stripe is broken for a cou-
ple of inches behind the head by the connection across
it of 5 of the dark spots on each side of it. Lateral stripe
on parts of 2" and 3" rows of scales pale olive-buff,
plainest for about 5 inches behind snout, not so evident
or so light as the dorsal stripe, and becoming lost about

FIGURE 8. Pure melanistic Maritime Gartesnake, Thamnophis sirtalios pallidulus, anomaly with yellowish-brown supralabials
fading to yellowish-white on underside of head and with blackish ventral scales and greyish subcaudal scales, from Big
Tancook Island, Lunenburg County, Nova Scotia, captured on 18 August 2013. Photo: R. Lloyd. ;

2014 GILHEN AND Scott: MELANISTIC DIVERSITY IN THE MARITIME GARTERSNAKE, IN NOVA SCOTIA — 71

anus (about 17 inches from snout). Two rows of obscure
black spots, the inner row about .10 inch diameter each
spot, the outer row about .15 inch diameter each spot,
arranged alternately (zig-zag fashion) on each side of
dorsal stripe (that is between dorsal stripe and lateral
stripe); these spots somewhat noticeable for 4 or 5 inch-
es behind snout, but soon become lost in the general
blackish colour of upper parts. Underside of head,
bluish-white. Belly and underside of tail plumbous, the
abdominal plates each with a black spot on the front
lateral part. “*

Abundismus

Abundismus or the mosaic condition is extremely
variable. All 9 individuals recorded differed in the
amount of black pigment. The spots can be small and
involve only a few scales, large patches, or extensive
grey-black areas, particularly on the posterior half of
the body and tail, and the spots or areas invade or cover
the normal spotted pattern and stripes (Figure 6).

Acknowledgements

We thank Francis R. Cook, Curator Emeritus and
Research Associate, Canadian Museum of Nature, for
his encouragement, and for prodding JG over many
years to complete the manuscript. Dr. Cook and Andrew
Hebda, Curator of Zoology, Nova Scotia Museum of
Natural History, pre-reviewed and made useful com-
ments on the manuscript. We are grateful to the follow-
ing persons who either collected or provided reports of
Nova Scotia Maritime Gartersnakes in a melanistic
condition: David Baker, Joseph Baker, Suzanne Barnes,
Francis R. Cook, James Callaghan, Gary Corbett, Stan
Corbett, Albert d’Entremont, Hillary Dione, Gert Ewelt
(Nova Scotia Herpetological Atlas no. 4883), Calvin
Fraser, Karen Gilhen, Michael Gilhen, David Gordon,
May Goring, Ross Hall (Nova Scotia Herpetological
Atlas no. 4352), Anne-Belle Hamilton, James Harding,
David E. Harris, Peter Hope, Andrew Kennedy, Randy
Lauff, Earl Luffman, George MacInnis, Norman Mal-
oney, Mike McCabe, Ronald Merrick, John Mills (Nova
Scotia Herpetological Atlas nos. 10072, 10073, and
11900), Dana Pierce, Russell Rogerson, and Wayne F.
Weller. We thank Jeffrey Rowell for permission to cite
his privately printed book on the snakes of Ontario.

Roger Lloyd, photographer, Nova Scotia Museum,
developed digital images of all melanistic Maritime
Gartersnakes from original colour transparencies and
pictures taken by Ronald Merrick, JG, and Hillary
Dione. The cover image is by Roger Lloyd. We thank
naturalists Graham Caswell, Mary MacDonald, and
Heather McKinnon for the professional care of the live
Maritime Gartersnake display in the Netukulimk Gal-
lery, giving us ample opportunity to study and photo-
graph Maritime Gartersnakes in melanistic conditions.
Kim Franklin, administrative assistant, and Roger Lloyd
assisted in the development of Tables |—4.

Literature Cited

Barnes, S. M., C. M. Dubesky, and T. B. Herman. 2006.
Ecology and morphology of Thamnophis sirtalis pallidulus
(Maritime Garter Snake) on Georges Island, Nova Scotia.
Northeastern Naturalist 13(1): 73-82.

Bleakney, J. S. 1958. A zoogeographical study of the amphib-
ians and reptiles of eastern Canada. National Museum of
Canada. Ottawa, Ontario. Bulletin 155: 1-119.

Cook, F. R. 1967. An analysis of the herpetofauna of Prince
Edward Island. National Museum of Canada. Ottawa, On-
tario. Bulletin 212. Biological Series Number 75.

Gilhen, J. 1984. Amphibians and reptiles of Nova Scotia.
Nova Scotia Museum. Halifax. 162 pages.

Gilhen, J. 2000. Amphibians and reptiles of Nova Scotia:
Species recorded in the accession books of Harry Piers
1899-1939. Unpublished report. Nova Scotia Museum,
Halifax, Nova Scotia. 162 pages.

Gilpin, J. B. 1875. On the serpents of Nova Scotia. Trans-
actions of the Nova Scotia Institute of Science 4: 80-88.

Jones, J. M. 1865. Contributions to the natural history of
Nova Scotia: Reptilia. Transactions of the Nova Scotia
Institute of Science I(iii): 114-128.

Klauber, L. M. 1972. Rattlesnakes: Their Habits, Life Histo-
ries, and Influence on Mankind. Second edition. Volume 1.
Published for the Zoological Society of San Diego by the
University of California Press, Berkeley and Los Angeles,
California.

Peters, J. A. 1964. Dictionary of Herpetology. Hafner Pub-
lishing Company, New York, New York.

Rowell, J. C. 2012. The Snakes of Ontario: Natural History,
Distribution and Status. Privately Printed. 411 + vi pages.

Received 27 September 2013
Accepted 18 November 2013

Notes
Thickness of Common Murre (Uria aalge) Eggshells in Atlantic Canada

DONALD W. Pirte-Hay and ALEXANDER L. Bonp!

Department of Biology, University of Saskatchewan, and Environment Canada, 11 Innovation Boulevard, Saskatoon,
Saskatchewan S7N 3H5 Canada
‘Corresponding author: alex.bond@usask.ca

Pirie-Hay, Donald W., and Alexander L. Bond. 2014. Thickness of Common Murre (Uria aalge) eggshells in Atlantic Cana-
da. Canadian Field-Naturalist 128(1): 72-76.

Reported values for eggshell thickness in Common Murre (Uria aalge) are few, and even fewer since the decline in use of
organochlorine pesticides and other environmental pollutants that caused significant thinning of shells. The eggshells of Common
Murres and Thick-billed Murres (Uria lomvia) are among the thickest and heaviest, proportionately, of any bird and this represents
a non-trivial maternal investment. We measured the length and breadth of Common Murre eggs collected from Machias Seal
Island, New Brunswick, in 2006, and Gull Island, Newfoundland and Labrador, in 2012, and we measured the thickness of the
eggshells. Shell thickness was not related to egg size or volume, and it varied in individual eggs. The shells of Common Murre
eggs from Machias Seal Island (mean and standard deviation [SD] (0.767, SD 0.078 mm) and Gull Island (0.753, SD 0.057 mm)
were significantly thicker than any previously reported value and among the thickest of all birds. Such thickness is likely a result
of nesting on rock substrate with no nesting material and, perhaps, high breeding densities.

Key Words: Common Murre; Uria aalge; eggshell thickness; egg size; Atlantic Canada; Bay of Fundy; Gull Island; Machias
Seal Island; Witless Bay; New Brunswick; Newfoundland and Labrador

Introduction The purpose of our study was to determine eggshell
The integrity of avian eggshells, which are composed __ thickness and its relation to egg size of Common Murres
mainly of calcium carbonate, is critical for successful in contemporary samples from two sites in Atlantic
breeding (Gill 2007). Eggshell thickness depends on Canada and to compare eggshell thickness with previ-
the dietary intake of calcium of the egg-laying female _ ously published values. We also investigated the rela-
(Uspenski 1958); thus, the need to produce thick shells _ tion between eggshell thickness and other physical char-
creates high calcium demand. Thickness is also affected —_acteristics (colour and speckling pattern).
by organochlorine pesticides, introduced in the 1940s.
These compounds caused widespread eggshell thinning Methods
in birds (Bitman ef a/. 1969; Ratcliffe 1970; Lundholm We collected Common Murre eggs from two sites in
1997), and the most detrimental, dichlorodiphenyltri- | Atlantic Canada: Machias Seal Island, New Brunswick
chloroethane (DDT), was banned in North America in (44°30'N, 67°06'W), where about 100 pairs of Com-
the 1970s. mon Murre breed beneath large boulders (Diamond
Common Murres (Uria aalge) are cliff-nesting sea- and Devlin 2003; Bond and Diamond 2006) and Gull
birds that typically breed in high-density colonies (Tuck —_ Island (47°16'N, 52°46'W) in the Witless Bay Ecolog-
1961). They build no nest. The shells of their ovate- ical Reserve, Newfoundland and Labrador, which sup-
pyriform to elliptical-ovate eggs are thicker at the nar- _ ports approximately 10 000 breeding pairs of Common
row end, a characteristic shared with Razorbills (A/ca | Murres (S. I. Wilhelm, Canadian Wildlife Service. un-
torda) and presumably an adaptation to nesting on bare _ published data, 2012).
rock (Belopolskii 1961; Ainley ef a/. 2002). Given the We collected 10 whole eggs from Machias Seal Is-
conditions of the nesting habitat, thick shells prevent —_ land in June 2006 and 69 intact and partly broken shells
egg loss and breakage and, therefore, thickness is an from eggs depredated by gulls (Larus spp.) on Gull
important factor in breeding success. The shape of Com- _ Island in July 2012. Because gulls could be targeting
mon Murre eggs is adapted to maximize the surface area younger inexperienced breeders, which have thinner
in contact with the brood patch to facilitate incubation eggshells (Hipfner et a/. 2003), we also collected 20
(Harris and Birkhead 1985; Ainley et al. 2002), and eggs from known established breeders near the centre
the small roll radius, which further decreases as the egg of nesting cliffs on Gull Island in June 2013.
develops, helps prevent the egg from rolling off the For mostly intact eggs from Gull Island, eggshell
ledge (Uspenski 1958). Murres, along with penguins _ thickness (including the egg membrane) was measured
(Spheniscidae), cormorants (Phalacrocoracidae), and at six locations: four equidistant points at the equator
guillemots (Alcidae: Cepphus spp.), have the heaviest — (i.e., largest circumference) and one at each end of the
eggshells relative to whole egg mass of any flying bird — egg. Thickness was measured to the nearest 0.0025 mm
(Schonwetter 1960-1992). using a spring-loaded micrometer. Measurements at

ie

2014

the ends of the eggs were taken on the flattest part to
avoid measuring a strongly curved surface, especially
at the narrow end. Any obvious exterior or interior sur-
face or shell irregularities were avoided. Average thick-
ness of the egg membrane was determined by measur-
ing the difference in shell thickness with and without
the membrane at a sample of adjacent spots on the egg
(n= 10, selected randomly from all eggs). The mean of
the four thickness measurements at the equator (minus
mean egg membrane thickness) was recorded as mid-
dle eggshell thickness. We assessed measurement error
by measuring 21 eggs twice on different days. A single
observer made all measurements.

We measured the length and breadth of intact egg-
shells (2 = 50 from Gull Island, not including any
collected in 2013) with dial callipers, to the nearest
0.1 mm. Because eggs were depredated by gulls, both
measurements could not be made on all eggs (breadth
n= 44, length n = 40). Ground colour and speckling
pattern of eggs were sorted into one of five classes, as
defined by Gaston and Nettleship (1981), with higher
numbers indicating more base and speckling pigmen-
tation in the eggs. As depredated eggs might be more
likely to come from young or inexperienced females
and, therefore, have thinner shells (Hipfner et a/. 1999),
eggshell thickness measurements from Gull Island in
2012 should be taken as a minimum estimate.

Statistical methods

We used repeated-measures analysis of variance
(ANOVA) to examine within-egg variation in shell
thickness between the bottom (round end), middle (lar-
gest circumference), and top (narrow end) of eggs col-
lected from Gull Island in 2012. Only eggs with meas-
urements for all locations were included.

We used Tukey’s honest significant difference (HSD)
post-hoc test for pairwise comparisons. To determine

NOTES 13

whether shell thickness at the top (the measurement
with the largest sample size, n = 58) increased with
increasing ground colour and speckling pattern, we
used Kruskal-Wallis rank-sum test.

We tested relations between egg size (length,
breadth, and volume) and shell thickness using linear
regression. A volume index was calculated as length
x breadth? (Hipfner and Gaston 1999).

Finally, we compared the thickness of our shell sam-
ples with previously published values (Gress e7 al.
1971; Henny ef al. 1982; Pyle et al. 1999; Zimmerman
and Hipfner 2007) and tested differences using ANOVA
and Tukey’s HSD. All tests were performed in R 3.0.2
(R Development Core Team 2013), and differences
were considered significant when P < 0.05. Data are
presented as the mean and standard deviation (SD).

Results

Differences between repeated measurements of the
thickness of the same shells were small (0.016, SD
0.021 mm) and ranged from zero to 0.145 mm or
0.0-8.6% of shell thickness. The membrane meas-
ured 0.070 mm (SD 0.049), or 8.2% of the combined
thickness of the shell and membrane. Among the 21
eggs assessed twice on different days, categorization
of ground colour and speckling pattern differed for two
and five eggs, respectively; in all cases, the assessment
differed in a single category.

Shell thickness varied within each egg at both sites
(Table 1). At Gull Island in 2012, repeated measures
(ANOVA, F, ,, = 310.3; P < 0.001) showed that the
mean thickness of the shell at the bottom (0.687, SD
(0.066 mm) was thinner than at the middle (0.753, SD
0.057 mm, P < 0.001), and the mean thickness of the
shell at the middle was thinner than at the top (0.906,
SD 0.062 mm, P < 0.001). We observed a similar pat-
tern in shells from Machias Seal Island (repeated meas-

TABLE 1. Shell thickness and size of Common Murre (Uria aalge) eggs from Gull Island, Newfoundland and Labrador

(2012) and Machias Seal Island, New Brunswick (2006).

Shell thickness n Range (mm) Mean (mm) SD (mm)
Gull Island

Top 58 0.761—1.040 0.906 0.062
Middle* 56 0.664—0.822 0.753 0.057
Bottom 54 0.568—0.860 0.687 0.066
Machias Seal Island

Top 9 0.780-0.932 0.829 0.058
Middle* 8 0.726—0.907 0.767 0.078
Bottom qi 0.653-0.866 0.684 0.064
Egg size

atte 40 74.2-90.8 82.2 3.6
Breadth* 44 47.6-53.9 50.7 1.4
] ias Seal Island

Spain ‘s 10 79.0-91.0 84.6 *S
Breadtht 10 49.4-52.9 50.0 Bis

e008 0 OOS

*At the “equator” or largest circumference.
+Largest diameter.

74 THE CANADIAN FIELD-NATURALIST

ures ANOVA, F,, = 9.29, P = 0.008); although the
mean thickness of the shell at the top and at the middle
did not differ (P > 0.05), both were thicker than at the
bottom (P < 0.05). On Gull Island, mean shell thickness
in 2012 did not vary with ground colour (y? = —1.91,
df = 4, P = 0.75) or speckling pattern (v7 = —1.31,
df= 4, P=0.86; Table 2).

The mean length and mean diameter of Gull Island
eggs in 2012 were 82.2 mm (SD 3.7) and 50.7 mm
(SD 1.4), respectively; these measures for Machias Seal
Island eggs were 84.6 mm (SD 3.3) and 50.0 mm
(SD 3.5). At both sites, there was no detectable cor-
relation between eggshell thickness at the six meas-
ured locations and length (all P > 0.08 and 7° < 0.08),
diameter (all P > 0.23 and 7° < 0.04), or volume index
(all P > 0.58, and 77 < 0.01).

Common Murre eggs collected from Gull Island in
2012 and Machias Seal Island in 2006 were signifi-
cantly thicker than any previously reported value
(Fe 293 = 57.55, all pairwise comparisons P < 0.05;
range of means: 0.54—0.716 mm; Figure 1). The mean
thickness of the shells of the 20 eggs collected from
known established breeders on Gull Island in 2013 was

Eggshell thickness (mm)

Vol. 128

TABLE 2. Ground colour and speckle pattern on Common
Murre (Uria aalge) eggs from Gull Island, Newfoundland
and Labrador (2012).

Ground colour No. of eggs

Class | (lighter) 14
Class 2 23
Class 3 12
Class 4 10
Class 5 (darker) 7

Unclassified 3

Speckle pattern

Black speckles with blotches at large end 14
Brown spots with a few scribbles 4

Faint brown scribbles 7

Black scribbles 18
Black blotches 24
Unclassified 2

significantly greater than the mean thickness of egg-
shells measured in 2012 on Gull Island (middle: 0.811,
SD 0.040 mm, P < 0.001), but was not significantly
greater than the mean thickness of eggshells on Machi-
as Seal Island (middle: 0.767, SD 0.078 mm, P = 0.58).

A. 8° ¢ D.. bor ean

Study

FIGURE 1. Shell thickness at the equator of Common Murre eggs in Atlantic Canada (G—I) compared with that reported in other
studies (A-F). Groups not sharing the same lowercase letter above are significantly different (Tukey’s HSD, P < 0.05),

Black horizontal lines indicate the mean, shaded boxes are the first and third quartiles, whiskers are the 5—95%

data

range, and open dots are extremes. A, Triangle Island, British Colombia (Zimmerman and Hi fner 2 ;

Islands, California, 1968 and 1970 (Gress et al. 1971); C, Island Rock and Gull Island. Daeeel fo ieee
1982), D, Southeast Farallon Island and Ano Nuevo Island, California, 1993 (Pyle et al. 1999): E, Farallon Islands,
California, 1913 (Gress et al 1971); F, Pacific Coast, northern California to Queen Charlotte Islands, British Colum:
bia, pre-1947 (Henny et al. 1982); G, Gull Island, Newfoundland and Labrador, 2012; H, Machias Seal Island, New

Brunswick, 2006: and I, Gull Island, 2013.

2014

Discussion

Like those of Thick-billed Murres (Uspenski 1958),
eggs of Common Murres are not of uniform thickness.
Greater thickness at the top allows the egg to withstand
the same amount of pressure there as at the bottom, but
over a smaller surface area. Egg size was not related to
shell thickness at any location on the egg, nor was there
any relation between thickness and level of pigmenta-
won.

The eggs collected in Atlantic Canada had the thick-
est shells ever recorded for Common Murres. In the
comparison, we excluded eggs measured at Gull Island
in 1977 and 1978 (0.43 mm, measured by vapour con-
ductance; Mahoney and Threlfall 1981), because they
were measured by a different method: vapour conduc-
tance.

The only other record of shell thickness after the rap-
id decline in use of organochlorine pesticides (Zimmer-
man and Hipfner 2007) also differs significantly from
our measurements. These eggs, collected from Triangle
Island, British Columbia, had a lower average shell
thickness (0.54, SD 0.005 mm), but egg length (85.5,
SD 3.5 mm) and diameter (50.5, SD 1.2 mm) were
similar despite a small sample size (7 = 6 for thick-
ness, n = 15 for length and breadth; Zimmerman and
Hipfner 2007). The thicker eggshells in the Atlantic
region suggest population differences between the two
oceans, where birds are known to experience different
ecological pressures (Ainley et a/. 2002). Differences
in contemporary eggshell thickness could be related to
salinity (Dyck and Kraul 1984) or the availability of
calcium in pre-breeding diets (Bientema ef al. 1997;
Brenninkmeijer ef a/. 1997).

Our study is the first to examine the relation between
egg size and shell thickness in this species. Although
we found no such relation, this is the first investigation
of its kind among Common Murres, and similar re-
search is recommended at other colonies, particularly
those where shell thinning is known to have occurred as
a result of environmental pollutants.

Williams ef al. (1982) found that, among Charadri-
iformes, eggshell mass, as a proportion of total egg
mass, is highest among murres (13.7% for Common
Murres and 14.4% for Thick-billed Murres), compared
with terns (Sternidae: 4.8—10.7%) and gulls (Larus spp.:
5.9-11.8%); this progression corresponds to decreas-
ing rigidity of nest sites (bare rock [murres], sand and
gravel [terns], and plant material [gulls], Williams e¢
al. 1982). Penguins, cormorants, and murres have the
heaviest proportional eggshells among seabirds (Wil-
liams et al. 1982). Although Razorbills (Alca torda) nest
on substrates similar to those used by Common Murres,
they aggregate much less densely (Birkhead 1978), and
their eggshells are proportionally less massive (8.66 +
(0.36%; Birkhead and Nettleship 1984). It 1s possible
that species breeding on a rigid breeding substrate with-
out nesting material and in dense aggregations (relative

NOTES 75

to other seabirds) combine to produce some of the rel-
atively thickest and heaviest eggshells.

Conclusions

Our results show that Common Murres from Atlantic
Canada have the thickest eggshells recorded for this
species and that even the eggshells of supposedly inex-
perienced breeders are thicker than those found else-
where. Thickness varied with location on the egg, and
thickness was not related to egg size or volume. Eggs
of established breeders had thicker shells than those
depredated by gulls, possibly produced by younger
murres. This is one of only a few studies to investigate
eggshell thickness in seabirds since the decline of
organochlorine pesticide use. We recommend further
research to investigate contemporary murre eggshell
thickness throughout the breeding range and possible
ecological, toxicological, or phylogenetic influences.

Acknowledgements

We thank the Canadian Coast Guard and Canadian
Wildlife Service for permission to work on Machias
Seal Island and the Parks and Natural Areas Division
of the Newfoundland and Labrador Department of
Environment and Conservation for permission to work
in the Witless Bay Seabird Ecological Reserve. Col-
lections were made with permission of the Canadian
Wildlife Service (permits ST 2455, SS 2784), and
whole eggs were collected with the approval of the
University of New Brunswick Animal Care Committee
(protocol 06032). Environment Canada, the Atlantic
Cooperative Wildlife Research Network, the Science
Horizons Youth Internship Program, Collaborative
Mercury Research Network, and the Natural Sci-
ences and Engineering Research Council of Canada
provided financial support. We thank G. M. Bouchard,
T. C. Clarke, A. W. Diamond, and A. Patterson (Bold
Coast Charter Co., Cutler, Maine) for logistic and field
support in New Brunswick and K. A. Hobson, G. J.
Robertson, S. I. Wilhelm, the Canadian Wildlife Serv-
ice, and Environment Canada for logistic support in
Newfoundland. Comments from A. J. Gaston, G. J.
Robertson, and an anonymous reviewer improved pre-
vious drafts of this manuscript.

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76 THE CANADIAN FIELD-NATURALIST

Birkhead, T. R. 1978. Behavioural adaptations to high-
density nesting in the Common Guillemot Uria aalge.
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Birkhead, T. R., and D. N. Nettleship. 1984. Egg size, com-
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Bond, A. L., and A. W. Diamond. 2006. The Common Murre
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Diamond, A. W., and C. M. Devlin. 2003. Seabirds as indi-
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Dyck, J., and I. Kraul. 1984. Environmental pollutants and
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Gaston, A. J., and D. N. Nettleship. 1981. The Thick-billed
Murres of Prince Leopold Island. Canadian Wildlife Serv-
ice, Ottawa, Ontario. Monograph Series, no. 6.

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Vol. 128

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Alcidae. Auk 124(1): 307-315.

Received | April 2013
Accepted 30 August 2013

2014 NOTES 77

Tie Most Northerly Black Witch (Ascalapha odorata): A Tropical
Moth in the Canadian Arctic

TORBJORN EKREM!:4, PETER G. KEVAN?, THOMAS S. Woopcock2, and PAUL D. N. HEBERT?

Department of Natural History, Norwegian University of Science and Technology (NTNU) University Museum, NO-7491
Trondheim, Norway

- Canadian Pollination Initiative, School of Environmental Sciences, University of Guelph, 50 Stone Road E., Guelph,
Ontario NIG 2W1 Canada

“Biodiversity Institute of Ontario, University of Guelph, 50 Stone Road E., Guelph, Ontario NIG 2W1 Canada

*Corresponding author: torbjorn.ekrem@ntnu.no

Ekrem, Torbjorn, Peter G. Kevan, Thomas S. Woodcock, and Paul D. N. Hebert. 2014. The most northerly Black Witch
(Ascalapha odorata): a tropical moth in the Canadian Arctic. Canadian Field-Naturalist 128(1): 77-79.

A specimen of the Black Witch (Ascalapha odorata) was collected in August 2006 near Churchill, Manitoba, at 58.7652°N.
This represents the most northerly record for this species. DNA barcode comparison of 93 specimens of A. odorata in the
Barcode of Life Data Systems revealed low genetic divergence even though these specimens were collected from a large
geographical area. The haplotype of the Churchill specimen was shared by only one other individual (collected in the
Yucatan Peninsula of Mexico) in the Barcode of Life Data Systems. A definite assignment of the geographic origin of the
Churchill specimen is not possible with current data, but more extensive analysis of Central American populations with
additional genetic markers might resolve this uncertainty.

Key Words: Black Witch; Mariposa de la muerte; Ascalapha odorata; DNA barcoding; distribution; genetic divergence;
migratory; migration; Manitoba

Ascalapha odorata (Linnaeus) (Black Witch) is one —2008—2014*) (Figure 1), the natural breeding range of
of the largest members of the superfamily Noctuoidea A. odorata is thought to be limited to Central America
in North America. Although it has been collected from and the southernmost United States, including Texas
Canada to Argentina and even from Hawaii (Quinn and Florida (Wagner et al. 2011; Quinn 2008—2014*).

170°W 160°W Ss 150°Wts«éd1OWsid130°Wtsiéd120°WeSsé110°W 100°W 90°wW 80°W 70°W 60°W 50°W 40°w

60°N

30°N 40°N 50°N

20°N

10°N

Churchill record
Quinn (2008-2014)

o

Global Biodiversity Information Facility

10°S

Barcode of Life Data Systems

20°S

FiGurE 1. Collection sites of specimens of Ascalapha odorata (Black Witch) currently registered in the Barcode of Life Data
S) stems (black dots, star), the Global Biodiversity Information Facility (red triangles), and by Quinn (2008-2014*)
ieaaiieo accessed January 2014, most northerly observed record from Bird Cove, 18 km east of Churchill (star).

78 THE CANADIAN FIELD-NATURALIST

The caterpillars of the Black Witch have also been ob-
served on introduced acacias in the greater Los Angeles
area (Wagner ef al. 2011).

This moth is a seasonal migrant to more northerly
regions of North America. Most Canadian records are
from summer and early autumn. Quinn (2008—2014*)
gives an excellent summary of records from the Unit-
ed States and Canada, and several additional records
are available through the Global Biodiversity Infor-
mation Facility (http://www.gbif.org). These are shown
in Figure |. Additional North American sightings can
be seen in the Butterflies and Moths of North America
website (Opler et al. 2013*) and the North American
Moth Photographers Group website (North American
Moth Photographers Group 2014*), although many of
these records are the same as those reported by Quinn
(2008-2014*).

Most past Canadian records derive from sites in
southern Canada, but specimens have a broad longitu-
dinal distribution, with records from Newfoundland
to British Columbia. In 2012 there were particularly
frequent incursions of this moth into Canada (Quinn
2008—2014*).

The species also appears to engage in southerly and
altitudinal migrations, as specimens are known from
as far south as Argentina and into Andean locales
(Henderson 2002). Its broad dispersal to temperate
and high-elevation sites may reflect migratory flights
to cooler environments, as is known for a number of
other noctuoids (Kevan and Kendall 1997).

Ascalapha odorata 1s a recurring visitor to Canada,
but most past records have been restricted to southern
regions of the country. Although one of its larval host
plants, Gymnocladus dioicus (Kentucky Coffee-tree),
is native to Carolinian Canada and adult food resources
of over-ripe fruit (e.g., Prunus spp.) are widespread
in Canada, this species is unlikely to become estab-
lished, as all life stages are believed to be intolerant
to prolonged freezes (Wagner et al. 2011).

The primary motivation for this paper lies in report-
ing the discovery of a live female Ascalapha odorata
on 18 August 2006 resting on Empetrum nigrum (Black
Crowberry) at Bird Cove, a rocky headland on Hud-
son Bay about 18 km east of Churchill, Manitoba
(58.7652°N, 93.8682°W) (Figure 2). This specimen
represents the most northerly record for any Black
Witch, displacing the previous record holder, a speci-
men collected at Auke Bay, 13 km north of Juneau,
Alaska (58.3833°N, 134.5822°W) (Spangler 1957).

The Churchill specimen is the first individual of A.
odorata found in the Low Arctic tundra. This individ-
ual probably arrived in the Hudson Bay area on strong
southerly winds about a week before its discovery (P.
Kevan, personal observation). The specimen is now
housed in the Biodiversity Institute of Ontario at the
University of Guelph, and its photograph is accessible
on the Barcode of Life Data Systems (Ratnasingham
and Hebert 2007): http://www.boldsystems.org/index
-php/Public_RecordView?processid=DSCNI024-07.

Vol. 128

FIGURE 2. The presence of Ascalapha odorata (Black Witch)
near Churchill, Manitoba. A and B: The worn but liv-
ing specimen collected near Bird Cove, 18 km east
of Churchill, Manitoba, on 18 August 2016. C: Paul
Hebert documents the most exciting catch of the day.
Photos: T. Ekrem.

Ninety-three sequences from the DNA barcode re-
gion of the cytochrome c oxidase | gene are currently
available for A. odorata on the Barcode of Life Data
Systems (Ratnasingham and Hebert 2007). The records
are from a wide geographical range (Figure 1), but all
are members of the same Barcode Index Number (clus-
ter) (BOLD:AAA5595*).

The average p-distance among these sequences is
0.28%, and the maximum distance is 1.38%. The p-
distance to the nearest neighbours is 6.6%. Most of the
A. odorata sequences fall into a single cluster, but the
Churchill specimen belongs to a rather divergent group,
represented by just three other individuals, two from
the Yucatan Peninsula (Mexico) and one from Costa
Rica (Figure 3).

Although it may never be possible to firmly estab-
lish the point of origin of the Churchill specimen, fur-
ther analysis of barcode variation in Central and South
American populations of 4. odorata coupled with the

e
89

2014

[|__|
0.2%

NOTES 719

CR (39)
Mex (22)

Arg (7)
US (7)
Bra (3)
Peru (2)
FG(1)
Ecu (1)

CR (4), US (2)

@ Mex (1)
O Can (1)

FIGURE 3. Neighbour-joining tree of DNA barcodes for Ascalapha odorata (Black Witch). Cluster with the specimen collected
near Bird Cove, 18 km east of Churchill, Manitoba, on 18 August 2016, is coded: Churchill (open circle), Yucatan
Peninsula (grey dots), Costa Rica (square). The tree was generated using Kimura 2-Parameter distances and pair-
wise deletion of gaps with MEGAS (Tamura ef al. 2011).

analysis of additional gene markers might permit the
resolution of this uncertainty.

Acknowledgements

We thank Megan Milton for producing the figure for
the neighbour-joining (NJ) tree and Marc Daverdin for
producing the distribution map. Fieldwork in Churchill
was supported by an International Polar Year grant to
PDNH and PGK from the Natural Sciences and Engi-
neering Research Council of Canada (NSERC). Se-
quence analysis was enabled by a grant from the Gov-
ernment of Canada through Genome Canada and the
Ontario Genomics Institute in support of the Inter-
national Barcode of Life project. We thank Winnie
Hallwachs, Cecilia Kopuchian, Daniel Janzen, Carmen
Pozo, and Carlos Lopez Vaamonde for allowing the
use of barcode records under their curation.

Documents Cited (marked * in text)

BOLD:AAA5595. 2014. Barcode Index Number cluster for
Ascalapha odorata. Analysed dataset accessible at: dx.doi
-org/10.5883/DS-BWMC.

North American Moth Photographers Group at the Mis-
sissippi Entomological Museum at Mississippi State
University. 2014. 930759 — 8649 — Ascalapha odorata
Black Witch Moth — (Linnaeus, 1758). http://mothphoto
graphersgroup.msstate.edu/species.php?hodges=8649.
(Accessed 5 January 2014). .

Opler, P. A., K. Lotts, and T. Naberhaus. 2013. Butterflies
and moths of North America. http://www. butterfliesand
moths.org/. (Accessed 5 January 2014).

Quinn, M. 2008-2014. The black witch moth: its natural
and cultural history. http://www.texasento.net/witch.htm.
(Accessed 5 January 2014).

Literature Cited

Henderson, C. L. 2002. Field Guide to the Wildlife of Costa
Rica. University of Texas Press, Austin, Texas. 539 pages.

Kevan, P. G., and D. M. Kendall. 1997. Liquid assets for fat
bankers: summer nectarivory by migratory moths in the
Rocky Mountains, Colorado, U.S.A. Arctic and Alpine
Research 29(4): 478-482.

Ratnasingham, S., and P. D. N. Hebert. 2007. BOLD: The
Barcode of Life Data System (www.barcodinglife.org).
Molecular Ecology Notes 7: 355-364. doi 10.1111/j.1471
-8286.2006.01678.x. (Accessed 28 December 2013).

Spangler, P. J. 1957. A record of the black witch, Erebus odo-
ra [sic] (Noctuidae), in Alaska. Journal of the Lepidopter-
ists’ Society 11(6): 205.

Tamura, K., D. Peterson, N. Peterson, G. Stecher, M. Nei,
and S. Kumar. 2011. MEGAS: Molecular Evolutionary
Genetics Analysis using Maximum Likelihood, Evolution-
ary Distance, and Maximum Parsimony Methods. Molec-
ular Biology and Evolution 28: 2731-2739. doi 10.1093
/molbev/msr121.

Wagner, D. L., D. F. Schweitzer, J. B. Sullivan, and R. C.
Reardon. 2011. Owlet Caterpillars of Eastern North Amer-
ica. Princeton University Press, Princeton, New Jersey.
576 pages.

Received 10 September 2013
Accepted 13 December 2013

80 THE CANADIAN FIELD-NATURALIST

Vol. 128

Multiple Crossings of a Large Glacial River by Canada Lynx (Lynx

canadensis)

DASHIELL FEIERABEND!>? and KNUT KIELLAND!*2

‘Department of Biology and Wildlife, University of Alaska Fairbanks, Fairbanks, Alaska 99775 USA
*Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, Alaska 99775 USA

‘Corresponding author: dfeierab@alaska.edu

Feierabend, Dashiell, and Knut Kielland. 2014. Multiple crossings of a large glacial river by Canada Lynx (Lynx canadensis).

Canadian Field-Naturalist 128(1): 80-83.

Rivers may act as barriers to the movement of terrestrial mammals, which could limit dispersal and gene flow. Glacial
rivers are particularly hazardous because of the cold water temperature and swift current. Yet, we determined that 2 Canada
Lynx (Lynx canadensis) equipped with GPS collars repeatedly swam across the main channel of the Tanana River in interior
Alaska in 2011 as late in the season as November, when the average minimum daily air temperature was —27°C. These
observations are consistent with the low level of genetic structure observed in Canada Lynx in northwestern North America
and suggest that even large rivers may pose less of a barrier to movement by Canada Lynx than expected.

Key Words: barrier; boreal forests; Canada Lynx; Lynx canadensis; dispersal; home range; river; Alaska

Introduction

Rivers may act as barriers to the movement of ter-
restrial mammals because of the risk of injury or death
from drowning (Storm ef al. 1976; Peres et al. 1996;
Garroway et al. 2011). By limiting dispersal into poten-
tially favorable habitats as well as constraining gene
flow, such barriers may have both ecological and genet-
ic consequences (Puth and Wilson 2001). In northern
regions, the near-freezing temperatures of glacial rivers
add to the dangers posed by swift currents and floating
debris.

Whereas large ungulates such as Moose (Alces amer-
icanus ) and Caribou (Rangifer tarandus) regularly cross
open glacial rivers during seasonal migrations (Grif-
fith et al. 2002; Kellie 2005), these rivers presumably
pose considerably more danger to medium-sized mam-
mals, such as Coyotes (Canis latrans) and Canada Lynx
(Lynx canadensis).

The only published report of a Canada Lynx cross-
ing an open river describes an individual that swam
across a 3.2 km wide section of the Yukon River (Koba-
lenko 1997). There are several unpublished reports of
Canada Lynx crossing open rivers in Alaska and Cana-
da, but the frequency with which individual Canada
Lynx make such crossings is unknown. Here, we report
the behavior of 2 Canada Lynx that repeatedly crossed
the glacially fed Tanana River in interior Alaska during
fall and early winter, before the river was sufficiently
frozen to allow them to cross on foot.

Methods
Study area

This study took place in the Bonanza Creek Long-
Term Ecological Research (LTER) site (65°, —148°)
and surrounding areas, approximately 20 km south-
west of Fairbanks, Alaska, U.S.A. Upland, lowland, and
floodplain boreal habitats in the study area included

Black Spruce (Picea mariana), White Spruce (Picea
glauca), Balsam Poplar (Populus balsamifera), mixed
White Spruce—Alaska Paper Birch (Betula neoalas-
kana), Trembling Aspen (Populus tremuloides), and
regenerating stands of Alaska Paper Birch, Trembling
Aspen, and shrub birch (Betu/a spp.) in areas that had
burned in the previous 30 years.

The Tanana River, which is the largest tributary of
the Yukon River, flows across the southern edge of
the site and is highly braided. The Tanana River has an
average discharge volume of 1600 m>- s! with a low
of approximately 560 m? - s! in spring and fall prior
to freeze-up, but it can reach a maximum of about
2300 m3 - s! during floods in late summer (USGS
2013*).

The river is typically frozen from late November
until April-May (KK, personal observation). Open-
water temperatures in late fall/early winter are just
above freezing (KK, unpublished data). Average maxi-
mum and minimum air temperatures in the study area
were 15°C and 4°C, respectively, in September 2011,
3°C and —6°C, respectively, in October 2011, and
—18°C and —27°C, respectively, in November 2011
(Bonanza Creek LTER 2013*).

Canada Lynx capture and collaring

Between June 2008 and November 2012, we used
padded Soft-catch® traps (No. 3, Woodstream Corp.,
Lititz, Penn.), Belisle foot snares (Wildlife Control
Supplies, East Granby, Conn.), and home-made cage
traps to live-capture 23 Canada Lynx. Scent lures and
animal carcasses were used to bait traps. We adminis-
tered Telazol (3 mg/kg) to trapped individuals using a

jab stick, then determined sex and weight, and we fit-

ted 12 individuals with timed-release collars equipped
with global positioning system (GPS) loggers, VHF
transmitters, and activity sensors (model G2110B.
Advanced Telemetry Systems, Isanti, Minn). Collars

2014

weighed 350 g (approximately 2-4% of the mass of
Canada Lynx in Alaska) (Alaska Department of Fish
and Game 2008*) and were not expected to affect
survival (Sikes er al. 2011). The activity sensor pro-
vided a count of the number of changes in the posi-
tion of a tilt switch on the collar and represented a
unitless measure of an animal’s activity. Collars were
programmed to record locations at 5-hour intervals.

Capture and handling procedures followed animal
care and use guidelines of the American Society of
Mammalogists (Sikes e¢ a/. 2011) and were approved
by the University of Alaska Fairbanks Institutional
Animal Care and Use Committee (Protocol 135202-3)
and the Alaska Department of Fish and Game (Permit
11-041).

Analysis

We report movements for 2 adult Canada Lynx: a
male collared from 2 September to 5 November 2011
and a female collared from 15 August to 21 Novem-
ber 2011. We recovered the male’s collar after it pre-
maturely released from the animal, and we recovered
the female’s collar after the animal was trapped by a
professional fur trapper. We counted the number of
times each Canada Lynx crossed the main channel of
the Tanana River as well as major braids and sloughs,
based on 0.6-m resolution DOQQ imagery (USDA
2007*) in ArcGIS 10.0 (ESRI, Redlands, Calif.). Cross-
ings were identified by consecutive GPS locations sit-
uated on opposite sides of the river. Because Canada
Lynx could conceivably have entered and exited the
river from the same bank, this count represents the min-
imum number of times each Canada Lynx was in the
river during the sampling period.

We estimated home range sizes using Brownian
bridge movement models (Horne ef al. 2007). Suc-
cess rates for scheduled GPS fixes were 49% and 61%
for the male and female Canada Lynx, respectively.
GPS accuracy varied, depending on the orientation of
the GPS antenna to the sky and the number and con-
figuration of satellites available.

We estimated location accuracy using two stationary
collars programmed to collect fixes at 5-hour intervals
in the field under dense vegetative cover and open
sky for 60 hours. Using the Standard Distance tool in
ArcGIS, we found the standard deviations from the
mean centers of GPS locations to be 6.7 m (n = 12)
and 3.1 m (m= 12) under cover and open sky, respec-
tively. When calculating Canada Lynx home ranges,
we used a conservative estimated location error of 30
m based on data from Horne ef al. (2007).

We used all GPS location fixes (146 for the male
and 287 for the female) over all days fixes were ob-
tained (64 days for the male and 98 days for the fe-
male). We calculated utilization distributions using the
kernelbb function (adehabitat package) in R (R Core
Team 2013*) and extracted isopleths using Geospatial
Modeling Environment (Beyer 2OU2F i

NOTES 8]

Home range boundaries were defined by 90% iso-
pleths (Borger e¢ a/. 2006), and core use areas were
defined by isopleths (male = 37%; female = 33%)
that divided intensively used areas from peripheral
areas of less intense use. This was done by fitting an
exponential regression to a plot of utilization distri-
bution (UD) area against utilization distribution vol-
ume (i.e., isopleth value) and identifying the utiliza-
tion distribution volume for which the slope of the
regression line was equal to | (Vander Wal and Rodgers
2012).

We quantified diel activity patterns using average
collar activity counts and movement rates derived from
distances between consecutive GPS locations. In order
to avoid biased movement rates over longer intervals,
we used only GPS locations that were 5 hours apart.
Because activity accrued and movements occurred over
5-hour periods, we calculated activity and movement
for a given time of day as the average of all 5-hour
periods containing that time (e.g., activity at 1300 was
calculated as an average of activity counts that spanned
the time periods 0900-1359, 1000-1459, 1100-1559,
1200-1659, and 1300-1759).

Results

We collected location data from a total of 12 Cana-
da Lynx equipped with GPS collars between Decem-
ber 2009 and January 2013. Of the 10 animals whose
data spanned at least one month, 2 Canada Lynx made
crossings of the Tanana River when it was frozen in
April, 1 male crossed the main channel twice in two
days when the river was still open in October, another
male crossed the main channel 6 times in November,
when the river was likely only partially frozen, and a
male and a female repeatedly crossed the unfrozen riv-
er between September and November. Only the home
ranges of the latter 2 individuals spanned the river.

Of those 2 Canada Lynx, the male made 14 cross-
ings of the main channel of the Tanana River between
4 September and 4 November 2011, and he swam
across smaller river braids and sloughs (15-50 m in
width) an additional 20 times (Figure 1). The female
made 11 crossings of the main river channel between
13 September and 21 November, and she swam across
smaller river braids and sloughs an additional 40 times
(Figure 1).

We could not determine the exact time of day of
most crossings. However, both Canada Lynx crossed
river channels multiple times in a given 24-hour period
and each animal crossed the main channel at least 3
times at night. The male exhibited a more diurnal pat-
tern of activity than the female (Figure 2). Home ranges
were 201 km? and 254 km? for the male and female,
respectively. River water constituted 6% of each home
range, and river water constituted 12% and 18% of core
use areas for the male and the female, respectively.
Both of the Canada Lynx made considerable use of is-
lands in the river (28% of locations for the male and
52% of locations for the female).

82 THE CANADIAN FIELD-NATURALIST

* GPS Location
— River Crossing
MM Tanana River %

OL ee
oes Kilometers

FIGURE |. Movement across the Tanana River near Fairbanks,
Alaska (65°, -148°), by (A) a female Canada Lynx
(Lynx canadensis) equipped with a GPS collar be-
tween 15 August and 21 November 2011 and (B) a
male Canada Lynx equipped with a GPS collar be-
tween 2 September and 5 November 2011. Lines con-
nect consecutive GPS locations for Canada Lynx on
either side of the river but do not indicate the actual
routes travelled.

Discussion

We documented the regular crossing of a large, swift
glacial river by 2 Canada Lynx during times of the
year when cold air temperatures and initial ice forma-
tion elevated the dangers of such behavior. Due to the
length of time between GPS locations, we could not
determine the exact entry and exit points of the Cana-
da Lynx on the river, but we can be reasonably sure
that both Canada Lynx spent considerable time travers-
ing sections of flowing water that were 100-300 m in
width.

Assuming an estimated maximum swimming rate of
50 m- min’! (the maximum sprinting speed of a human
swimmer is 120 m- min!) (Toussaint and Trultens
2005) and an actual travel distance twice the width of
the river channel (to allow for the current, which was
1-2 m-s'') (Johnson et a/. 2010*), we estimate that
it would have taken a Canada Lynx 4-12 minutes to
cross the main stem of the Tanana River, depending on
channel configuration. Moreover, in October and No-
vember, shelf ice along the banks of the river could
have made exiting the water difficult. Sub-freezing air

Vol. 128

~~ —Male- Movement “Female - Movement
—-Male - Activity ---Female - Activity |
207 ~~ a 4.9000
A
= ho gem - g000
& 1S =
= 6000 8
a 1M) z
= 4000 2
4 :
90.5 :
s 2000

0:0) ASSASSINS ee O
0 37 “6 9) 9 129915. “2s P20
Hour

FicurE 2. Diel activity and movement patterns of a male
Canada Lynx (Lynx canadensis) equipped with a
GPS collar between 2 September and 5 November
2011 and a female Canada Lynx equipped with a GPS
collar between 15 August and 21 November 2011 near
Fairbanks, Alaska. Activity is quantified as the num-
ber of changes in the position of a tilt switch on the
collar. Black lines along the x-axis indicate periods
of sunrise and sunset.

temperatures surely caused the animals’ fur to freeze,
drastically reducing its insulative capacity, yet both
Canada Lynx often crossed the river multiple times in
a single day. We can only speculate as to why these
individuals swam across the cold river.

The abundance of Snowshoe Hares (Lepus ameri-
canus) in the study area reached a cyclical peak in the
fall of 2009 and had declined by nearly 50% by the fall
of 2010 (Feierabend 2013). We do not have a direct
measure of Canada Lynx abundance during this time,
but Canada Lynx tracks remained common in the study
area through the winter of 2010-2011 despite the declin-
ing numbers of “sealed” Canada Lynx pelts (pelts with
an official marker or locking seal tag (seal) placed by
the Alaska Department of Fish and Game) (all Canada
Lynx taken in Alaska must be sealed) in interior Alaska
following the winter of 2008-2009 (Alaska Department
of Fish and Game 2012*). It may be that Canada Lynx
were foraging over increasingly large areas as Snow-
shoe Hare populations declined. The islands used by
the Canada Lynx in this study were up to 4 km? in size
and could have harbored populations of Snowshoe
Hares, but most GPS locations were on much smaller
islands that were more likely used as stepping stones
during river crossings than for foraging.

Alternatively, the territoriality of neighboring Cana-
da Lynx may have forced the 2 observed individuals
to concentrate their movements along the margins of
the river. However, given the apparent risks of crossing
the Tanana River, one might expect a Canada Lynx sim-
ply to cross once in order to explore more suitable habi-
tat in the expansive Tanana Flats to the south. In fact.
both individuals did travel up to 30 km southeast of the

2014

river for as long as 12 days at a time before returning to
their previous locations, where they again crossed the
river.

These observations suggest that even hazardous
rivers in freezing conditions may not represent signif-
icant barriers to highly mobile mesocarnivores such
as Canada Lynx and that foraging movements as well
as dispersal that include river crossings may take place
under such conditions. This ability to traverse apparent
physical barriers is consistent with observations of
long-distance dispersal capacity in the Canada Lynx
(Mowat et al. 2000) and the low level of genetic struc-
ture among Canada Lynx populations in northwestern
North America (Rueness er a/. 2003; Row et al. 2012).

Acknowledgements

This research was supported by the Bonanza Creek
Long-Term Ecological Research program funded joint-
ly by the National Science Foundation (DEB-1026415)
and the U.S. Forest Service, U.S. Department of Agri-
culture (PNWO1-JV11261952-231). We thank K. Olson
for help in the field and K. Hundertmark, T. Jung, E.
Koen, and M. O’ Donoghue for critical comments on
the manuscript.

Documents Cited (marked * in text)

Alaska Department of Fish and Game. 2008. Lynx: Wildlife
Notebook Series. http://www.adfg.alaska.gov/static/edu
cation/wns/lynx. (Accessed 16 January 2014).

Alaska Department of Fish and Game. 2012. Trapper
Questionnaire. Statewide Annual Report: 1 July 2010-30
June 2011. http://www.adfg.alaska.gov/index.cfm?adfg=
trapping.reports. (Accessed | June 2013).

Beyer, H. L. 2012. Geospatial Modeling Environment (Ver-
sion 0.7.1.0). http://www.spatialecology.com/gme.

Bonanza Creek LTER. 2013. Bonanza Creek LTER Data.
http://www. lter.uaf.edu/data.cfm. (Accessed | September
2013).

Johnson, J. B., P. Duvoy, K. Moerlein, A. E. Rosenberg-
er, J. W. Schmid, A. E. Seitz, and H. A. Toniolo. 2010.
Assessment of the Tanana River (Nenana, AK) for hydro-
kinetic turbine operations. http://www.akenergyauthority
.org/OceanRiver/2010-2-16_ACEPLecture.pdf.
(Accessed 21 September 2013).

R Core Team. 2013. R: A language and environment for sta-
tistical computing. R Foundation for Statistical Computing,
Vienna, Austria. http://www.R-project.org.

USDA. 2007. Natural Resources Conservation Science Geo-
spatial Gateway. http://datagateway.nrcs.usda.gov. (Ac-
cessed 25 July 2013).

USGS. 2013. Surface-Water Data for Alaska. http://water
data.usgs.gov/ak/nwis/sw. (Accessed 22 September 2013).

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Received 24 October 2013
Accepted 15 November 2013

A Tribute to Kenneth William Stewart, 1936-2011

FRANCIS R. COOK

Curator Emeritus and Research Associate, Canadian Museum of Nature, Ottawa, Ontario KOG 1RO Canada; email: frcook

@ripnet .com

Cook, Francis R. 2014. A tribute to Kenneth William Stewart, 1936-2011. Canadian Field-Naturalist 128(1): 84-90.

Kenneth William Stewart, Manitoba herpetologist
and ichthyologist, died at Victoria General Hospital in
Winnipeg, Manitoba, on 4 July 2011 after an extended
battle with the HIN1 virus. Ken was born 30 December
1936 in Madison, Wisconsin, to Drs. John and Gertrude
Stewart. Growing up in Clinton, Iowa, he spent much of
his youth developing a passion for fishing and nature
on the waters of the Mississippi River (Figure |). He
earned a B.Sc. in Dendrology from the Colorado State
College of Agriculture and Mechanic Arts (renamed
Colorado State University in 1957), Fort Collins, Col-
orado; an M.Sc. in Marine Biology from the University
of Miami, Coral Gables, Florida; and a Ph.D. in Zool-
ogy from the University of British Columbia, Vancou-
ver, with a thesis on hybridization between two species
of minnow in different genera. Bessie Wenzel (born |
April 1939) married Ken in Bessie’s birth place of
Camanche, Iowa, 11 September 1956. Bessie assisted
him on many of the field trips during his student years
at UBC and was one of the lucky souls to decipher his
hand-writing in order to type up his dissertations.

In July 1966, Ken accepted a position in the Depart-
ment of Zoology at the University of Manitoba as Assis-
tant Professor. From his arrival, he made a major contri-
bution to the remarkable student and staff fellowship
of a largely newly recruited zoology department under
Harold Welch. Welch had been lured away from the fed-
eral Department of Agriculture in the mid-1960s with
the challenge of expanding the department, and had
quickly blended established researchers like the recent-
ly added Casimir Lindsey and veterans like George
Lubinsky and Fred Ward, with a bunch of newly mint-
ed Ph.D.s from various sources that included, besides
Stewart, Mike Aleksiuk, Roger Green, Jack Gee, Roger
Evans, and, shortly after, the established W. O. (Bill)
Pruitt and others.

“Do ye ken Dr. Stewart at the break of day” (the
refrain from a rollicking rendition of an old Scottish
melody laced with laudatory original verses depicting
Ken’s return from all-night frog collecting) still rings
heartily in my ears as I write this, decades later. This
occasion was an introduction for a departmental seminar
presentation by Ken in the winter of 1968-1969 on the
developing results of his frog research in Manitoba.
This introduction was in partial retaliation for one by
Ken for an earlier seminar by Cas Lindsey. Like many
endeavours in the zoology department of the University

84

FIGURE 1. A young Ken Stewart: beginnings of a lifetime of
scientific curiosity at about | or 2, probably at Madi-
son, Wisconsin about 1938. Photo: Stewart family
album.

of Manitoba at the time, it originated with Lindsey incit-
ing the collaboration of a crew of enthusiastic younger
professors and graduate students. This model department
generated fresh excitement and productivity virtually
daily, a utopia for a student. Not the least in creating this
atmosphere of fellowship in the zoology department
were the sometimes scathing and always humorous and
good-natured barbs contributed by Ken, whose presence
ensured that no one lost perspective. Ken was also an
enthusiastic participant in (and sometimes victim of)
the annual departmental Christmas party skits, where
students and professors parodied each other, a tradition
brought from the University of British Columbia by
Lindsey and his former students. ;
In 1972, Ken was promoted to Associate Professor
and in 1984 to Professor. He retired in 2000 after teach-
ing for 34 years. He gave courses on the biology of
fishes, the biology of amphibians and reptiles, the prin-

2014

ciples and process of evolution (jointly with B. J. Hann),
the ethics and principles of animal care, and an intro-
duction to chordate zoology. On retirement, he was ap-
pointed a Senior Scholar at the University of Manitoba.

Former student Doug Watkinson who co-authored
The Freshwater Fishes of Manitoba, providing 100 of
the 160 photographs commented: “Ken had an amaz-
ing memory and was able to cite the Latin names of
fishes from all over the world. He was interested in
many different fields of science; I know physics was
one of his favourite reading topics. | guess another way
of putting it is he had a tremendous breadth and depth
of knowledge on numerous topics. He loved to teach
and was a great story teller. When you were in one of
your classes you felt inspired by his passion on the top-
ic. He really seemed to enjoy the people he worked
with and the students. He was an avid outdoors per-
son, enjoying fishing and hunting whenever he could
find the time. He was keen to share a fishing trip with
new University staff and students in his boat on the Red
River. If the person did not fish at all that was often all
the more reason to take them out. Really he was gen-
erous both with his time and sharing his pastime. He
had no fear of ethanol, isopropanol or formaldehyde
and the possible effects they had on his skin. It was very
typical for him to plunge his hands and arms into a vat
of fish with 20 year old isopropanol running into his
shirt and dripping on his pants. His office was a disor-
ganized disaster to look at but if you asked him for
something he would instantly reach into the piles of
papers and pull out what he needed. In contrast to his
office organization he was meticulous with his notes on
whatever project he was working on.”

Another former student Gavin Hanke recalls that
Ken’s lectures on anatomy and lab discussions includ-
ed tests of attention by mention of the foramen Reming-
ton (i.e., bullet holes) in skulls. Gavin also recalled,
“Rarely was Ken tricked in return, but it did happen,
and he took it with the usual humour we knew to ex-
pect. On one occasion he boldly stated that only the
extinct acanthodian fishes had pectoral fin spines. Then
a student pointed out that his favourite fishing quarry
(Channel Catfish) had pectoral fin spines. The look on
Ken’s face was priceless,” as was his response. His ear-
ly lectures annually featured a guest lecture by Cas
Lindsey to present his classic paper on the phylogeny
of dragons to demonstrate classification methods.

Ken was an active participant in many zoology de-
partment and university committees, developing guide-
lines as a member (later chair) of animal care and serv-
ing on committees at the University of Manitoba and
representing it on interuniversity committees through-
out his tenure. Included in his other committee contri-
butions was membership and later chair of zoology
curriculum review committees.

Ken was also interested in reaching out to the public.
He presented lectures and demonstrations related to
amphibian and reptile biology at primary and secondary

Cook: A TRIBUTE TO KENNETH WILLIAM STEWART, 1936-2011 85

schools, Manitoba Schools Science Symposium, and
community organizations, such as Scouts, averaging
12 per year; he made major contributions to conser-
vation in Manitoba by conducting tours of Red-sided
Gartersnake (Thamnophis sirtalis parietalis) dens in
the Interlake district for the Manitoba Naturalists
Society (now Nature Manitoba) every year; he was an
advisor to the British Broadcasting Corporation on
Gartersnake (Thamnophis sirtalis) biology during the
production of David Attenborough’s “Life on Earth”
series (approximately 5-minute portion of programme
5) and he was an advisor to film crews from the Cana-
dian Broadcasting Corporation and the Royal Ontario
Museum on Gartersnake biology for the production of
educational films. He presented interpretive lectures for
Manitoba Provincial Parks on Manitoba amphibians
and reptiles (Spruce Woods Provincial Park 1981—
1991). He served as advisor to the provincial Wildlife
Branch on amphibian and reptile biology in 1971-1972
on the status of and protection extended to amphibians
and reptiles by revision of the Wildlife Act; in 1972 and
subsequently, he provided continuing advice on the
regulation of the commercial harvest of Gartersnakes
and Northern Leopard Frogs (Lithobates pipiens) dur-
ing a period when biological supply houses were inten-
sively harvesting from the large populations of these
species in Manitoba; with the Manitoba Naturalists,
he advised on the protection of communal Red-sided
Gartersnake dens at Narcisse, where a cluster of dens
was designated a Provincial Wildlife Management Area;
he advised the Manitoba Conservation Data Centre on
the status of fish, amphibian, and reptile species in
Manitoba; in 1989 and after, he advised the Wildlife
and Fisheries branches regarding control of nesting
Double-crested Cormorant (Phalacrocorax auritus)
populations on Lake Winnipegosis.

During the two years (1968-1970) I spent in Win-
nipeg, to fulfill the university residence requirement for
a Ph.D., and afterward, when I returned to the National
Museum of Canada to complete my thesis (among my
other duties), Ken doggedly supported my efforts, and
the degree was finally obtained in 1978. As a Ph.D.
supervisor, Ken was ideal, always generating sugges-
tions for additional approaches and then leaving the stu-
dent the independence to decide what advice to take and
what to use of his own ideas. But he allowed little argu-
ment over ruthlessly cutting the verbosity in his stu-
dents’ theses drafts. (If Ken were alive, this tribute
would be a fraction of its present length.)

Ken joined field trips with awesome enthusiasm. |
recall these with special relish for the successes and
comradeship as we investigated the fascinating biolog-
ical transition zone east of Winnipeg to the Ontario
border. Ken and I vied, often separately, in nightly
spring outings to plug holes in our understanding of
Manitoba amphibian distribution, especially the dis-
tribution of call types (“species”) of the Gray Treefrog
(Hyla versicolor). Each morning in the lab, we would

86 THE CANADIAN FIELD-NATURALIST

exchange our latest respective successes and individual
theories and differing interpretations. He also promoted
seldom studied winter herpetology by in-season scuba
diving for Mudpuppies (Necturus maculosus) in the
waters of the Brokenhead River and skidooing on
frozen Lake Manitoba to investigate the Leopard Frogs
found in the bottom meshes of fisher’s nets far out from

shore. Jim Johnston took his B.Sc. at the University of

Manitoba and was Ken’s field assistant until he ac-
cepted an offer to be herpetological technician at the
National Museum of Natural Sciences in 1972. He
stayed until 1979, when his interest in photography led
to a position with the Canadian Museum of Science and
Technology.

Patrick Gregory arrived in Manitoba a year after |
did, after completing his B.Sc. at the University of Tor-
onto. He left with not only a master’s degree (1971) but
also a doctoral degree (1973), both based on research
assisted by the Great Dane Jason on the incredible Red-
sided Gartersnake dens in the Interlake area, until then
unstudied. It is a tribute to Ken’s stimulating role as
supervisor that Gregory, Tom Vincent (M.Sc., 1971),
and Don Hart (M.Sc., 1975) completed graduate de-
grees based on snake populations from this area (four
theses; eight publications) (Figure 2). Subsequently,
Vincent continued his Interlake snake studies in a Ph.D.
thesis under Diane Secoy at the University of Regina.
Further, Ken interested departmental colleague and
physiologist Mike Aleksiuk in the Red-sided Garter-
snakes, and Mike with his students added another 14
papers. Then David Crews from Harvard University
with Mike Garstska and other students subsequently
analyzed the behaviour, pheromones, and other aspects
of the Red-sided Gartersnake to extend studies there by
over 30 additional papers. Gregory went on to the Uni-
versity of Victoria, where he remains to this day as Pro-
fessor, with a term served as department head and a
succession of papers and students extending the studies
of Gartersnake ecology to British Columbia, Alberta,
and the Northwest Territories and to other snakes, tur-
tles, and even salamanders and frogs from British Col-
umbia.

Ken did have other graduate students in reptiles and
amphibians after Pat and me, so it is untrue that shep-
herding the two of us to doctorates completely wore out
his enthusiasm for herpetology. His change in emphasis
was more likely triggered by the departure of ichthyol-

ogist Cas Lindsey, who returned to the University of

British Columbia. After that, Ken resumed work on his
initial interest, fish (even including some palaeontol-
ogy—a true measure of versatility and flexibility), For-
mer graduate student Gavin Hanke has pointed out that
it was fitting that Ken’s paleontological interests fo-
cused on fishes from the Devonian, the Age of Fishes.
A succession of papers and graduate students followed
for these lower vertebrates, and Ken extended his field

experience occasionally beyond Manitoba with trips to
Fiji and Africa.

Vol. 128

FIGURE 2. Ken Stewart at a Red-sided Garter Snake (Tham-
nophis sirtalis parietalis) den in the Interlake region,
April 1983. Photo: D. Berezanski.

Though brought up farther south and with only an
initial introduction to Canada in British Columbia, Ken
embraced the shockingly different climate of Manitoba
with the same unnerving cheer with which he met any
challenge. He added a brace of Malamutes (a breed of
Huskies he had come to admire during vacation fishing
trips to the Northwest Territories) to his family. These
dogs were regularly harnessed to sleds he built himself
in the traditional native way. Ken drove them on the
streets in his residential Winnipeg suburb and beyond
to the surrounding windswept spaces most of us avoid-
ed from October until April.

Ken and I shared a commitment to dogs (though we
differed greatly on breeds favoured), but Ken drew a
personal line on cats. He never could gain acceptance
by Tasha, our independent-minded Siamese, who may
have realized this, as she invariably quietly welcomed
his visits by jumping up on the couch beside him and
biting him under the arm. It was a measure of his toler-
ance that he invariably took this with good humour and
restraint, although not without increasing apprehension.

Along with Bessie, he furthered our education beyond
herpetology by cooking up batches of fresh- caught local
crayfish and catfish (Figure 3) and treating my wife,
Joyce, and I to feasts on these. Ken relished both: a taste
brought north from his early upbringing in the Missis-
sippi drainage. But I missed out on his sausage-making,

2014

Cook: A TRIBUTE TO KENNETH WILLIAM ST! WART, 1936-201] 87

FIGURE 3. Ken Stewart and Channel Catfish (/cta/urus punctatus) on the Red River. Photo: Stewart family.

for which Pat Gregory vividly recalls a session : “Ken
attached 20 feet (or so) of sheep intestine to a (manual,
I think) meat grinder, into which he put his previously
prepared mix of sausage meat and seasoning, etc. As
the grinder worked away, the sausage mix was forced
into the intestine, so Ken had to quickly move to the
output end of the machine to twist the lengthening coil
here and there to form sausage links. The links were all
of different lengths and the whole chain seemed to
slowly wind its way around the kitchen like some in-
vading organism.”

Cas Lindsey has summed up Ken’s contributions:
“Ken was a man with a wide range of interests, which
he pursued with impressive expertise. Amongst his col-

leagues and his students he was the accepted source for

astonishingly well-informed opinion on almost any
topic. He was as versatile in his skills as in his written
knowledge; in subtropical seas he was a highly com-
petent scuba diver; in subarctic (Winnipeg) climes he
reared husky dogs. He was a mine of information on
the academic side of ichthyology, but he was also a
skilled and enthusiastic angler. He loved hunting as
well as fishing, and he knew a great deal about guns.

(He even constructed a firing range in the basement of

his Winnipeg home.) Despite his great store of knowl-
edge, he was singularly modest and uncompetitive.
Disinterested in self-aggrandizement, Ken preferred
teaching to writing. He did not often publish through-

out most of his teaching career. But fortunately in 2004
he more than made up for a slow start, and poured a
cornucopia of experience into the authoritative and
attractive volume The Freshwater Fishes of Manitoba
(K. W. Stewart and D. A. Watkinson, University of
Manitoba Press) (Figure 4). Although this may not be
the best way to get NSERC [Natural Sciences and Engi-
neering Research Council of Canada] grants, the book
demonstrates there is much to be said for writing your
great book after you have built up years of experience,
rather than the other way around.”

Ken and I remained in touch through correspondence
and during occasional visits Ken made to Ottawa. I often
called on Ken for perceptive and constructive reviews
of papers on fish, reptiles, and amphibians when I served
as chair of the Committee on the Status of Endangered
Wildlife in Canada (COSEWIC) Subcommittee on Am-
phibians and Reptiles and as editor of The Canadian
Field-Naturalist.

Ken is survived by his wife, Bessie, of Winnipeg,
daughters Kathy (Guy) Plett of Winnipeg and Anne
(Rob) Moniuk of Oakville, Ontario, and son John
Stewart of Vancouver, British Columbia. Ken also
leaves grandchildren Elizabeth (Don Ross) Plett, Guy
Plett, Wendy Moniuk, and Victor Moniuk, and great-
grandchildren William and Cameron Plett and Alyssa
and Ella Ross. A memorial was held in his honour on
8 October 2011 at the University of Manitoba.

88 THE CANADIAN FIELD-NATURALIST

Vol. 128

FIGURE 4. Ken Stewart (right) and coauthor D. A. Watkinson at the media book launch for The Freshwater Fishes of Mani-

toba in 2004. Photo: University of Manitoba Press.

Acknowledgements

An obituary appeared in the Winnipeg Free Press,
6 July 2011. Appreciation is owed to C. C. Lindsey,
Gavin Hanke, Patrick Gregory, Doug Watkinson, and
Beverly Horn for their recollections. Darren Gillis, at
the University of Manitoba, and Ken’s daughters Anne
Moniuk and Kathy Plett contributed photographs and
further information. D. A. Watkinson advised on con-
tacts and he and Bruce McCulloch provided additional
documentation for photographs.

Bibliography of Kenneth William Stewart

Publications

Stewart, K. W. 1962. Observations of the morphology and
optical properties of the adipose eyelid of fishes. Journal of
the Fisheries Research Board of Canada 19(6): 1161-1162.

Peden, A., and K. W. Stewart. 1964. Extension of the known
range of the antherinid fish Atherinops affinis. Copeia
1964(1): 239-240.

Stewart, K. W., and C. B. Levin. 1968. A method of obtain-
ing permanent dry mounted chromosome preparations from

fish. Journal of the Fisheries Research Board of Canada
25(5): 1091-1093.

Lane, E. D., and K. W. Stewart. 1968. A revision of the genus
Hoplunnis Kaup (Apodes, Muraenesocidae), with a descrip-
tion of a new species. Contributions in Marine Science
(Marine Science Institute, University of Texas, Port Aransas,
Texas) 1968 (13): 51—64.

Stewart, K. W., and C. C. Lindsey. 1969. First specimens of
the stonecat, Noturus flavus, from the Hudson Bay drain-

age. Journal of the Fisheries Research Board of Canada

27(1): 170-172.

Stewart, K. W., and F. R. Cook. 1970. [Abstract] Distribu-
tion of tree frogs of the Hy/a versicolor complex in Man-
itoba. Pages 27-28 in American Society of Ichthyologists
and Herpetologists Fifteenth Annual Meeting Abstracts,
New Orleans, Louisiana, 26-30 March 1970. Tulane Uni-
versity, New Orleans, Louisiana.

Aleksiuk, M., and K. W. Stewart. 1971. Seasonal changes in
the body composition of the garter snake ( Thamnophis sir-
talis parietalis) at northern latitudes. Ecology 52(3): 485
490).

2014 Cook: A TRIBUTE TO KENNETH WILLIAM STEWART, 1936-2011 &9

Gregory, P. T., and K. W. Stewart. 1975, Long distance dis-
persal and feeding strategy of the red-sided garter snake
(Thamnophis sirtalis parietalis) in the Interlake of Mani-
toba. Canadian Journal of Zoology 53(3); 238-245.

Stewart, K. W., and C. C. Lindsey. 1983. Postglacial disper-
sal of lower vertebrates in the Glacial Lake Agassiz region.
Pages 391-419 in Glacial Lake Agassiz. Edited by J. T.
Clayton and L. Clayton. Geological Association of Canada
Special Paper 26. Geological Association of Canada, c/o
Memorial University of Newfoundland, St. John’s, New-
foundland and Labrador.

Stewart, K. W. 1984. Maintenance of captive reptiles. Jn Care
and Use of Experimental Animals: A Guide for Canada,
Vol. Il. Edited by F. H. Flowers and J. P. W. Gillman.
Canadian Council on Animal Care, Ottawa, Ontario.

Stewart, K. W. 1984. Maintenance of captive amphibians.
In Care and Use of Experimental Animals: A Guide for
Canada, Vol. Il. Edited by F. H. Flowers and J. P. W. Gill-
man. Canadian Council on Animal Care, Ottawa, Ontario.

Stewart, K. W., I. M. Suthers, and L. K. Leavesley. 1985,
New fish distribution records in Manitoba and the role of
man-made interconnection between two drainages as an
avenue of dispersal. Canadian Field-Naturalist 99(3): 317—
326.

Hubicht, S. M., W. G. Franzin, and K. W. Stewart. 1988.
New distributional records for the minnows Hybognathus
hankinsoni, Phoxinus eos, and P. neogaeus in Manitoba.
Canadian Field-Naturalist 102(3): 475-484.

Stewart, K. W. 1988. First collections of the weed shiner,
Notropis texanus, in Canada. Canadian Field-Naturalist
102(4): 657-660.

Stewart, K. W., and C. Lindsey. 1990. Report to Fort Whyte
Outdoor Education Centre on Fish Species Composition
and Management Recommendations for the Fort Whyte
Ponds. August, 1990.

Stewart, K. W. 1990. Preliminary Assessment of the Impact
of the Dam on the Red River Proposed in the Pembina
Triangle Water Supply Study on Fish Communities in the
Red River. Submitted to the Manitoba Department of Nat-
ural Resources and to the Manitoba Environmental Coun-
cil, October 1990.

Campbell, K. B, A. J. Derksen, and K. W. Stewart. 1991.
First specimens of the Rainbow Smelt, Osmerus mordax,
from Lake Winnipeg. Canadian Field-Naturalist 105(4):
568-570.

Hanke, G. F., and K. W. Stewart. 1994. Evidence for recent
dispersal of fishes in Lake Winnipeg. Proceedings of the
North Dakota Water Quality Symposium, Part III: 133-149.

Duncan, J., E. Bredin, G. Hanke, W. Koonz, R. Larche, K.
Leavesley, W. Preston, D. Ross, C. Scott, D. Stardom,
K. Stewart, and P. Taylor. 1994. Estimated status of Mani-
toba amphibians based on criteria used by the Nature Con-
servancy’s Conservation Data Center Network. Proceed-
ings of the 4th Meeting of the Task Force on Declining
Amphibian Populations in Canada, Part IV: 93-103.

Hanke, G. F., K. W. Stewart, and G. A. Lammers. 1996.
Eastmanosteus lundarensis, a new arthrodire from the Elm
Point Formation of Manitoba. Journal of Vertebrate Pale-
ontology 16(4): 606-616.

Hanke, G. F., K. W. Stewart, and G. A. Lammers. 1996.
Squamatognathus steeprockensis, a new inferognathal from
the Elm Point Formation of Manitoba. Journal of Vertebrate
Paleontology 16(4): 617-622.

McCulloch, B. R., and K. W. Stewart. 1998. Range exten-
sions and new locality records for the stonecat, Noturus

flavus, in Manitoba: evidence for a recent natural invasion.
Canadian Field-Naturalist |12(2): 217-224.

Steinhilber, M., and K. W. Stewart. 1999. The species diver-
sity of fishes in the Prairies Ecozone of Canada. Report
Commissioned by Environment Canada.

Stewart, K. W., W. G. Franzin, B. R. McCulloch, and G. F.
Hanke. 2001. Selected case histories of fish species inva-
sions into the Nelson River system in Canada. Pages 63-81
in Science and Policy: Interbasin Transfer of Aquatic Bio-
ta. Edited by J. A. Leitch and M. J. Tenamoc. Institute for
Regional Studies, North Dakota State University, Fargo,
North Dakota.

Franzin, W. G., K. W. Stewart, G. F. Hanke, and L.
Heuring. 2003. The fish and fisheries of Lake Winnipeg:
the first 100 years. Canadian Technical Report of Fisheries
and Aquatic Sciences 2398. Central and Arctic Region,
Department of Fisheries and Oceans, Winnipeg, Manitoba.
53 pages.

Stewart, K. W., and D. A. Watkinson. 2004. The Freshwater
Fishes of Manitoba. University of Manitoba Press, Win-
nipeg, Manitoba. 276 pages.

Theses supervised

Schweitzer, R. D. 1968. Evidence of two sympatric forms of
ciscoes (subgenus Leucichthyes) in Cedar Lake. M.Sc. the-
sis, University of Manitoba, Winnipeg, Manitoba.

Case, B. E. 1970. An ecological study of the tadpole madtom,
Noturus gyrinus (Mitchell), with special reference to move-
ments and population fluctuations. M.Sc. thesis, Univer-
sity of Manitoba, Winnipeg, Manitoba.

Gregory, P. T. 1971. Overwintering populations and homing
of the red-sided garter snake (Thamnophis sirtalis pari-
etalis) at a northern latitude. M.Sc. thesis, University of
Manitoba, Winnipeg, Manitoba.

Vincent, T. K. 1971. Resistance to cold stress in the red-sided
garter snake (Thamnophis sirtalis parietalis) at a northern
latitude. M.Sc. thesis, University of Manitoba, Winnipeg,
Manitoba.

Levin, C. B. 1972. Karyotypic analysis of somatic chromo-
somes of Noturus gyrinus (Mitchell) (Ictaluridae: Teleos-
tei). M.Sc. thesis, University of Manitoba, Winnipeg, Man-
itoba.

Gregory, P. T. 1973. Life history parameters of a population
of red-sided garter snakes (Thamnophis sirtalis parietalis)
adapted to a rigorous and fluctuating environment. Ph.D.
thesis, Univiversity of Manitoba, Winnipeg, Manitoba.

Clarke, R. MeV. 1973. Systematics of ciscoes (Coregonidae)
in central Canada. Ph.D. thesis, University of Manitoba,
Winnipeg, Manitoba.

Hart, D. R. 1975. Quantitative niche comparison of the west-
ern plains garter snake (Thamnophis radix haydeni) and the
red-sided garter snake (7hamnophis sirtalis parietalis) in
allopatric and sympatric regions of Manitoba’s Interlake
District. M.Sc. thesis, University of Manitoba, Winnipeg,
Manitoba.

Eddy, S. B. 1976. Population ecology of the leopard frog,
Rana pipiens pipiens Schreiber, at Delta Marsh, Manitoba.
M.Sc. thesis, University of Manitoba, Winnipeg, Manitoba.

Cook, F. R. 1978. An analysis of toads of the Bufo ameri-
canus group in a contact zone in central North America,
Ph.D. thesis, University of Manitoba, Winnipeg, Manitoba.

Yaremchuk, G. 1981. The effects of exposure to predation on
mean vertebral and fin ray counts in the fathead minnow
(Pimephales promelas). M.Sc. thesis, University of Man-
itoba, Winnipeg, Manitoba.

90 THE CANADIAN FIELD-NATURALIST

Guinn, B. 1982. A morphological and ecological comparison
of round whitefish (Prosopium cylindraceum) and moun-
tain whitefish (P. williamsoni) in allopatric and sympatric
areas in northwestern Canada. M.Sc. thesis, University of
Manitoba, Winnipeg, Manitoba.

Day, A. C. 1982. Ecology of burbot, Loa Jota and lake trout,
Salvelinus namaycush in Lake Athapapuskow, Manitoba.
M.Se. thesis, University of Manitoba, Winnipeg, Manitoba.

Leavesley, L. Ik. 1987. Ecological study of the western hog-
nose snake, Heferodon nasicus nasicus in Spruce Woods
Provincial Park, Manitoba. M.Sc. thesis, University of
Manitoba, Winnipeg, Manitoba.

Wallace, R. 1989. Multivariate analysis of habitat parameters
affecting occurrence, abundance and productivity of lake
trout, Salvelinus namaycush, in lakes in northern Saskat-
chewan. M.Sc. thesis, University of Manitoba, Winnipeg,
Manitoba.

Harbicht, S. A. 1990. Biology of the shorthead redhorse,
Moxostoma macrolepidotum in Lake Dauphin, Manitoba.
M.Sc. thesis, University of Manitoba, Winnipeg, Manitoba.

Horn, B. M. 1993. Geographic variation of populations of
Mimic Shiners, Notropis volucellus (Cope 1865) and Sand
Shiners, Notropis stramineus stramineus (Cope 1865)
(Cyprinidae), in Manitoba and southern Ontario. M.Sc.
thesis, University of Manitoba, Winnipeg, Manitoba.

Rowes, K. D. 1994. Temporal and spatial distribution of
pelagic larval fishes of Dauphin Lake, Manitoba. M.Sc.
thesis, University of Manitoba, Winnipeg, Manitoba.

McCulloch, B. 1994. Dispersal of the Stonecat (Noturus
flavus) in Manitoba and its interactions with resident fish
species. M.Sc. thesis, University of Manitoba, Winnipeg,
Manitoba.

Ogutu-Ohwayo, R. 1994. Adjustments in fish stocks and in
life history characteristics of the Nile Perch, Lates niloticus
L. in Lakes Victoria, Kyoga and Nabugabo. Ph.D. thesis,
University of Manitoba, Winnipeg, Manitoba.

Doolgindachbaporn, S. 1994. Development of optimal rear-
ing and culturing systems for the Climbing Perch, Anabas
testudineus (Bloch) (Perciformes, Anabantidae). Ph.D. the-
sis, University of Manitoba, Winnipeg, Manitoba.

Johnson, J. D. 1995. Description and comparison of two pop-
ulations of Saffron Cod (Eleginus gracilis) from western
Canadian Arctic coastal waters. M.Sc. thesis, University of
Manitoba, Winnipeg, Manitoba.

Tyson, J. D. 1996. The effect of thermal effluent on overwin-
tering Channel Catfish (/ctalurus punctatus) in the lower
Red River, Manitoba. M.Sc. thesis, University of Manitoba,
Winnipeg, Manitoba.

Hanke, G. 1996. A survey of the fishes of Lake Winnipeg and
interactions of the introduced White Bass with the native

Vol. 128

ichthyofauna of Hudson Bay drainage: with emphasis on
young-of-the-year in nearshore environments. M.Sc. thesis,
University of Manitoba, Winnipeg, Manitoba.

Stephenson, S. A. 1996. An analysis of meristic, morphome-
tric and biochemical variation in the Johnny Darter, Etheo-
stoma nigrum Rafinesque, in the northern United States
and Canada with biogeographic and systematic consider-
ations. M.Sc. thesis, University of Manitoba, Winnipeg,
Manitoba.

Kristofferson, A. H. 2003. Identification of Arctic char stocks
in the Cambridge Bay area, Nunavut Territory and evidence
of stock mixing during overwintering. Ph.D. thesis, Uni-
versity of Manitoba, Winnipeg, Manitoba.

Publications arising from graduate student research

that Stewart supervised or advised

(in chronological order)

Hlynka, L. 1969. The biology of Rana pipiens Schreber and
Bufo hemiophrys Cope in the Delta marshes. University of
Manitoba Delta Marsh Field Station report.

Hlynka, L. J. 1970. Helminths in Rana pipiens Schreber, and
Bufo hemiophrys Cope, from the Delta Marshes, Manitoba.
M.Sc. thesis, Department of Zoology, University of Mani-
toba. 110 pages. (Supervisor, G. Lubinsky)

Aleksiuk, M., and P. T. Gregory. 1974. Regulation of sea-
sonal mating behaviour in Thamnophis sirtalis parietalis.
Copeia 1974(3): 681-89.

Gregory, P. T. 1974. Patterns of spring emergence of the red-
sided garter snake (Thamnophis sirtalis parietalis) in the
Interlake region of Manitoba. Canadian Journal of Zoology
52(8): 1063-69.

Gregory, P. T. 1977. Life-History Parameters of the Red-
sided Garter Snake (Thamnophis sirtalis parietalis) in an
Extreme Environment, the Interlake Region of Manitoba.
National Museums of Canada, Ottawa, Ontario. 44 pages.

Hart, D. R. 1979. Niche relationships of Thamnophis radix
haydeni and Thamnophis sirtalis parietalis in the Interlake
District of Manitoba. Tulane Studies in Zoology and Botany
21: 125-140.

Hart, D. R. 1982. Growth of Painted Turtles, Chrysemys picta,
in Manitoba and Louisiana. Canadian Field-Naturalist 96:
127-131.

Cook, F. R. 1983. An analysis of toads of the Bufo americanus
group in a contact zone in central northern North America.
National Museum of Natural Sciences, National Museums
of Canada, Ottawa, Ontario. 89 pages.

Received 27 October 2013
Accepted 21 February 2014

Book Reviews

c qi j %% . > ore ec j j fi b j
ata Review Editor’s Note: We are continuing to use the current currency codes. Thus Canadian dollars are CAD, U.S.
dollars are USD, Euros are EUR, China Yuan Remimbi are CNY, Australian dollars are AUD and so on.

ZOOLOGY

Into the Night: Tales of Nocturnal Wildlife Expeditions

Edited by Rick A. Adams. 2013. University of Colorado Press, 5589 Arapahoe Avenue, Suite 206C, Boulder, Colorado 80303

USA. 194 pages, 26.95 USD, Cloth.

Except in urban settings, most people avoid the
night. But humans are not typical mammals, because
most species in this class of animals are nocturnal.
However, the eight contributors to this edited book on
evening wildlife experiences are not your average per-
son, they are biologists and naturalists fascinated with
what goes on when the sun goes down.

The first story recounts the camping adventures of
naturalist Stephen Jones in the Sandhill prairies of
Nebraska. He studies owls, which has brought him in
close encounters with these nocturnal birds. They have
visited him in the wild on numerous occasions over the
past three decades and have been his personal window
on nature. The long-eared owl and its survival has been
a particularly poignant species for him and 1s a harbin-
ger for the progress of civilization.

Dr. Frank Bonaccorso is a wildlife biologist who
specializes on studying bats. His story focuses on re-
search in Kruger National Park in South Africa and is
interspersed with cat and mouse games of avoiding
large mammals such as lions and elephants. He puts
little transmitters on fruit bats caught in his nets set near
big sycamore fig trees — a keystone species that pro-
vides food and shelter for many different animals in an
otherwise predominantly open grassland region. It turns
out that the only mammal capable of self-powered
flight eats a lot of fruits but also is a good disperser of
seeds that pass through the digestive tract and are defe-
cated while flying. So I suppose one more thing to
watch out for during the night but not a potentially life
threatening danger as other things that lurk in the night.

Although not all exclusively nocturnal, the tales of
diving in the Galapagos by Christina Allen are certain-
ly dark and mysterious, especially when sharks come
into play. But there are two incidents she shares of night
time experiences. One is being mesmerized by a mass
of bioluminescent marine organisms seen during the
first evening on her research vessel that is investigating
changes to the island archipelago since Darwin’s time.
The other is a night dive to find the bizarre and aptly
named red-lipped batfish flapping on the ocean floor.

The fourth chapter is written by the editor of the
book, Dr. Rick Adams, who is a biology professor at
the University of Northern Colorado and a bat re-
searcher. Most of his stories take place in the nearby
Rocky Mountains, up to around 4,000 metres in ele-
vation, as he studies the distribution, abundance, and
resource use of bats. Some experiences were eerie, such
as ghostly images lurking in the shadows, but others
were humorous, such as the trailer-eating porcupine
that wakes up his field team in the middle of the night.

The next writer takes us to Sulawesi and macaque
monkeys, but also reveals her distain for mosquitoes.
Dr. Ann Kohlhaas is a professor of biology at Califor-
nia State University in Stanislaus but dreams of her
fieldwork in southeast Asia. But it’s more than just pri-
mates, as she reminisces of wildlife encounters with
bats, rats, and snakes. And there are also potentially
threatening encounters with earthquakes and malaria.

More tropical adventures are told by Dr. Lee Dyer,
a biologist at the University of Nevada in Reno. But
these ones take place on the other side of the world in
Costa Rica and Ecuador. They are equally as odd with
stories of fellow field biologists getting high one night
on the local flora and stripping naked before walking
into a web of thousands of spiders. Another time after
a long night’s work, the author gets bit by “a big red and
hairy spider” on his private parts and blood is drawn.
It ends happily when he wakes up, after stumbling back
to his cabin and passing out, to see the stars are still
shining.

Not to be left out, a desert encounter with a rattle-
snake is told by Dr. James Halfpenny while studying
packrats that have been dusted with a fluorescent pow-
der enabling them to be tracked at night to find their
middens and hopefully also the beetles they eat that are
good indicators of historical changes in climate. But he
also tells of snowy adventures as far north as the Arctic,
where “nocturnal” work and avoidance of polar bears
takes place in the “land of the midnight sun”.

The final chapter is left to bat biologist Dr. Scott
Pedersen at South Dakota State University in Brook-
ings. He gets to do fieldwork on the Caribbean island of

9]

92 THE CANADIAN FIELD-NATURALIST

Montserrat. But it is not all paradise, as he has to deal
with erupting volcanoes and pooping bats at night. How-
ever, he does survive to give us a recipe for “mountain

Owls

Vol. 128

chicken stew”, which is actually a big frog for you
foodies out there.
BURTON K. LIM

Department of Natural History, Royal Ontario Museum,
100 Queen’s Park, Toronto, Ontario MSS 2C6 Canada

By Marianne Taylor. 2012. Cornell University Press (Comstock Publishing Associates), Box 6525, 750 Cascadilla Street,
Ithaca, New York 14851-6525 USA. 224 pages, 35.00 USD, Cloth.

In the introduction to Ow/s, British illustrator, pho-
tographer, and author Marianne Taylor speculates on
the reasons why these birds capture the human imag-
ination so powerfully. She points out that since eye
contact is critical to human relations, we are drawn to
animals who can return our gaze with two forward-
pointing eyes. Most birds, she stresses, have side-
mounted eyes for a broader field of vision. She con-
cludes that “owls can truly look at us as we look at
them.”

The first section of her book, consisting of nine
chapters describing owl diversity and natural history,
provides insight into how owls see, and explains addi-
tional owl senses, attributes and capacities, hunting
behaviour and techniques, habitat, breeding biology,
conservation issues, and human-owl relations. The sec-
ond section of the book offers profiles of 41 individual
owl species from around the world.

A discussion of the cognitive capacities of owls early
in the book adds significant interest. Owl intelligence
is apparently little studied, but research has discovered
that Barn Owls form extraordinarily detailed auditory
maps in their brains, and that the birds have remark-
ably accurate memories for sounds and their locations.
Berndt Heinrich’s stories of a hand-reared Great Horned
Owl learning to wake him for early meals, gently take
food from his hands, and play with inanimate objects
for hours provide anecdotal evidence of owl intelli-
gence.

Another intriguing fact, described in a later chapter,
further demonstrates owl intelligence, specifically the
capacity of the birds to solve problems. Taylor writes
about the novel way screech owls deal with flies and
other small animals attracted to waste in the birds’ nest-
ing cavities. The owls bring live slender blind snakes
into the nest to eat the flies, usually leaving the snakes
uneaten in return for their services — a fascinating rela-
tionship.

Taylor devotes an entire chapter to another relation-
ship, with humans, pointing out that since owls are dis-
tributed so widely around the world, most countries
have developed distinct owl-related myths and legends.
These include owl-like gods and goddesses, owl com-

panions to certain deities, and sundry other links to the
supernatural. Some owl symbolism has grown out of
the latter connections — for example, early Christian
associations of owls with evil, resulting in the unfortu-
nate persecution of owls in related cultures. In other
parts of the world, owls — in particular their vocaliza-
tions — can be either positive or negative, depending on
the type or frequency of the sound

Taylor also writes about owls in literature, televi-
sion, and film — for example, Winnie-the-Pooh books,
Sesame Street, and the Harry Potter series — and about
owls in falconry, where they are valued despite (or per-
haps because of) the fact that they are known for their
reluctance to relinquish prey to their handlers.

The second part of Owls starts with a very clear
overview of the various owl genera, followed by spe-
cies profiles describing the bird’s range, evolution,
relationships, physical features, geographical variation,
movements and migration, voice, habitat, behaviour,
hunting-diet, and status-conservation.

This part of the book introduced me to unfamiliar
owl species such as the Pharaoh Eagle Owl, a large bird
of arid and rocky landscapes in north-west Africa, where
it typically nests among rocky structures, including the
pyramids. I learned that the Collared Owlet of south-
east Asia’s evergreen forests is the continent’s small-
est species, but a powerful predator that eats birds up
to its own size or even larger — likely the reason it is
fiercely mobbed by forest birds when discovered. I
was intrigued to read about the fish owls of Asia, who
hunt at the water’s edge, and whose diet consists main-
ly of aquatic animals; unfortunately some of these owls
are not well known, while the Blakiston’s Fish Owl of
eastern Russia, China, and Japan (north Hokkaiso Is-
land) is considered endangered.

Owls is a large and beautiful work of photography
and writing. Naturalists intrigued by the mysterious
lives and uncanny abilities of owls will find the book.
with its comprehensive general introduction to the birds,
intriguing facts about individual species, and plentiful
photographs a worthwhile addition to their libraries.

RENATE SANDER-REGIER
3, 11th Line, Bristol, Quebec, Canada

2014

Book REVIEWS 93

A Pocket Guide to Salamanders of Pennsylvania

By Walter E. Meshaka, Jr., and Joseph T. Collins. 2012. The Pennsylvania Heritage Foundation, 225 State Street, Suite 302,

Harrisburg, Pennsylvania 17101 USA. 52 pages, 4.71 USD.

This booklet is a companion to 4 Pocket Guide to
Lizards and Turtles of Pennsylvania published the
same year (reviewed in The Canadian Field-Naturalist
126(4): 345-346). It is an identical 15.2 by 9.5 in height
and width and only marginally thicker with 12 more
pages. The authors are also identical in the two guides
but this one is dedicated to Collins, “Whose passion
for herpetology was equaled only by his love for shar-
ing it’, who had died of a heart attack at 72 before it
was printed. The sponsors are the same: The State
Museum of Pennsylvania, Pennsylvania Historical &
Museum Commission; Wild Resource Conservation
Program, Pennsylvania Department of Conservation &
Natural Resources; Dickinson College Biology Depart-
ment; Forgotten Friend Reptile Sanctuary; Friends of
Wildwood Lake Nature Center, Inc.; Liberty Environ-
mental, Inc.; Powdermill Nature Reserve; Shippens-
burg University; and The Center for North American
Herpetology.

Like the earlier guide to lizards and turtles, this guide
opens with an introduction. This one points out the pro-
tected species in the state, officially two endangered,
and one threatened. Mentioned as well is the concern
expressed by some conservationists over four addition-
al ones. Following this are acknowledgements.

Accounts for the 23 salamander species recorded in
Pennsylvania occupy two facing pages for each. On the
left are presented the English name, scientific name,
total length, and brief highlights on range (in Pennsyl-
vania), habitat, habits, breeding, diet, and conservation
considerations. On the right are two illustrations of
transformed individuals. For the Eastern Newt both
mature adult and immature terrestrial eft stages are
depicted. Also on the illustration page is a minute range
map with counties in which the species is present in
green based on information from the Pennsylvania Her-
petological Atlas web site and Amphibians and Reptiles
of Pennsylvania and the northeast by Arthur C. Hulse.
C. J. McCoy and Ellen J. Censky. The distributions vary
from covering the whole state to barely entering it.

The guide concludes with one photograph of a Spot-
ted Salamander egg mass, another of spermatophores
of the same species, and two contrasting the pond larva

Snapper

of the Spotted Salamander with the stream larva of the
Spring Salamander, and a checklist of species in the
five families represented. A page gives sources of fur-
ther information on herpetology on websites and pub-
lications relevant to Pennsylvania and comments on
herpetoculture which stresses current state regulations.
The inside back cover has information on the authors
and a quote from Af the Planning Commission by Bar-
bara Meyn in The Abalone Heart, 1988: “I forgot to tell
them about the salamanders, dark as chocolate, torpid
with cold, they move ... about this time every year, how
easy it is to drive right over them if you are unaware.”

Overall, this publication is less than successful for
species identification partly due to the darkness of many
photographs but primarily because colour and pattern
alone are not sufficient to distinguish between some
similar forms. Examples are Jeffersonian and Blue-
spotted salamanders, and the many dark or red wood-
land or stream assemblages in Plethodontidae. It
should, however, reliably serve for distinguishing the
other three Ambystomatidae ([ Yellow-] Spotted Sala-
mander, and Marbled [not recorded in Canada] and
Tiger [recorded only once in eastern Canada] sala-
manders), and the single Protidae (Mudpuppy), Sala-
mandridae (Eastern Newt) and Cryptobranchidae (Hell-
bender) [the latter not occurring in Canada]. Fortunately
for use in eastern Canada, 10 of the 15 Pletodontidae do
not range this far north. The particularly confusing pairs
that do occur in Canada are the plethodontid Northern
and Allegany dusky salamanders and the ambystom-
atid Jeffersonian and Blue-spotted salamanders. A po-
tential source of confusion for Canadian users is that
the Red-backed Salamander is represented in the pock-
et guide only by the red-back phase (red dorsum and
gray sides) whereas lead-back (lacking red) and ery-
thristic (all red) phases which might be confused with
a southern species, as well as well as other colour vari-
ants, occur in Canada with the lead-back reaching
100% in some populations.

FRANCIS R. COOK

Canadian Museum of Nature, Ottawa, Ontario KIA 6P4
Canada

By Brian Kimberling. 2013. Pantheon Books, The Knopf Doubleday Group, 1745 Broadway, New York, New York 10019

USA. 224 pages, 28.96 USD, Cloth.

A few months ago, I read a disquieting article in the
New York Times on declining bird populations written
by Brian Kimberling. While the article was interesting,
and well-written, and of course depressing, it also con-
tained a reference to a book by Kimberling entitled

Snapper. | was immediately on alert. A book about
“my” species? Was I being scooped? Or did this have
nothing to do with snapping turtles? | rushed to Ama-
zon, a few keystrokes south, and to my horror, yes it
was about snapping turtles, though apparently not en-

94 THE CANADIAN FIELD-NATURALIST

tirely. Nevertheless, a few more excited keystrokes
and in the next day’s mail there arrived a small book
with an attractive dust jacket decorated by colourful
Audubon-like drawings of birds, which turned out to
be Audubon drawings. Not a dust jacket one would
associate with snapping turtles. | hurriedly opened the
book to scan the Table of Contents. There was none,
but there were chapters and Snapper was the title of
chapter 2. I settled back in my Lazyboy and started on
chapter | to build up suspense and prejudice for chap-
ter 2. Thus began a wonderful read of many vignettes
by a most accomplished writer and, I surmise, natural-
ist, on the life and times of a budding biologist/ornithol-
ogist named Nathan Lochmueller. Although this is an
autobiographical novel, one cannot help feel it is far
from entirely fiction. The “snapper” takes up only one
chapter and it is obscure to me how the author chose
this title for the book. “A Biologist’s Adventures among
the Rednecks of Southern Indiana” would get more
attention and reflect the content better. Even “Ram-
blings about an Ordinary Youth” or “My Hopeless Love
Life while Studying Eagles” might be more attractive
to potential readers and still better reflect the story.

Lochmueller is a 20’s-year-old biologist, born, raised
and sort of maturing in southern Indiana. The book
chronicles bits and pieces of his life. Like Bob Dylan’s
recent “autobiography”, Chronicles, this book leaps
about with no respect for time and skitters among top-
ics and times without rhyme, reason or warning. To me
the writing style, the incorporation of a smattering of
interesting natural history, and the description of the
lonely, impoverished, slavish life of the underpaid,
overworked, dedicated and unappreciated neophyte
naturalist makes this a fascinating read. Indeed, I will
say without qualm, anyone who is not captivated by the
end of chapter 1, is no one I really need to know, and
vice versa.

I read the whole book in one short day. I like short
books. I liked this one enough that I have since read it
more slowly and deliciously twice more. One would
like to say it is slightly reminiscent of Stephen Lea-
cock, but sharper, more poignant and critical, edging
toward Mark Twain, or even Jon Stewart of the Daily
Show. I like his preface quote, “While there is a lower
class, I am in it, while there is a criminal element, I am
of it, and while there is a soul in prison, I am not free”.
Or the opening sentence. “I got my job by accident.”

The book swerves and slips between amusing yet
awed descriptions of Nathan’s ornithologist boss, who
reminds me of many brilliant biologists I have known
and whom I found awesome, and amusing, but never
close to. Other memorable characters populate the nar-
rative including, his rogue aunt and uncle from Texas,
the Hoosier natives of redneck Indiana, the ivory tower
residents of Bloomington, home of Indiana University,
and a few feckless friends. But the narrative always
returns to Lola, the Maggie May (a la Rod Stewart)
love of his life, or at least youth. No male, regardless

Vol. 128

of his sexual orientation can read this book, and not
admit he has known a Lola. Especially if that male was
somewhat aimless, and spending six days a week scour-
ing woodlands to map the frail lives of obscure wildlife
in the undergrowth. The recent bird researchers in Al-
gonquin Park kept appearing in my picture of young
Nathan.

Consider his description of finding the nest of a
Kentucky Warbler; “It’s like staking out the girl’s show-
er block at summer camp. It can be done, but it takes
skill.” Oh, hello Hugo.

My problem in reviewing this book is that I want to
quote vignettes to reveal the clever humour, the wry
regret, the observer’s forgiveness, the blatant stabs, the
dazzling mix of arrogance and self-deprecation and the
pithy commentary on modern humans and our institu-
tions. And the touching and hilarious depiction of a lost
opportunity with a woman afflicted badly by Lyme’s
disease. Nathan attributes to her the description of the
Mississippi as “a vast national sow prone to rolling over
her young”. Nathan then pauses to refer to the ivory-
billed woodpecker as Elvis in feathers for both gaudy
plumage and regular sightings since declared extinct
in 1944. We ecologists all have our Elvis.

Nathan ends his saga of Dana the Lyme’s girl by
comparing her to his great love, Lola. Lola “belonged
to that air conditioned world, Dana understood the
squalid and menacing nature of things...the snap of
events, Lola’s independence was a vain inglorious
thing, and my own spotty affair with Lola struck me
as trivial, something that dropped out of my sleeve or
back pocket, probably not worth picking up.” Needless
to say, Lola was falling from grace at this point, but she
repeatedly reappears, although it is not clear whether
it is before or after the above conclusion.

Chapter ix is titled ‘Proof’. It describes how Nathan
marked a forest of trees by carving Lola’s name on
all those marked for felling by the US Forest Service
plus many others (to temporarily confuse the USFS, a
hopeless tactic I confess to using in my youth). I trust
that all we Luddite nature lovers have done similarly
hopeless things to thwart developers a la Edwin Albee.
Nathan then describes a court proceeding utterly remi-
niscent of my experiences before the Ontario Munici-
pal Board. In this case, a tribunal hears arguments from
both sides in a dispute (the cutting or not of trees in a
“nature reserve”). This proceeding before a tribunal
occurs , “when one side has no prospect of winning”.
Nathan recounts that the case failed because “it was
tried in Indiana, where science, education and Darwin
are all equally deplored”. The defendants were the US
Forest Service whose primary function is “to facili-
tate timber sales”, and the US Fish and Wildlife Serv-
ice which “truly reveres all woodland creatures provid-
ed they can be shot, trapped, hooked, netted or clubbed”.
At least we know where he stands. Nathan goes on to
describe how the lawyers for the defense systemati-
cally destroy his credibility and the affidavits of sev-

2014

eral leading ecologist and explains, ‘a lawsuit is nota
work of art, and its aims are to obfuscate what it cannot
denigrate until money is free to speak”. Such cynicism!

In the end, Nathan comes to some, for me, surpris-
ing conclusions, and the fearsome spectre of Darwin
rears its ubiquitous embrace. But earlier, our hero thinks
back to his days following avian domestic lives, recall-
ing his time, “wandering vast tracts of Indiana wood-
land and riverbank, taking orders from no one, chron-
icling the lives, births and deaths and domestic disputes
of forest songbirds for biology departments and gov-

The Crossley ID Guide to Britain and Ireland

BOOK REVIEWS 95

ernment agencies. | reveled, like ... John James Audu-
bon, two hundred years before me, in the same extraor-
dinary beauty and variety there — reduced every day, by
human encroachment, but resilient and resplendent
nevertheless”. I can’t think of a better summary of my
own life in the wilds of Ontario and I am sure this
echoes the experiences of many readers of the Cana-
dian-Field Naturalist. This is a book to be treasured.

RON BROOKS

221 Ferguson Street, Guelph, Ontario NIE 2Z1 Canada

By Richard Crossley, and Dominic Couzens. 2013. Princeton University Press, 41 William Street, Princeton, New Jersey,

08540-5237 USA. 304 pages, 27.95 USD, Paper.

I have to admit to a positive bias for Crossley’s
method of illustration. I know others find the crammed
pictures confusing and distracting. But you never see
a bird facing right against a plain white background.
Well maybe a Shorelark (Horned Lark) on fresh snow
is a possibility. Crossley takes computer-manipulated
photographs of birds, mounted on to a typical back-
ground. This illustration looks like a jigsaw puzzle,
with several examples of the species in question dotted
throughout the habitat. Most common species have a
full page, while the rarer birds have half or a quarter of
a page. The author does not follow the current taxo-
nomic order so, for example, all the waterbirds are in
sequence. This is a very sensible approach and far more
valuable in the field.

My copy arrived a week before | left for England.
So I avidly used it to review species I was likely to see.
I reminded myself of the difference between Rooks and
Carrion Crows. I read through the key points separating
Willow and Marsh Tits. I read that Marsh Tit is “One
of the worst named birds ...” as it prefers deciduous
damp woodland — not marshes. The Willow Tit likes
scrubby bushes around lakes and that does include wil-
lows.

Rather than being put off by the illustrations I found
them particularly charming. Swallows (Barn Swallows
to us) are shown in a typical farmyard; bright birds
against cold, grey stone. There is a quaint flower-filled
village as the background for Blackbird and typical
churchyard for the iconic (Eurasian) Robin. While the
Marsh Tit flits in its woodland but the Bearded Tit is
in a marsh.

In Britain I visited Greenwich, Dulwich, Hampton
court, Kensington Gardens and Hampton court. I was
amazed to realise that after four days I had made sev-

eral sightings of Rose-ringed Parakeets and yet I had
not seen a single House Sparrow. Crossley gives the
current estimate of populations where available. For
the parakeets he quotes 8,600 pairs, while the sparrows
rank at 5.3 million pairs. I eventually saw 20 House
Sparrows near Lincoln.

There are other species included that add to the
sense of the habitat and give a more rounded idea of
the species environment. For example, there is a man
raising his arm in defence against Great Skuas. A Badg-
er peeks out from the trees holding Tawny Owls. A
Eurasian Red Squirrel, that shares the same limited dis-
tribution, adds to the Crested Tit page.

Last week a friend sent a photo of a flock of Com-
mon Redshank with their bright coral “shanks” with
a slightly smaller yellow-legged bird in the centre. It
was one of the over 250 records for Lesser Yellowlegs
for the UK. This species does not make it into this book.
It covers about 300 out of 596 species on the official
British record. Regularly occurring rarities that arrive
every year, but in very small numbers are covered, but
the more sporadic vagrants are not. As such this guide
is more useful to new birders and occasional visitors.
Indeed the introduction provides an excellent approach
for the new birdwatcher. There are many practical
examples of what to do in the field to get the best out
of this guide.

So I recommend this book as a first guide for begin-
ners. You can move on to a more comprehensive guide
after a couple of year’s experience. Visitors will get
an introduction to both the birds and the countryside
of the British Isles.

Roy JOHN

2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada

96 THE CANADIAN FIELD-NATURALIST

The Warbler Guide

Vol. 128

By Tom Stephenson, and Scott Whittle. 2013. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-

5237 USA. 514 pages, 29.95, Paper.

Warblers are the crown jewels of the Americas avi-
fauna. Exquisitely plumaged in spring, many assume
cryptic plumages in fall, morphing from among the
most beautiful of birds to tricky identification chal-
lenges. Indeed, for many Canadian observers learning
fall warblers is their first great birding milestone. Add
to this an array of songs and you have the complete
avian package. As an added bonus, warblers are a key
part of our summer ecosystem; and witnessing mixed
species migration fallout is among the greatest spec-
tacles that await the Canadian birder.

Warblers are well treated in standard field guides,
which are more than adequate for most identifications.
However, due to their intrinsic beauty and the interest
they invoke, several family monographs have been
produced, of which this is the most recent. At over 500
pages it is a stay-at-home reference work, not a field
guide. It starts with an extensive introductory section
including an easy to follow “topographic tour’, tips on
what to look for, and an extensive segment on how to
use sonograms. Indeed, a unique feature of this book
is its extensive treatment of vocalizations. Every species
has a comprehensive collection of sonograms, and there
are stand alone sections visually comparing the songs,
chip notes and flight calls of similar species. Not every-
body “gets” sonograms, but if you are one of those
who does, this will be a selling point. There is also an
extensive and well-organized photographic “quick find-
er guide”. Particularly useful for suffers from “warbler
neck” are plates portraying undertail patterns. The end
pieces include a fun test quiz and an intriguing section
on hybrids.

Most of the book consists of species accounts. Every
regularly occurring species in North America is exhaus-
tively treated, with multiple photographs of all distinct
plumages, close ups of key features, useful studies of
comparison species, a treatment of aging and sexing,
accurate, easy to read range maps, and sonograms. To

provide a random example: the species account for
Canada Warbler has 37 photographs, nine sonograms
and two range maps (one each for fall and spring migra-
tion). Additional photographs and sonograms of the
species are found elsewhere in the book. On the other
hand, the text is minimalist, consisting mostly of con-
cise descriptions of the key features illustrated in each
figure. Rarely occurring species are afforded less space
but are more extensively treated than in a standard field
guide.

The audience for this book are more experienced
observers; neophytes will be better served by standard
field guides. The current work will help you figure out
if that Common Yellowthroat you are saw was a first
winter male or female, sort out American Redstart ag-
ing and sexing, or help with that tricky Oporornis iden-
tification. I “test drove” the book several times this fall,
checking it against birds seen that day or testing to
see if it was up to the challenge of identifying “mystery
bird” photographs posted to various web groups, which
it was, often using second order identification clues
beyond those included in standard works.

How does it stack up? The most important compari-
son is “A Field Guide to Warblers of North America”
by Jon Dunn and Kimball Garrett. In effect the two
works are complementary, Dunn and Garrett has a more
textual approach with detailed written descriptions
and greater detail on ecology and range, whereas the
current work is unabashedly visual. Warbler offician-
dos will want to own both. The bottom line? Get this
book if you are an experienced observer looking to take
your warbling to the next level. Like me, you will prob-
ably find yourself turning to it from time to time, either
to verify particularly tough identifications or to admire
its many fine photographs.

MARK GAWN
25 Forsyth Lane, Ottawa, ON, Canada

Yellowstone Wildlife: Ecology and Natural History of the Greater Yellowstone Ecosystem

By Paul A. Johnsgard. 2013. University of Colorado Press, 5589 Arapahoe Avenue, Suite 206C. Boulder, Colorado 80303

USA. 248 pages, 31.29 USD, Paper.

Paul Johnsgard is a lifetime naturalist who has pub-
lished numerous works on taxonomy, natural history,
and ecology in the American West especially the Great
Plains and the state of Nebraska. His long publication
list is filled with natural history monographs of birds
of the Rocky Mountains and Great Plains can be found
on the University of Nebraska, Lincoln website Digital
Commons. His last book on the Wyoming mountain
area was published in 1982 entitled Teton Wildlife:

Observations by a Naturalist, and focussed on different
bird species of the area.

The above work, in collaboration with his former stu-
dent and wildlife photographer Thomas Mangelsen, is
a coffee-table sized, soft-cover book of 224 pages with
many full and double paged photographs spectacular in
their beauty and grand perspective. Landscapes and
natural areas as well as birds and mammals in their nat-
ural landscapes are featured in Mangelsen’s photog-

2014

raphy and many sidebar drawings by Johnsgard himself
are included. A quick count rendered 46 photographs
and 26 drawings which fill up the book and provide
relief in the large pages of text on natural history in
the area.

The area featured is the first confusing aspect of the
book. The Greater Yellowstone Ecosystem is not only
Yellowstone National Park, a name well-known and
recognized by tourists and almost all readers, but in-
cludes much more regional geography. Chapters are
named by their landform areas but only four of the
fourteen chapters are located in or near the park and
the other ten chapters are located far to the south in
regions named Jackson Hole, Grand Teton Park, the
National Elk Refuge, and the Bridger Teton National
Forest. To the north and east there are Montana and
Wyoming cattle ranching areas outside of the parks.
A couple of small maps at the beginning of the book
locates the chapters in their relative areas but there is
no large map to orient the unfamiliar reader with the
whole area. The visitor who has had one or two experi-
ences of visiting Yellowstone will quickly realize that
these are not areas remembered from driving the ring
roads of neither Yellowstone nor day-hiking nearby
trails.

The strength of the book is the extensive research
which Johnsgard has accumulated on population and
conservation data of many of the large mammal species
of elk, bison, deer and wolves. Years of population stud-
ies on timber wolves as they rose and fell coinciding

Book REVIEWS Q7

with the health of the elk population and comments on
current herd and wolf pack management are included
along with comments on population control decisions.
Johnsgard is in favour of annual culls to keep the pop-
ulations healthy and allowing wolf packs to contribute
to the population control. Natural history patterns of
other mammals are also included with life history ac-
counts of beaver, bear, bighorn sheep and various small-
er mammals. His life history stories of the various ani-
mals makes interesting reading for the naturalist who
has encountered these species at Yellowstone or in other
natural habitats.

The birds of course are Johnsgard’s strength and
most of his life’s work. The book includes many ac-
counts of bird-life written in a story-book fashion fol-
lowing an individual or mating pair forming pairs,
nesting-building, and home range defence. Lists of res-
ident birds, mammals, amphibians, reptiles, and fish are
all included to complete the vertebrate counts.

I spent many hours reading the text and leafing back
and forth through the book to view and revisit the pho-
tographs comparing the mammals and birds featured
with the appendices giving more research details. It is
an attractive book to read in short life-story passages
by a masterful story-teller or look at brilliant pictures
by an accomplished photographer. As a good story will
do, I enjoyed my time with it and thought of more tour-
ing in the Yellowstone Plateau region.

JIM O’ NEILL
26095 Taft Road, Novi, Michigan 48374 USA

Yellowstone Wildlife: Ecology and Natural History of the Greater Yellowstone Ecosystem

By Paul A. Johnsgard. 2013. University Press of Colorado, 5589 Arapahoe Avenue, Suite 206C, Boulder, Colorado 80303

USA. 248 pages, 31.29 USD, Paper.

The beautiful portrait of a coyote on the cover of
Yellowstone Wildlife immediately captured my atten-
tion since it directly relates to my two books: Subur-
ban Howls (on eastern coyotes) and My Yellowstone
Experience. With my background as a wildlife biolo-
gist and an avid traveler to Yellowstone, I was a prime
candidate to read this book, which is described in the
inset as “a detailed natural history of the wildlife spe-
cies that call Yellowstone National Park (YNP) and the
Greater Yellowstone Ecosystem their home. Illustrated
with stunning images by renowned wildlife photogra-
pher Thomas Mangelsen, Yellowstone Wildlife describes
the lives of species in the park, exploring their habi-
tats from the Grand Tetons to Jackson Hole.” Herein,
I find one of the major confusions of the book: while it
initially seems like a book about YNP, in actuality only
three of the 14 chapters actually concentrate on Yellow-
stone with most of the remaining focusing on Grand
Teton National Park (GTNP). A slight modification to
the title, Greater Yellowstone Wildlife, would have more
accurately characterized this book which has a more

regional approach, as indicated in the “Preface and
Acknowledgments” section. There, Johnsgard defines
the Greater Yellowstone Ecosystem as 18 million acres
of national parks, national forests, national wildlife re-
fuges, Bureau of land Management lands, plus state
and private landholdings.

The cover inset additionally summarizes the book,
noting “From charismatic megafauna like elk, bison,
wolves, bighorn sheep, and grizzly bears, to smaller
mammals like bats, pikas, beavers, and otters, to some
of the 279 species of birds, Johnsgard describes the
behavior of animals throughout the seasons, with sec-
tions on what summer and autumn mean to the wildlife
of the park. Enhanced by Mangelsen’s wildlife pho-
tography, Yellowstone Wildlife reveals the beauty and
complexity of these species’ intertwined lives and that
of Yellowstone’s greater ecosystem.” I counted 73
images from cover to cover including 27 illustrations
by Johnsgard and 46 photographs by Mangelsen. While
these images clearly add to the text, | would have pre-
ferred the pictures to have been more closely aligned

98 THE CANADIAN FIELD-NATURALIST

with the text along with many additional photos that
directly related to the stories (locations and animals)
that unfolded in each chapter. Further, the existing
images (some being two page spreads) could have been
smaller so more could have been squeezed within the
existing length of the book. For example, in the discus-
sion of the Sagebrush Sea of GTNP, there are two 2-
page spreads of coyotes, both from YNP, yet there is not
one picture of the scenery from that area of the Tetons.

Each of the 14 chapters focused on specific areas of
YNP and GTNP and a natural history story was woven
around the lives of notable animals inhabiting those
regions, such as: coyotes and pronghorn in sagebrush
habitats, gray wolves in Lamar Valley YNP, ravens and
other birds (including golden eagles) by the Grand
Canyon of the Yellowstone, woodpeckers within recent-
ly burned areas, bison around geyser basins, sandhill
cranes near willow flats, trumpeter swans and water-
fowl in and around ponds, bald eagles and ospreys by
oxbows, beavers and elk near aspen stands, grouse and
martens in spruce forests, dippers in mountain streams,
and picas and bighorn sheep in high elevation areas.
The stories in each chapter are very detailed and accu-
rate, for the most part, and Johnsgard has a lengthy sec-
tion of notes and references at the end of the book to
support the behaviours he describes. There were only a
few things I questioned, such as the claims that white-
tailed deer are bigger than mule deer (I always thought
it was the opposite) and male martens help raise the
young (I thought that only female Mustelids raised off-
spring). The notes and references section is impressive
with many sources reviewed; the only odd thing was
the number of very old literature sources, like a 1934
reference being the main one used for bald eagle nat-
ural history, and 1950 and 1959 sources for dippers.
However, this did not affect the content of each chap-
ter, so maybe I am being a tad nitpicky here.

Because each chapter was so in depth, I found my-
self repeatedly trying to find the ultimate meaning of
each section other than gaining a glimpse of the daily
activities of some of the prominent animal residents in
each region. Each chapter usually came to an abrupt
end, with a new and unrelated chapter following it,
making it difficult for the book to maintain a proper
flow. While the descriptions were impressive, there was
so much detail in each chapter I felt bogged down with
information on landmarks (e.g., streams, rivers, moun-
tain ranges) and animals in each region. A small map
of YNP and GTNP at the beginning of the book gave
a general location for each chapter, but I believe that
each individual chapter needed its own map so the
reader could appreciate all of those names and places
mentioned and described in detail throughout the book.
For instance, | have to admit that I have spent much
more time in YNP than GTNP so I found myself get-
ting lost in all of the unfamiliar descriptions of GTNP
because I had no diagram to refer to for each section.

Vol. 128

This was especially evident in Chapter 1, History of the
Greater Yellowstone Ecoregion, which would more
accurately be called Geologic History of the region
since it is a very heavy read about how the area was
created and has changed over millions of years. I found
it very difficult to focus on all of the text in that section
without seeing some image(s) of the landscape features
that “resembles the imprint of a raccoon’s right fore-
foot” (page 2). I was also disappointed that only two
paragraphs at the end of that chapter dealt with human
use of the area over the past 10,000 years, since there
was no other mention of it.

The book has only minor grammatical errors, and
most descriptions seem accurate and realistic for the
location discussed. The appendices provide necessary
information on the wildlife of the region including
checklists of mammals, birds, amphibians, reptiles, and
even invertebrates such as dragonflies and butterflies.
However, and sort of like the majority of the paperback,
I felt that a lot of appropriate information was provided
but the organization could have been better. For exam-
ple, birds are mentioned in Appendices 1-3 including a
checklist, short descriptions for many birds, and longer
descriptions for fewer, more recognizable birds. Thus,
the common birds were mentioned in all three appen-
dices. While each list in and of itself was fine, it seems
that there could have been a more concise summary
of the information presented. Also, while all species
were grouped taxonomically to related species, the
lists were not made in alphabetical order and it seems
that an easier scheme would have aided the reader,
like maybe having a sub-column for different groups
(e.g., waterfowl, raptors, songbirds). Finally, and to my
point above about the need for more coherent struc-
ture, some of the appendices include Red Rock Lakes
and Grays Lake National Wildlife Refuges, yet I fail to
understand the reason for their inclusion since they do
not appear in the text up to that point, even if they may
technically be located within the Greater Yellowstone
Ecoregion (GYE). I had to resort to learning about these
refuges by conducting internet searches. I surmise that
these places were added because the author compiled
all available information on the GYE and included
everything even if it was random or repetitive.

Overall, this is a fine natural history book of the GYE
and is worthy of putting on your bookshelf, especially
for travelers to that region. It provides lots of detailed
information on many different resident wildlife species
ranging from large mammals to inconspicuous birds,
even if a better focus to the book would make it easier
and more enjoyable to read. As it appears now, the
paperback can certainly be read in sections and used
as a reference. I suggest focusing on specific chapters
when visiting the GYE to add to your experience of
those ecosystems.

JONATHAN Way
89 Ebenezer Road, Osterville, Massachusetts 02655 USA

2014

BOTANY

Field Manual of Michigan Flora

Book REVIEWS 99

By E. G. Voss, and A. A. Reznicek. 2013. The University of Michigan Press, c/o Chicago Distribution Center, 11030 South
Langley Avenue, Chicago, Illinois 60628 USA. 990 pages, 25.00 CAD, Cloth.

Many field naturalists in eastern North America will
be familiar with Edward Voss’s classic, three-volume
Michigan Flora, But since its publication beginning in
1972, extensive taxonomic changes led many of us to
supplement this flora with various updated but often
less accessible keys, sometimes photocopied from oth-
ers, and maybe inconveniently stuffed into a filing cab-

inet. With the 2012 publication of the Field Manual of

Michigan Flora, a single pared-down volume offers a
reference that compiles the many changes in nomen-
clature and plant relationships that have occurred in the
last four decades. Over 200 new species for Michigan,
a large majority of them non-native, have also been
described in this new work.

To achieve the synthesis of three volumes into one
while including new species, several important changes
have been made. Perhaps most notably, the botanical
line drawings of selected species have been removed.
This turns out to be less of a limitation than I had an-
ticipated, since I have come to rely upon other sources
for most illustrations anyway. Distribution maps for
Michigan counties are still present but smaller, and
thanks to this more compact form, maps are now phys-
ically closer to their corresponding species description.
Finally, the concise text is now restricted to identifi-
cation and distribution, with taxonomic debate refer-
enced rather than treated directly. Like others, I have
a fondness for Voss’ thorough and often witty intro-
ductory text to many genera in the original flora, and
will continue to refer to it. However, the advantage of
this single volume is undeniable.

As with the preceding Michigan Flora, the stand-out
feature of this work continues to be the clarity and high
quality of the keys, which are more accessible to begin-
ners and amateurs than any other volume I know. Cou-
plets consist of precise but clear language, almost al-
ways with multiple characters. As before, certain keys
even permit identification with only vegetative mate-
rial (e.g. Potamogetonaceae). Practical identification
tips for the field botanist are present in many descrip-
tions. Ontario botanists benefit from native-born co-
author Tony Reznicek’s extensive knowledge of this
area with a number of direct references to Ontario oc-
currences. The glossary is also substantially more com-
prehensive, although it would still benefit from a few
illustrations for more difficult concepts.

Those already familiar with flora of the area will note
many, many nomenclature and taxonomic changes.
Some of these follow Flora of North America (FNA)
treatments, and some do not. The large number of fam-
ilies and species assignments that have changed since
the publication of Michigan Flora are helpfully sum-
marized in a three-page appendix (and take note of

Liliaceae!). The taxonomic concept is relatively con-
servative in some difficult and highly variable groups
(e.g. Rubus, Amelanchier), with quite broadly defined
species or complexes.

Organizational changes are mostly welcome. The
listing of families by alphabetical order (rather than by
taxonomic order) makes the manual more accessible
to the non-taxonomist. Following this pattern, genera
are also given alphabetically within families, and spe-
cies within genera. For the most part, this makes the
book much easier and faster to use.

I have found one significant exception to this, and
that is in the organization of the genus Carex. In gen-
eral, the Cyperaceae keys are a great strength of these
authors, and are among the most-used pages in my
copy. However, the alphabetic listing of all Carex spe-
cies descriptions, rather than by section as in the old
Michigan Flora, does not allow for easy comparison
of the descriptions of closely related species (e.g. all
Ovales). The location of the key for each species is
also not referenced in the index. Since many sections
and their constituent species have changed since the
publication of Michigan Flora, this has left me to flip
through over a dozen pages to locate the appropriate
section key. The overall result, in my experience, is
much less user-friendly than Michigan Flora. Fortu-
nately, this shortcoming appears limited to Carex, since
other large families are sufficiently diverse at the gener-
ic level that their keys are more compact.

A minor disappointment relates to the identifica-
tion of non-native species. Given the large number of
new additions of non-native species and the increasing
attention paid to them in recent years, I would have
liked to see the addition of symbols in each key or
description, so they could be obvious at-a-glance. As
before, this information is available only in the text
of each description.

Still, these issues do not detract greatly from the
overall utility of this book. As I expected, this has be-
come my main reference for flora in southern Ontario,
which has a species assemblage very similar to Michi-
gan. Weighing in at about 3.5 lbs, the durable hardcov-
er could even fit into a backpack — at least on shorter
trips. Thanks to a foundation grant that supported its
publication, it is available at an extremely affordable
price. Whether you are a recent beginner ready for more
information or an experienced field botanist, this prac-
tical work is the new standard identification manual for
vascular plants not only in Michigan, but in adjacent
areas.

HOLty J. BICKERTON

143 Aylmer Avenue, Ottawa, Ontario KIS 2Y1 Canada

100

THE CANADIAN FIELD-NATURALIST

Vol. 128

North Pacific Temperate Rainforests: Ecology and Conservation

Edited by Gordon H. Orians, and John W. Schoen. 2013. The University of Washington Press, P.O. Box 50096, Seattle,
Washington 98145-5096 USA. 383 pages, 60.00 USD, Cloth.

It’s only truly possible to appreciate the geographi-
cal complexity of the region covered in this book by
flying over it, as I had the opportunity to do this past
summer while flying between Seattle, WA, and Fair-
banks, AK. There are more than 6000 islands, ranging
in size from less than | ha to more than 6000 sq km,
between the north end of Vancouver Island and Yaku-
tat, near the Alaska- Yukon border. The adjacent main-
land is dissected by a handful of large river valleys
draining the interior, and numerous small, steep water-
ways cascading down from the glaciers of the coastal
ranges. The biological complexity is correspondingly
enormous and of global importance.

The genesis of this book was a science cruise and
tightly structured workshop to incorporate conserva-
tion biology concepts into management strategies for
conserving the biodiversity and ecological integrity of
the Tongass National Forest in southeast Alaska, which
at 6.8 million ha encompasses 80% of the land area in
that region. During the workshop, it was decided to
produce a book and to expand the scope to include the
broader North Pacific temperate rainforest region.
While much of the text still focuses on the Tongass,
there is considerable input from the British Columbia
part of the rainforest and the concepts are broadly appli-
cable. Each of eight chapters is written by one or more
experts in the topic under discussion, which are then
synthesised in a final chapter which includes conclu-
sions and recommendations.

Much of the biodiversity and endemism of the region
stems from its glacial history when some of the area,
particularly along the outer continental shelf, remained
ice-free during recent glacial advances (chapter 1).
Species colonised the rest of the region from these refu-
gia or along a tidewater route from the north and south,
or later along ice-free corridors from the interior of the
continent. Preservation of these corridors and linkage
sites should be the goal of management plans because
gene flow is a critical process with benefits ranging
from persistence of small populations to introduction
of traits necessary to adapt to fluctuating environments.
And fluctuate it will, as the region is expected to expe-
rience significant climate change in the future, partic-
ularly for the small, steep watersheds where water and
nutrients are transported quickly to estuaries (chapter
2). Recent modelling forecasts a shift from hydrolog-
ical processes dominated by ice and rain-on-snow
events to purely rain-dominated responses; discharge
will become more seasonal, with lower discharge dur-
ing summer (the result of fewer glaciers) and higher
discharge during winter (more rain than snow). These
changes are expected to have regional-scale influences
across much of the North Pacific rainforest, and impact
estuarine and marine production through effects on
amount and timing of carbon and nutrient export. Most

riparian management focuses on fish-bearing streams,
often ignoring headwater streams, but recent research
shows that the structure and function (e.g., inputs of lit-
ter, invertebrates and nutrients) of headwaters is impor-
tant to lower stream reaches. Changes in the life his-
tory of invertebrates due to climate change, as well as
the magnitude, frequency and duration of floods affect-
ing stream morphology, have huge implications for the
adaptation and survival of salmon, a mainstay of wild
food webs, aboriginal culture and economic opportu-
nities in the region.

Natural disturbance patterns (chapter 3) are typified
by mass wasting events on steep, unstable slopes with
thin soils, flooding, avalanches, and many small scale
gap disturbances due to wind throw or stem breakage
from insects/disease, but rarely from fire. Large-scale
clear-cut logging contrasts sharply with this historical
pattern of disturbance because of its location on less-
steep slopes, larger patches of disturbance, and typical
short time to re-harvest. One of the greatest challenges
to developing a scientific approach to conservation in
these forests is to reconcile the temporal scale and pat-
tern of timber harvesting with the long-term patterns
of forest dynamics, complicated by the likely effects of
climate change (e.g., it may be drier, so more fires).

Chapter 5 discusses the Tlingit and Haida tradition-
al use of trees, other plants and animals (salmon was
especially important), and minerals, and the transition
to commercial exploitation. Mining began with the gold
rush in the 1880s, followed by silver, copper, lead, zinc,
barite and marble, but the early operations were mostly
over by the 1940s. High prices for rare earth elements
are driving more recent explorations. Large-scale tim-
ber harvesting of old-growth forests didn’t take off until
the 1950s, and are currently uneconomic and require
subsidies, while second-growth harvests will likely re-
quire even greater subsidies because they are compet-
ing with lower-cost products from many other regions
around the world.

The short-term economic and social benefits of
industrial logging will be paid for by long-term eco-
logical consequences as the result of forest succession
and roads — termed succession debt — even though there
may be short-term (0-19 years after logging) benefits
for some wildlife species, such as deer (chapter 6).
Heavy logging reduces winter range (the old growth
intercepted snowfall), causing a decline in deer num-
bers, which humans blame on predators, and the roads
allow increased legal and illegal harvest of wolves ....
and the cycle continues. It is a perverse irony of the
supply system in the US that timber harvesting, with
attendant new roads, is necessary for the Forest Service
to get funds to decommission, restore and monitor old
ones!

2014

Chapters 7 and 8 address the issues of choosing
areas for protection in reserves. The concepts of con-
servation biology are used to discuss three categories
of reserve design criteria: reserve content (species, eco-
Systems and processes internal to a reserve), reserve
context (interactions between a reserve and its sur-
roundings, i.e., transboundary processes), and emergent
properties (interactions among different components,
and between content and context). The authors also
raise the question — are intact watersheds good re-
serves? — and make the determination that they have
a better chance of maintaining ecological integrity over
the long term without human subsidies, but that small-
er size reserves may be beneficial in some cases. Under
climate change scenarios, reserves that incorporate a
range of climates, site types and elevations will allow
for movement within the reserve, and managing the
matrix habitat around protected areas to facilitate
species migration will be an important component.
Conservation strategies that rely on habitat reserves
and roadless patches often fail because they ignore the
importance and function of the intervening matrix of
unprotected lands (e.g., a wolf pack’s range is larger
than most reserves). Variable retention harvesting
(chapter 9), particularly group or aggregated retention
rather than dispersed retention, is very important in
landscapes that are unlikely to have enough protected
areas and reserves to maintain biodiversity on their
own. It is important, though, that monitoring of trends
and conditions of major indicators (forest structure,
windthrow, growth and regeneration, species and bio-
logical diversity) be done over an entire landscape,
since it is critical to adaptive management practices.
However, the authors caution that most adaptive man-
agement examples have failed because: 1) lack of fund-
ing for monitoring, 2) failure of decision makers to

OTHER

Book REVIEWS

lO]

embrace the concept, and 3) lack of leadership to sus-
tain what is a complex, lengthy process.

The final chapter reviews and synthesises the con-
cepts presented, describes lessons learned from years
of research, highlights uncertainties and gives sugges-
tions how these might be reduced, evaluates potential
new avenues of research, and provides conclusions and
recommendations relating to the challenges and prom-
ises. Forest managers tend to seek solutions that are a
compromise among complementary and competing
objectives, which are generally highly technical, sci-
entific solutions to what are truly political problems,
and they need to start factoring in global implications,
whether or not they are beyond local control. The cor-
nerstone of management or rehabilitation should be to
maintain an ability to adjust in the face of changing
natural conditions; restoring and maintaining processes
that generate habitat, rather than reference conditions,
will enable organisms to adapt to change. The authors
contend that if managers get the scale of their interven-
tions right, as well as ensuring functional and econom-
ic connectivity, they will be well on their way.

Overall, this is a very valuable book which covers
the basics of conservation biology and island biogeog-
raphy theory as applicable to the region, aboriginal and
commercial uses of its natural resources, management
challenges, and opportunities to find a better way for-
ward. Researchers, students and managers will find it
very useful. The centre-piece of 23 black-and-white
photographs has excellent captions, some annotated to
illustrate concepts or issues with forests. There is a good
index and extensive bibliography. Some figures are
reproductions from scientific papers, and are small and
almost illegible, and having a more legible full-page
map of the area would have been helpful.

Cynp!I M. SMITH
P.O. Box 70, Mountain View, Alberta TOK 1NO Canada

A Love Affair With the Birds: The Life of Thomas Sadler Roberts

By Sue Leaf. 2013. University of Minnesota Press, Suite 290, 111 Third Avenue South, Minneapolis, Minnesota 55401

USA. 274 + xviii +16 pages, 29.95 USD, Cloth.

Thomas Sadler Roberts was the right man in the
right place at the right time. His first passion was birds;
in high school in the frontier town of Minneapolis in
1875 he formed, with six other teens, the “Young Nat-
uralists Society.” Roberts was the first president; many
of their meetings were held in his bedroom, which also
served as their library and museum. Only two years
after foundation of the Minnesota Academy of Natu-
ral Sciences at St. Paul, the lads published original arti-
cles on the Black Tern, and compiled a list of 122 spe-
cies of Minnesota birds and a state list of plants. At
their meetings they discussed the works of Elliott Coues
and the theory of evolution proposed by Charles Dar-

win and Alfred Russell Wallace. Roberts was the vale-
dictorian of the class of 11 that graduated from Min-
neapolis High School in June 1877. He then entered
classes at the relatively new University of Minnesota.
In his first summer he studied birds at Herman and
Lake Minnetonka, publishing the results in the Jour-
nal of the Nuttall Ornithological Club, the predeces-
sor of The Auk. The second summer he took part in a
survey of the Lake Superior south shore from Duluth
to Grand Marais and began correspondence with key
ornithologists George Bird Grinnell, J. A. Allen, Wil-
liam Brewster, and Henry Henshaw. In 1880 he worked
as a land surveyor, but watched birds as he surveyed.

102

From 1882 to 1885, Roberts studied medicine at the
University of Philadelphia medical school. His room-
mate was a young lawyer, Carroll Williams, for whom
Roberts’s second child was later named. William Osler
was one of his clinical teachers, so impressed by
Roberts that he visited the Roberts home for break-
fast one morning when he was in Minneapolis years
later. Roberts then did a residency at the Children’s
Hospital, Philadelphia, the first pediatric hospital in
the United States. There was little time for bird study
until he returned to Minneapolis to establish his med-
ical practice in the fall of 1886. He married his friend
Jennie Cleveland in October 1887; by March 1892
they had three children.

When he had a patient warranting special expertise
in diagnosis or treatment, Roberts would sometimes
accompany that patient by train to Rochester, about
two hours distant, to consult with his friends, doctors
Will and Charlie Mayo. The Mayo brothers later rated
Roberts as “one of the five best diagnosticians in the
United States.” Will Mayo, as a regent of the Univer-
sity, later wrote a letter to University of Minnesota
President George Vincent to support the appointment
of Roberts to the Minnesota Natural History Survey.

For 25 years, Roberts was a “hard-driving hard-
working physician,” transported by John Nordquist,
living nearby on 24-hour call to drive Roberts’ horse
and buggy — and later automobile — to home visits.
Roberts’ nurse would often join in his visits to the sum-
mer homes of wealthy industrialists on nearby Lake
Minnetonka, often to deliver their babies there. Roberts
was the chief of staff at the first Minneapolis hospital,
St. Barnabas, from 1893 to 1900. In 1901 he became
the unpaid Professor of Pediatrics at the University of
Minnesota medical school and then, from 1906, the un-
paid Clinical Professor of Children.

Roberts delivered his last baby in 1915, the year the
University elected him as the unpaid (for four years!)
Professor of Ornithology. He solicited funds from his
wealthy friends merely to pay his expenses for speci-
men collecting trips; a few special families kept him on
retainer for medical care. James Ford Bell and John
Crosby (heads of the Washburn-Crosby Company that
later became multi-national General Mills) were two
of his wealthy industrialist friends. Bell himself said:
“Roberts has so many friends who have money.”

Roberts was an innovator. His first bird photos were
exposed onto glass plates at Heron Lake in 1898 with a
heavy mahogany camera powered by a small genera-
tor. These photographs were acclaimed at the Ameri-
can Ornithologists’ Union (AOU) meeting in Washing-
ton that year. In the fall of 1916, his second year at the
fledgling University museum, Roberts showed Sun-
day afternoon movies on a projector donated by Bell.
That year his first annual ornithology class began with
17 lectures and lab demonstrations followed thereafter
by field outings. He helped a bird club organized by the
Woman’s Club evolve into the Minneapolis Audubon

THE CANADIAN FIELD-NATURALIST

Vol. 128

Society. He encouraged Frank and Marie Commons in
their unusually intensive bird banding operations on
their farm at Lake Minnetonka. The main thrust of the
museum in the 1920s was the construction of large
dioramas. In February 1926 Roberts began weekly
radio broadcasts about Minnesota birds on Washburn-
Crosby’s new radio station WCCO.

His major obligation was to write the landmark two-
volume, heavily subsidized Birds of Minnesota, illus-
trated with colour paintings by Allan Brooks, George
M. Sutton, Francis Lee Jacques and Walter A. Weber.
It was finally published to great acclaim in June 1932.
His beloved wife Jennie died that October. Roberts
then resumed making medical house calls to sick pa-
tients to earn some money when income-producing
bonds defaulted during the worldwide financial crisis.

In 1937, over fifty years after his first visit there with
his roommate, Roberts revisited the Williams farm at
New Hope Pennsylvania, where Roberts proposed to
Carroll’s sister Agnes, twice widowed, and she accept-
ed.

In 1938, at his final AOU meeting, held in Washing-
ton, Roberts’ two-volume magnum opus received the
prestigious Brewster Medal from the AOU. That year
work had finally begun on a new museum building de-
ferred in 1931 when Minnesota, hit hard by the depres-
sion, delayed for 9 years its equal match of the large
donation offered by James Ford Bell. The elegant Bell
Museum of Natural History was not opened until
December 1939.

Roberts’ stamina persisted well past eighty. In spite
of a slight stroke in October 1944, he did not miss giv-
ing a single class in ornithology in the spring of 1945.
That summer he had another slight stroke. In October
Agnes had surgery for hemorrhage and thereafter re-
ceived long-term care in Franklin Hospital until her
death on New Year’s Eve. Roberts himself died of a
heart attack 19 April 1946 at age 88.

For centuries a disproportionate number of physi-
cians have shown a serious interest in birds. Thomas
S. Roberts was one of these. My personal interest in
birds was added to my desire to be a physician, when
my two aunts in Manitoba joined together to buy me
Taverner’s Birds of Canada in 1940, an expensive
book, selling for three dollars, which had seemed too
much money for the first aunt to give to a boy only
12 years old. Later my parents bought me the heavily
subsidized two-volume Birds of Minnesota which also
sold for only three dollars per volume; in those two
volumes I read about Roberts himself and his medical-
ly-trained colleagues P. L. Hatch, Elliottt Coues, E. A.
Mearns, and J. C. Hvoslef. These two books and his
bird identification key so valuable to bird banders. have
held prominent places in my library ever since.

Leaf is too polite to mention the Roberts family fi-
nances. Where did Roberts’ father, John, gain the finan-
cial resources that allowed him to move from a farm in
Pennsylvania to reside 25 years in Minneapolis? How

2014

could Thomas and Jennie employ a nurse, a maid, and
a chauffeur throughout thirty years of medical practice,
and then receive no salary whatever for his first four
years at the small initial University Museum?

Sue Leaf has located and reproduced magnificent
early and later photographs, from throughout Roberts’
life, from 1859 at age 18 months through to 1942. Yet,
until he closed his medical office in 1915 after thirty
years of medical practice, she makes it clear that, un-

Bootstrap Geologist: My Life in Science

By Gene Shinn. 2013. University Press of Florida, 15 NW
34.95 USD, Cloth.

Much of Gene Shinn’s professional life has been
spent outdoors in the field. And his life in fieldwork
reads as if it has been both highly enjoyable and deeply
relished. He has spent his time close to, on, in, and
especially under warm sea-water as he investigates car-
bonate reef systems and nearshore environments. In
Bootstrap Geologist, Shinn describes his career, the
projects and questions that have concerned him, the
places he has visited and studied, and the people that he
has met and worked with. Why “bootstrap”? Eighty-
year-old Shinn positions his autobiography as “the life
story of a successful self-taught geological scientist,
initially handicapped by lack of advanced degrees, who
rose to the top of his profession and received its high-
est award”, the Twenhofel Medal of the Society for
Sedimentary Geology in 2009. Throughout his account,
he emphasizes the practical aspects of fieldwork, des-
cribing the many different types of apparatus he has
helped to design and build, from underwater coring
devices to samplers and lo-tech monitoring equipment.
This is the career of a skilled and intuitive “hands-on”
technical specialist, an expert at spotting a problem, and
devising a means and an apparatus to investigate it.

Following college studies in zoology, Shinn’s early
career, between 1958 and 1974, was in the oil industry,
when he worked for Shell, first in Florida and Texas,
then in Qatar and the Persian Gulf, then in Louisiana
and Texas again. In 1974, he moved to work for the
United States Geological Survey (USGS), where he
continued his offshore research, notably as the director
of a specialized marine research work group based in
Florida. Here, his practical skills were put to good use
in keeping the group operational and busy, despite
budgetary and other challenges. Retiring from the USGS
in 2006, Shinn’s career has continued through associ-
ation with the University of South Florida. So he has
seen the research world from three quite different per-
spectives: as an employee of a large multinational com-
pany, as a government servant working for a major
federal agency, and as an affiliate with a mid-sized uni-
versity. His account shows how, despite working under
diverse administrative and funding regimes, he has
always managed to follow and maintain his research

BOOK REVIEWS

103

less he was away from Minneapolis, his patients took
precedence over birds. She deserves warm plaudits for
her careful scrutiny of a mass of data, both ornitho-
logical and medical, that she did not know existed in
adjacent cabinets when she did her zoology graduate
studies in the same room 35 years earlier.

C. STUART HOUSTON

863 University Drive, Saskatoon, Saskatchewan S7N 0J8
Canada

15th Street, Gainesville, Florida 32611 USA. xii + 297 pages,

interests in marine carbonate ecosystems, through tai-
loring his projects to the circumstances of his employ-
ment and taking advantage of opportunities to apply
his skills as they present themselves, including such
odd applications as exploring hydrogen bomb craters
off Pacific atolls.

Taking the mantra of “the present is the key to the
past” seriously, Shinn has focused his research efforts
on investigating modern carbonate systems, especially
subtropical offshore reefs and tidal flats, in the waters
around Florida and the Gulf of Mexico, in the Carib-
bean, off the Philippines, and in the Persian Gulf. Born
as a “‘salt-water conch” on Key West, he spent his late
teenage years as much in sea-water as by it, becom-
ing an accomplished spear-fisher and starting a long
acquaintanceship with coral reefs. A major thread run-
ning through his work has been the way in which
marine organisms build up reefs and other biogenic
structures and the factors that influence the growth and
lithification of reefs. His work is situated at the inter-
face between biological and geological systems, and
is therefore notable for the light that it sheds on both.
The initial motives for undertaking this modern ana-
logue research were pragmatic, springing from the need
to better understand ancient carbonate rocks and espe-
cially reef systems because of their role as host rocks
for hydrocarbon deposits. He has used the insights
gained from work on modern reefs to interpret ancient
reefs and carbonate rocks, for example, by coring algal
reef mounds in New Mexico. Other applications in-
clude the documentation of modern stromatolites off
the Bahamas, the recognition of hurricane and storm-
derived mud layers in tidal channels near the Bahamas,
the elucidation of sabkha stratigraphy in Qatar, and the
identification of algal mud production in Florida Bay.

The results from several of Shinn’s investigations
have challenged conventional views. He describes stud-
ies in which his team concluded that “drill mud had
not affected the growth of corals” near the Philippines,
discovered that “if there was an abundance of junk at
the site, there was more fish” when investigating old
drill sites off the west coast of Florida, and found “lit-
tle lasting harm” from offshore drilling near the Florida

104

Keys. He also studied the movement of groundwater
through the carbonate bedrock underlying southern
Florida, an issue that has significant implications for
assessment of anthropogenic impacts on the environ-
ment and the long-term viability of drinking water sup-
plies. This work started in the late 1980s with an inves-
tigation of shallow groundwater near a garbage dump
in south Florida. He discovered that “pollutants were
restricted to freshwater in a highly porous zone above
a regional impermeable layer just seventeen feet below
the [ground] surface”. In other words, the landfill con-
tainment leaked. His later studies in the 1990s, funded
in part due to concerns over water levels in the Ever-
glades, showed that pollutants and sewage from ex-
panding developments on the Keys were moving
through the highly porous limestone and into sea-waters
around the offshore reefs. His team conducted dye-
injection experiments to show that the flow of the shal-
low groundwater beneath the Keys is towards the
Atlantic, a phenomenon he attributes partly to “tidal
pumping”. However, despite a network of monitoring
wells, Shinn and his team were unable to “prove that
sewage contaminated groundwater was killing the reefs”
due to enhanced algal growth stimulated by nutrient
influxes from sewage.

In recent years, Shinn has considered other reasons
for reef die-offs. Noting a correlation between die-offs
since the mid-1980s and increased input of African
dust, he is convinced that, at least in the Florida and
Caribbean reefs, this is more than coincidence. Shinn’s
research associates have analyzed incoming dust show-
ing that it carries, besides fine-grained clastic material,
various biotic materials including bacteria and spores,
some of them potential pathogens. Yet dust has been
a factor in these ecosystems for millennia; Charles
Darwin noted dust fall in mid-Atlantic in 1832 during
the second Beagle voyage and reported on its organic
content (Darwin 1846, Geological Society Journal 2:
267-274). What is not clear to me from Shinn’s discus-
sion here is why the dust should have become more
deadly for reefs in recent decades.

The results of Shinn’s studies have not always been
well received. Conclusions from his dust research are
controversial. As he remarks for the Florida Keys drill
sites study, his team “didn’t find and document the
widespread death and destruction that was expected”

Tracks and Shadows: Field Biology as Art

By Harry W. Greene. 2013. University of California Press,
+ 280 pages, 29.95 USD, Cloth.

Harry Greene, a professor at Cornell University, is
the author of the classic herpetology book Snakes:
The Evolution of My. stery in Nature. His new book is
part memoir, part profile of the great herpetologist
Henry Fitch, and part meditation on snakes and the
quest to understand them.

THE CANADIAN FIELD-NATURALIST

Vol. 128

nor did they “find what the public, the media, and the
academic community wanted”. Though he has followed
the evidence where it led, he does paint himself as a bit
of a contrarian. Shinn expresses little interest or con-
fidence in modelling or theory, highlighting that his
work has been “very field oriented”. He also has a jaun-
diced view of permits and regulations that he sees as
strangling the capacity to do research, especially in na-
tional parks or protected areas. His memoir is fasci-
nating for highlighting changes in research methods
between the late 1950s and now. Some of the method-
ological quandaries we grapple with today were sim-
ply not considered decades ago. For example, Shinn
writes about using dynamite to create “explosures”,
holes in coral reefs, in the early 1960s when “few cared
about the environmental effects of dynamite”. It is dif-
ficult to imagine explosives as an approved research
tool for reefs now! Perhaps not all changes in the re-
search milieu are bad.

Shinn’s writing style is laboured in places and
sprinkled with superfluous exclamation marks, but it
becomes more readable as the book goes on. There are
early digressions into tangential matters that would
have been better omitted; the distasteful story of a
deeply disturbed physician being one. Shinn’s narrative
is more assured and interesting when he is writing
about his professional career. I was intrigued by his
descriptions of ex-pat life in Doha, Qatar, for example.
I enjoyed his stories about various field exploits, in-
cluding a recurrent fascination with “whitings”, lime
mud precipitated from sea-water that forms white
cloudy patches in the ocean. Only in recent years has
the role of planktic cyanobacteria in these precipitates
been recognized. Exciting new research suggests that
such ancient mud may be implicated in the origin of
some Middle Eastern oil deposits. Shinn hopes this
“hypothesis survives the test of time and future inves-
tigation”. Certainly, his field observations and sub-
stantial research contributions will remain valuable.
And anyone interested in modern and ancient reef
systems will find much to captivate them in Shinn’s
entertaining memoir of his industrious and energetic
life in sunshine and sea-water.

ALWYNNE B. BEAUDOIN

Royal Alberta Museum, 12845-102 Avenue NW, Edmon-
ton, Alberta TSN 0M6 Canada

2120 Berkeley Way, Berkeley, California 94704-1012 USA. xiii

In many ways this is a tale of two herpetologists:
Fitch and Greene. Henry Fitch was born in 1909 and
although educated in a one-room school-house he went
on to become a professor at the University of Kansas.
Before arriving in Kansas, he was drafted into the army
during World War II, serving as a medic in Europe.

2014

After the war, Fitch eventually landed a job with the
University of Kansas in 1948. A dedicated field biol-
ogist, Fitch published nearly 200 scientific papers and
although he is mainly known as a herpetologist he also
published papers on plants, snails, spiders, mammals
and birds. His most amazing contribution is a 50-year
study of the snakes of the Natural History Reservation
owned by the University of Kansas (and now re-named
the Fitch Natural History Reservation in his honour),
He continued conducting field work into his 90s. When
queried by Greene about his favourite animals, Fitch
replied “Alligator lizards, copperheads, and garter
snakes, because of their interesting natural histories”
(page 228).

Harry Greene’s father was in the US air force and
the family moved repeatedly. Before Greene finished
high school he had lived in five different states. He had
an early and profound interest in nature, particularly
snakes. At age 12, having met herpetologist Charles
Carpenter, Greene announced to his parents that some-
day he too would be a professor of herpetology. His
first publication, a brief note on the vestigial defensive
display of a species of blind cave salamander, was pub-
lished when he was just 16 years old. After graduation
from high school he got to spend the summer work-
ing with Henry Fitch. To help pay his way through uni-
versity, Greene got a job as an ambulance worker, deal-
ing with car crashes, heart attacks and more than his
share of death. Greene was drafted into the army during
the Vietnam War and, like Fitch, served as a medic.
Greene fully expected to be sent to Vietnam, but ended
up being stationed in Germany, where he spent nearly
two years before returning to civilian life and graduate

BooK REVIEWS

LOS

school back in the USA. Fascinated by snake behav-
iour, Greene examined behavioural ecology from an
evolutionary point of view, first in coral snakes for his
master’s thesis and then in constrictors for his PhD.

While part one of the book explores the background
of Fitch and Greene, part two, focuses more on snake
biology, ecology and evolutionary history. Greene has
managed to turn his research on snakes into a global
pursuit resulting in stories about Terciopelos in the rain-
forests of Costa Rica, Green Anacondas in Brazil, Green
Bush Vipers in Uganda, and a 15 year study of Black-
tailed Rattlesnakes in Arizona. Along the way Greene
muses about the co-evolution of primates and snakes.
The first snakes evolved over one hundred million years
ago. It doesn’t seem unreasonable that the early con-
strictors preyed upon early primates, possibly shaping
their evolution and contributing to people’s fear and
fascination with snakes.

There are many books filled with empty facts about
animals and there are also many about people’s person-
al experience with nature, but lacking intellectual depth.
Greene’s book offers fascinating insights into snake
biology and evolutionary history from a true expert, but
also portrays the human side of exploring nature, con-
ducting science and struggling through life. This fusion
would not work without Greene’s evocative writing
that captures both the richness of biological life and the
fragility of our own lives. This is not just a book for
snake lovers, but for anyone enthralled by nature and
life itself.

DAVID SEBURN

Seburn Ecological Services, 2710 Clarenda Street, Ottawa,
Ontario K2B 7S5 Canada

Alfred Russel Wallace (2013) On the Organic Law of Change: A Facsimile Edition and
Annotated Transcription of Alfred Russel Wallace’s Species Notebook of 1855-1859

Annotated by James T. Costa. 2013. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA.

xii + 573 pages, 49.95 USD, Cloth.

Let me say it right up front: I love this book! Read-
ing it is a bit like listening in on the musings of an emi-
nent colleague, trying to follow their train of thought,
catching bits of their reasoning and ideas, and being
impressed by their knowledge and insights. Tracing the
development of Wallace’s thinking on biogeography
and evolution as shown through this notebook is both
a challenge and a pleasure. Watching him tussle with
concepts, such as the definition of a species or the dis-
tinction between variety and species, is fascinating.
These are concepts that biologists and palacontologists
still struggle with and discuss. Reading about his ener-
getic collecting activities is also absorbing, even though
his accounts of orangutan hunts are harrowing. As with
any colleague, you don’t always agree with everything
they do. On the other hand, his strenuous and uncom-
fortable travels and efforts to conserve and ship his col-
lections elicit wry smiles of empathy.

Wallace is an endearing character. He started life
disadvantaged by social circumstances and poverty but
by dint of intelligence and incredible hard work turned
himself into an outstanding naturalist and scientific
thinker. Costa describes him “as the greatest biogeog-
rapher of his century.” The story of how he came up
with the idea of natural selection while recovering from
fever in a hut on a remote island is well-known. In a
rather touching display of trust, he sent his resulting
essay “On The Tendency of Varieties to Depart Indef-
initely from the Original Type” to Charles Darwin, who
received it in June 1858 and was devastated to realize
that someone else had had the same idea as him. Ina
hastily-organized presentation facilitated by Charles
Lyell and Joseph Hooker, Wallace’s essay and a sum-
mary of Darwin’s thoughts were read together at a
meeting of the Linnaean Society in London on July 1,
1858. Darwin, who was on the spot and had the better

106

PR machine, has always had the lion’s share of the
glory for coming up with the idea of natural selection.
Yet, to his enormous credit, Wallace seems never to
have harboured a grudge or expressed any resentment
toward Darwin. He seems to have had a rare modesty
and generosity of spirit.

Wallace lived for 31 years longer than Darwin, dying
in 1913 at the age of 90. In later life, his reputation
became somewhat diminished, in part because of his
interest in topics such as spiritualism. His scientific
stature was not widely recognized during the 20"
century, nor was the range of his achievements well
known. He was mainly remembered for defining Wal-
lace’s Line. This important biogeographic division be-
tween the fauna and flora of the Asian and Australian
regions was another discovery that arose from his
work in the Malay Archipelago recorded in this note-
book. However, in recent years, Wallace has been re-
ceiving more recognition for his independent discov-
ery of natural selection and evolution. Perhaps this is
because the Darwin bicentennial in 2009 stimulated
much in-depth re-examination and re-evaluation of the
history of thought concerning evolution in the 19" cen-
tury. Perhaps it is because with the clarity of hindsight
and temporal distance we can now evaluate the con-
tributions of both men more dispassionately. As Costa
comments in his introduction, today both “are equally
honoured as pioneers and discoverers of one of the most
profound insights into nature — and ourselves — yet
grasped by humanity.”

This facsimile edition Wallace’s important notebook
has been released in time to commemorate the cente-
nary of his death and as a celebration of his life. It
showcases one of ten surviving notebooks generated
during his travels in the Malay Archipelago between
1854 and 1862, one of the most formative and influ-
ential intervals in his life. The other notebooks com-
prise field notes, specimen records, and journals. This
Species Notebook, started some months after his arrival
in Singapore in April 1854, includes accounts of his
fieldwork and collecting. But it is more than that. In
the interstices between his field activities, Wallace read
deeply, especially Lyell’s Principles of Geology, and
recorded his thoughts about what he was reading in this
notebook. For Waltace as for Darwin, Lyell seems to
have acted as a catalyst for thought; indeed, this incite-
ment to thought was arguably Lyell’s most important
contribution to science. Wallace was especially con-
cerned to document the points where he disagreed with
Lyell — and there were a lot of them — and work out his
Own counter-arguments. These arguments could have
formed the basis for his own book on evolution, a book
he never wrote, although, as Costa sadly remarks, it is
“the book that should have been.” From the perspective
of his thinking on natural selection, Wallace’s lengthy
musings on the fossil and rock records and their impli-

cations and the nature and relationship of plant parts
are particularly interesting.

THE CANADIAN FIELD-NATURALIST

Vol. 128

The breadth of Wallace’s interests and observations
is impressive. He described his search for birds of par-
adise and comments on their varying morphologies and
behaviours. He considered and developed a “Plan to
stop the further increase of Synonyms,” the underlin-
ing perhaps emphasizing his frustration with the state
of taxonomy, by proposing that a central authority
should decide on precedence of names and adjudi-
cate disputes over synonyms. As Costa points out,
this anticipated the establishment of the International
Commission of Zoological Nomenclature by several
decades. Wallace identified and collected many species
of insects, primarily beetles and butterflies, especially
large and showy taxa, and this notebook contains many
observations of their behaviour and habitat preferences,
as well as some delightful and detailed drawings.

Wallace also expended much thought on the distinc-
tion between instinct and learned behaviour, especial-
ly as regards the construction of birds’ nests. There are
many pages of notes and commentary on this topic in
this notebook. It was this train of thought, moreover,
that eventually led him to consider the problem of
human cognition. In 1869, after returning from his
travels in the Malay islands, Costa recounts that Wal-
lace “publicly declared that the human brain could not
be explained by natural selection, and that therefore
human consciousness and cognitive abilities must be
the product of a divine plan of some kind.” Unsurpris-
ingly, this viewpoint led to a split with Darwin, and
probably in part accounts for his diminished reputa-
tion in later years.

Although it spans the interval when he wrote his fa-
mous essay, the Species Notebook contains no com-
ments, drafts, or obvious notes related to that work.
Readers will share Costa’s frustration at the “madden-
ingly little information” from Wallace during “the very
period in which he had his insight into natural selec-
tion and penned the landmark paper.” His notes do
record his bouts of illness and the many “hot rain
night[s]” he endured. Perhaps this is explanation enough
for his rather perfunctory record of his stay on Ternate
and the neighbouring island of Gilolo, now called
Halmahera, the place where he actually wrote his essay.
“If only,” laments Costa, “Wallace had jotted some-
thing about his momentous breakthrough.” Well, he
didn’t, but the notes he did leave us make for fasci-
nating if demanding reading nonetheless.

This book is beautifully designed and laid out. The
left-hand page has a facsimile of a page from Wallace’s
notebook, with the handwritten text transcribed next
to it. Some facsimile pages include Wallace’s drawings
or even, in a few cases, pasted in specimens. It is re-
markable how clear Wallace’s handwriting is, whether
he is working out the proper dimensions for specimen
labels, writing an account of a day in the field, or tal-
lying his collections. There are few scribbled, illegible,
perfunctory or blotched notes here. The facing right-
hand page has annotations compiled by Costa, includ-

2014

ing additional information about the life-forms, people,
places, and incidents mentioned in Wallace’s notes.
These are sometimes accompanied by illustrations from
Wallace’s later publications or other near contemporary
sources. Costa also provides background contextual
information about Wallace’s ruminations on species
questions and what he has been reading. These anno-
tations are insightful, informative, and often lengthy.
They add greatly to the value of this book and, indeed,
many of Wallace’s notes would not be comprehensible
without these extensive and knowledgeable comments.
Costa has done a tremendous job of ferreting out and

BooK REVIEWS 107

elaborating the sometimes opaque references in the
notes, especially the mention of people who were prob-
ably well known at the time but are now obscured by
history. In presenting the Species Notebook to us, Costa
has produced a work of admirable scholarship. This
book will certainly help to elevate Wallace to his right-
ful place in the pantheon of 19" century natural scien-
tists and garner him additional respect as an original
and perceptive thinker.

ALWYNNE B. BEAUDOIN
Royal Alberta Museum, Edmonton, Alberta, Canada

108

NEw TITLES

Prepared by Roy John

+ Available for review * Assigned

THE CANADIAN FIELD-NATURALIST

Vol. 128

Currency Codes — CAD Canadian Dollars, USD U.S. Dollars, EUR Euros, AUD Australian Dollars.

ZOOLOGY

* Amphibians of Ohio. Edited by Ralph A. Pfingsten,
Jeffrey G. Davis, Timothy O. Matson, Greg Lipps, Jr.,
Doug Wynn, and Brian J. Armitage. 2013. Ohio Bio-
logical Survey, P.O. Box 21370, Columbus, OH, USA,
43221-0370. 916 pages, 90.00 USD, Cloth.

Animal Earth. By Ross Piper. 2013. Thames & Hud-
son, 181A High Holborn, London, UK, WC1V 7QX.
320 pages, 45.00 USD, Cloth.

Animals of the Serengeti and Ngorongoro Conser-
vation Area. By Adam Scott Kennedy, and Vicki Ken-
nedy. 2014. Princeton University Press, 41 William
Street, Princeton, NJ, USA, 08540-5237. 152 pages,
27.95 USD, Paper.

A Field Guide to the Ants of New England. By
Aaron M. Ellison, Elizabeth J. Farnsworth, and Nicho-
las J. Gotelli. 2012. Yale University Press, P.O. Box
209040, New Haven, CT, USA, 06520-9040. 416
pages, 33.46 CAD, Paper.

Phillipps’ Field Guide to the Birds of Borneo, Sabah,
Sarawak, Brunei, and Kalimantan (Third Edition,
Fully Revised). By Quentin Phillipps, and Karen Phil-
lipps. 2014. Princeton University Press, 41 William
Street, Princeton, NJ, USA, 08540-5237. 384 pages,
35.00 USD, Paper.

Birds of Kenya’s Rift Valley. By Adam Scott Kennedy.
2014. Princeton University Press, 41 William Street,
Princeton, NJ, USA, 08540-5237. 264 pages, 29.95
USD, Paper.

Birds of the Serengeti and Ngorongoro Conserva-
tion Area. By Adam Scott Kennedy. 2014. Princeton
University Press, 41 William Street, Princeton, NJ,
USA, 08540-5237. 224 pages, 27.95 USD, Paper.

Ten Thousand Birds. By Tim Birkhead, Jo Wimpen-
ny, and Bob Montgomerie, 2014. Princeton University
Press, 41 William Street, Princeton, NJ, USA, 08540-
5237. 544 pages, 45.00 USD, Cloth.

Beetles of Eastern North America. By Arthur V.
Evans. 2014. Princeton University Press, 41 William
Street, Princeton, NJ, USA, 08540-5237. 544 pages,
35.00 USD, Paper.

Bumble Bees of North America — An Identification
Guide. By Paul H. Williams, Robbin W. Thorp, Leif

L. Richardson, and Sheila R. Colla. 2014. Princeton
University Press, 41 William Street, Princeton, NJ,
USA, 08540-5237. 208 pages, 24.95 USD, Paper.

The Fauna of British India Including Ceylon and
Burma — Butterflies Vol. 1. By G. Talbot. 2013. Today
& Tomorrows Printers and Publishers Karol Bagh,
Delhi, India, 110005. xxx + 600 pages, 1000.00 INR
(about 18.00 CAD).

Wildlife of the Caribbean. By Herbert A. Raffaele,
and James W. Wiley. 2014. Princeton University Press,
41 William Street, Princeton, NJ, USA, 08540-5237.
304 pages, 19.95 USD, Paper.

Conus of the Southeastern United States and Carib-
bean. By Alan J. Kohn. 2014. Princeton University
Press, 41 William Street, Princeton, NJ, USA, 08540-
5237. 528 pages, 99.50 USD, Cloth.

* Deer. By John Fletcher. 2014. The University of
Chicago Press, 1427 East 60th Street, Chicago, IL,
USA, 60637. 207 pages, 16.02 USD, Paper.

+ Dolphin. By Alan Rauch. 2014. The University of
Chicago Press, 1427 East 60th Street, Chicago, IL,
USA, 60637. 207 pages, 16.02 USD, Paper.

Britain’s Dragonflies — A Field Guide to the Dam-
selflies and Dragonflies of Britain and Ireland —
Third Edition. By Dave Smallshire, and Andy Swash.
2014. Princeton University Press, 41 William Street,
Princeton, NJ, USA, 08540-5237. 208 pages, 25.95
USD, Paper.

Lost Animals — Extinction and the Photographic
Record. By Errol Fuller. 2013. Princeton University
Press, 41 William Street, Princeton, NJ, USA, 08540-
5237. 240 pages, 29.95 USD, Cloth.

The Amazing World of Flyingfish. By Steve N. G.
Howell. 2014. Princeton University Press, 41 William
Street, Princeton, NJ, USA, 08540-5237. 48 pages,
12.95 USD, Cloth. .

Life on the Rocks: A Portrait of the American Moun-
tain Goat. By Bruce L. Smith. 2012. University of
Colorado Press, University Press of Colorado. 5589
Arapahoe Ave., Suite 206C, Boulder, CO, USA. 80303.
192 pages, 34.95 USD, Cloth.

2014

A Field Guide to the Larger Mammals of Tanzania.
By Charles Foley, Lara Foley, Alex Lobora, Daniela
De Luca, Maurus Msuha, Tim R. B. Davenport, and
Sarah Durant. 2014. Princeton University Press, 41
William Street, Princeton, NJ, USA, 08540-5237. 320
pages, 29.95 USD, Paper.

* Octopus. By R. Schweid. 2014. The University of
Chicago Press, 1427 East 60th Street, Chicago, IL,
USA, 60637. 207 pages, 16.02 USD, Paper.

Introduction to Marine Plankton. By Abhijit Mitra,
Kakoli Banerjee, and Avijit Gangopadhyay. Daya Pub-
lishing House, A Unit of Astral International Pvt Ltd.,
81, Darya Ganj, Near Hindi Park, Delhi Medical Asso-
ciation Road, New Delhi, India, 110002. 102 pages,
12.00 USD.

* Rabbit. By V. Dickenson. 2014. The University of
Chicago Press, 1427 East 60th Street, Chicago, IL,
USA, 60637. 207 pages, 16.02 USD, Paper.

The Extreme Life of the Sea. By Stephen R. Palumbi,
and Anthony R. Palumbi. 2014. Princeton University
Press, 41 William Street, Princeton, NJ, USA, 08540-
5237. 256 pages, 27.95 USD, Cloth.

* How Snakes Work: Structure, Function and
Behaviour of the World’s Snakes. By Harvey Lilly-
white. 2014. Oxford University Press, 198 Madison
Avenue, New York, NY, USA, 10016. 24 pages, 54.05
CAD, Cloth.

A Sparrowhawk’s Lament — How British Breeding
Birds of Prey Are Faring. By David Cobham, and
Bruce Pearson. 2014. Princeton University Press, 41
William Street, Princeton, NJ, USA, 08540-5237. 256
pages, 35.00 USD, Cloth.

BOTANY

* Braiding Sweetgrass: Indigenous Wisdom, Scien-
tific Knowledge, and the Teachings of Plants. By
Robin Wall Kimmerer. 2013. Milkweed Editions, 1635
West 12th Avenue, Vancouver, Canada, V6J 2E3. 320
pages, 24.00 CAD.

BooK REVIEWS

109

Trees of Western North America. By Richard Spel-
lenberg, Christopher J. Earle, and Gil Nelson 2014.
Princeton University Press, 41 William Street, Prince-
ton, NJ, USA, 08540-5237. 448 pages, 29.95 USD,
Paper, 60.00 USD, Cloth.

Trees of Eastern North America. By Gil Nelson,
Christopher J. Earle, and Richard Spellenberg. 2014,
Princeton University Press, 41 William Street, Prince-
ton, NJ, USA, 08540-5237. 656 pages, 29.95 USD,
Paper, 65.00 USD, Cloth.

OTHER
* Tracks and Shadows: Field Biology as Art. By
W. Harry. 2013. University of California Press, 1445

Lower Ferry Road, Ewing, NJ, USA, 08618. 296 pages,
29.95 USD, Cloth.

* A Love Affair with Birds: The Life of Thomas
Sadler Roberts. By Sue Leaf. 2013. University of
Minnesota Press, Suite 290 111 Third Avenue South,
Minneapolis, MN, USA, 55401. 29.95 USD.

* The dismal state of the Great Lakes. By James
Pinson Ludwig, 2013. Xlibris LLC, 1663 Liberty Drive,
Suite 200, Bloomington, IN, USA, 47403. 273 pages,
20,95 USD.

Marine Community Ecology and Conservation. By
Mark D. Bertness, John F. Bruno, Brian R. Silliman,
and John J. Stachowiczi. 2013. Sinauer Associates, Inc.,
23 Plumtree Road, P.O. Box 407, Sunderland, MA,
USA, 01375-0407. 675 pages, 109.95 USD, Cloth.

* On the Organic Law of Change — A Facsimile
Edition and Annotated Transcription of Alfred
Russel Wallace’s Species Notebook of 1855-1859.
By Alfred Russel Wallace. (Introduction and notes by
James T. Costa). 2013. Harvard University Press, 79
Garden Street, Cambridge, MA, USA, 02138. 592
pages, 49.95 USD, Cloth.

News and Comment

Upcoming Meetings

Meeting of the Canadian Botanical Association 2014

The 50" annual meeting of the Canadian Botanical
Association hosted by the IRBV, an institute for re-
search in plant biology, the Montreal Botanical Garden,
Space for life, and the Université de Montréal to be held
15-18 June 2014 at the Montreal Botanical Garden,

Montreal, Quebec. The theme of the meeting, “50 years
of Botany in Canada”, will focus on the important
contribution of Canadian researchers to the develop-
ment of plant biology. More information is available
at http://www. irbv.umontreal.ca/cba-meeting?lang=en.

International Conference on Biodiversity & Sustainable Energy Development 2014

The 3™ International Conference on Biodiversity
& Sustainable Energy Development to be held 24—26
June 2014 at the Valencia Conference Centre, Valencia,
Spain. The theme of the meeting is: Milestones of inno-
vative scientific research in biodiversity and its allied

Botany 2014 Meeting

The annual Botany conference to be held 26-30 July
2014 at the Boise Center in Boise, ID. This annual
meeting is a multiple scientific society conference
(American Bryological and Lichenological Society;
American Society of Plant Taxonomists; Botanical

areas. Registration is currently open. More informa-
tion is available at http://www.omicsgroup.com/bio
diversity-sustainable-energy-conference-20 | 4/index

.php.

Society of America; American Fern Society; Interna-
tional Association of Plant Taxonomists) serving over
6000 plant scientists whose research and practice span
the globe. Registration is currently open. More informa-
tion is available at http://www.botanyconference.org/.

110

TABLE OF CONTENTS (concluded) Volume 128, Number | 2014

Book Reviews

ZOOLOGY: Into the Night: Tales of Nocturnal Wildlife Expeditions — Owls — A Pocket Guide to Salamanders of

Pennsylvania — Snapper — The Crossley ID Guide to Britain and Ireland — The Warbler Guide — Yellowstone

Wildlife: Ecology and Natural History of the Greater Yellowstone Ecosystem — Yellowstone Wildlife: Ecology and

Natural History of the Greater Yellowstone Ecosystem 91
Botany: Field Manual of Michigan Flora — North Pacific Temperate Rainforests: Ecology and Conservation 99

OTHER: A Love Affair With the Birds: The Life of Thomas Sadler Roberts — Bootstrap Geologist: My Life in Science
— Tracks and Shadows: Field Biology as Art — Alfred Russel Wallace (2013) On the Organic Law of Change: A
Facsimile Edition and Annotated Transcription of Alfred Russel Wallace's Species Notebook of 1855-1859 10]

New TITLES 108

News and Comment

Meeting of the Canadian Botanical Association 2014 110
International Conference on Biodiversity & Sustainable Energy Development 2014 110
Botany 2014 Meeting 110

Mailing date of the previous issue 127(4): 14 January 2014

THE CANADIAN FIELD-NATURALIST Volume 128, Number |

Articles

Distribution and abundance of benthic macroinvertebrates and zooplankton in lakes in Kejimkujik
National Park and National Historic Site of Canada, Nova Scotia
CHRISTINA NUSSBAUMER, NEIL M. BurGESS, and Russ C. WEEBER

An illustrated key to the mandibles of small mammals of eastern Canada
DOMINIQUE FAUTEUX, GILLES LUPIEN, FRANCOIS FABIANEK,
JONATHAN GAGNON, MARION SEGUY, LOUIS IMBEAU

Characteristics of Barred Owl (Strix varia) nest sites in Manitoba, Canada
Topp M. WHIKLO and JAMES R. DUNCAN

Yellow Warblers (Setophaga petechia) rear second broods in some years at Delta Marsh, Manitoba
SPENCER G. SEALY

Asynchronous breeding and variable embryonic development period in the threatened Northern
Leopard Frog (Lithobates pipiens) in the Cypress Hills, Alberta, Canada: conservation and man-
agement implications

LEA A. RANDALL, LYNNE D. CHALMERS, AXEL MOEHRENSCHLAGER, and ANTHONY P. RUSSELL

Diet of the Pacific Sand Lance (Ammodytes hexapterus) in the Salish Sea, British Columbia, in the
1960s J. MARK HIPFNER and MorrRA GALBRAITH

Melanistic diversity in the Maritime Gartersnake, Thamnophis sirtalis pallidulus, in Nova Scotia,
Canada JOHN GILHEN and FRED W. SCOTT

Notes

Thickness of Common Murre (Uria aalge) eggshells in Atlantic Canada
DONALD W. PirtE-HAy and ALEXANDER L. BOND

The most northerly Black Witch (Ascalapha odorata): a tropical moth in the Canadian Arctic
TORBJORN EKREM, PETER G. KEVAN, THOMAS S. WoopcOCckK, and PAUL D. N. HEBERT

Multiple crossings, of a large glacial river by Canada Lynx (Lynx canadensis)
DASHIELL FEIERABEND and KNUT KIELLAND

Tributes
A tribute to Kenneth William Stewart, 1936-2011 FRANCIS R. COOK

2014

25

38

44

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63

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80

84

(continued on inside back cover)

ISSN 0008-3550

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The CANADIAN
FIELD-NATURALIST

Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

April-June 2014

Volume 128, Number 2

The Ottawa Field-Naturalists’ Club

FOUNDED IN 1879

Patron
His Excellency the Right Honourable David Johnston, C.C.,C.M.M., C.O.M.,C.M.
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these
fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-
ments of high quality for living things.
Honorary Members

Ronald E. Bedford Paul M. Catling John M. Gillett E. Franklin Pope
Edward L. Bousfield Francis R. Cook Peter W. Hall Joyce M. Reddoch
Charles D. Bird Bruce Di Labio Christine Hanrahan Allan H. Reddoch
Fenja Brodo Anthony J. Erskine C. Stuart Houston Dan Strickland
Irwin M. Brodo J. Bruce Falls Theodore Mosquin John B. Theberge
Daniel F. Brunton Jay Fitzsimmons Robert W. Nero Sheila Thomson

Michael D. Cadman

2014 Council
President: Fenja Brodo Daniel F. Brunton Don Hackett Karen McLachlan-Hamilton
1 Vice-President: vacant Carolyn Callaghan David Hobden Lynn Ovenden
24 Vice-President: Jeffrey H. Skevington Barbara Chouinard Diane Kitching Rémy Poulin
Recording Secretary: Annie Bélair Owen Clarkin Diane Lepage Henry Steger
Treasurer: Ken Young Barry Cottam Ann MacKenzie Eleanor Zurbrigg

Ian Davidson

To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate
P.O., Ottawa, ON, K1Z 1A2, or e-mail: ofnc@ofnce.ca. For information on Club activities, go to www.ofnc.ca

The Canadian Field-Naturalist

The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this
journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.

Website: www.canadianfieldnaturalist.ca/index .php/cfn

Editor-in-Chief: Dr. Carolyn Callaghan Assistant Editor: Dr. Trina Rytwinski
Copy Editor: Sandra Garland Typographer: Wendy Cotie
Journal Manager: Dr. Jay Fitzsimmons: subscriptions @canadianfieldnaturalist.ca or via postal address above.

Book Review Editor: Roy John

Associate Editors: Charles D. Bird Anthony J. Gaston David Nagorsen Jeffery M. Saarela
Paul M. Catling Thomas S. Jung Claude B. Renaud David C. Seburn
Francis R. Cook Donald F. McAlpine Marty Obbard Jeffrey H. Skevington
Jennifer R. Foote Garth Mowat

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The Canadian Field-Naturalist J) °

Volume 128, Number 2

i April-June 2014 Y

Effectiveness of Stream Sampling Methods in Capturing Non-native
Rusty Crayfish (Orconectes rusticus) in Ontario

Scott M. Retp!:? and JANE DEVLIN!

Aquatic Research and Monitoring Section, Ontario Ministry of Natural Resources, Trent University, 2140 East Bank Drive,
Peterborough, Ontario K9J 7B8 Canada
“Corresponding author: reid.scott(@ontario.ca

Reid, Scott M., and Jane Devlin. 2014. Effectiveness of stream sampling methods in capturing non-native Rusty Crayfish
(Orconectes rusticus) in Ontario. Canadian Field-Naturalist 128(2): 111-118.

Habitat alteration and species introductions have contributed to the decline of native crayfish in Ontario. Although lake populations
of crayfish in Ontario are monitored, there is no corresponding program for streams. We used removal-based sampling to
evaluate the efficacy of three sampling techniques (backpack electrofishing, hand capture, and seining) to characterize native
and non-native crayfish populations in six streams in the Kawartha Lakes region and five streams in the Muskoka/Haliburton
Lakes region. All types were effective at detecting non-native Rusty Crayfish (Orconectes rusticus). Rusty Crayfish were
collected from 65% of samples, constituted 90% of the total catch, and were the only species present in 30% of streams.
Compared with other methods, electrofishing was more likely to capture additional crayfish species. Removal-based sampling
was not a reliable approach for estimating capture probability and population size. Failure of the removal model was due to
increases in the number of crayfish captured after the first pass or too few individuals collected. Mean capture probabilities
for electrofishing (0.30) and hand capture (0.31) did not result in reliable population estimates. Compared with seining, electrofish-
ing and hand capture resulted in more sexually active males (fewer sexually inactive males) and more small (< 25 mm carapace
length) individuals. For each method, there were differences in capture probability among length classes. A combination of
electrofishing and seining (with multiple passes) would maximize species detection, permit sampling of a range of habitat types,
and be easily integrated into existing stream fish surveys.

Key Words: invasive species; monitoring; Rusty Crayfish; capture success; streams; Orconectes rusticus; Ontario; electrofishing:
hand capture; seining; Appalachian Brook Crayfish; Cambarus bartonit, C alico Crayfish; Orconectes immunis; Allegheny
Crayfish; Orconectes obscurus; Northern Clearwater Crayfish, Orconectes propinquus; Virile Crayfish; Orconectes virilis

Introduction

Almost half of the freshwater crayfish species in
North America are considered imperilled, making cray-
fish second only to freshwater mussels in terms of con-
servation concern (Butler et a/. 2003; Taylor ef al.
2007). Worldwide, habitat alteration and non-native
crayfish are considered to be the key threats to native
crayfish species (Taylor et al. 2007). In Ontario, urban-
ization, draining of wetlands, and acid rain have hada
negative effect on crayfish habitat (Guiasu 2007, 2009;
Edwards et al. 2009). Also contributing to the decline
of native crayfish in Ontario is the spread of Small-
mouth Bass (Micropterus dolomieu) and the non-native
Rusty Crayfish (Orconectes rusticus) (Edwards et al.
2009; Phillips et al. 2009).

The Rusty Crayfish has moved or been transported
(via bait buckets) into C anadian waters from the north-
ern limits of its natural range in the Ohio River basin
of the United States (Rosenburg et a/. 2010*). In other
parts of the Laurentian Great Lakes basin, the major
methods of Rusty Crayfish introduction include the re-

lease from bait buckets by recreational anglers, the inten-
tional release by aquarium hobbyists, and their intro-
duction by lake users to control nuisance weeds (Olden
et al. 2011). Where introduced, the Rusty Crayfish has
caused dramatic changes to aquatic ecosystems includ-
ing the replacement of native crayfish, damage to mac-
rophyte beds, and shifts in macroinvertebrate and fish
assemblages (Phillips ef a/. 2009).

The Rusty Crayfish was first reported in Canada dur-
ing the 1960s in Lake of the Woods in northwestern
Ontario and a small number of south-central Ontario
lakes (Crocker and Barr 1968). It has subsequently been
captured in numerous lakes and rivers in other regions
of the province (Berrill 1978: Momot 1996; Edwards
et al. 2009). Although trends in the status of native and
non-native crayfish have been monitored across hun-
dreds of south-central Ontario lakes (Edwards et al.
2009: Somers and Reid 2010*), a corresponding pro-
gram has not been developed for Ontario streams and
rivers. Given the labour-intensive nature of removing
the Rusty Crayfish, a highly fecund species, the like-

111

lihood of successful eradication or control is largely
dependent on early detection (Hamr 2010; Lieb et al.
201 1a). However, the absence of a standardized,
coordinated monitoring program prevents a defensi-
ble assessment of the status of native crayfish and the
impact of Rusty Crayfish (and another non-native
species, Allegheny Crayfish, O. obscurus) in flowing
waters and limits opportunities for timely remedial
action at newly invaded locations.

Selection of sarnpling method is an important part
of the design of monitoring programs. In this study,

we assessed the effectiveness of three methods of

stream sampling (backpack electrofishing, hand cap-
ture, and seining) in capturing Rusty Crayfish. These
methods were selected because of their past use to sur-
vey stream crayfish assemblages (Jezerinac 1991;
Guiasu et al. 1996; Sibley and Néel 2002; Heath er
al. 2010), their current use to monitor stream fish in
Ontario (Stanfield 2005*; Portt et a/. 2008*), and their
suitability for sampling in habitats where wading is
possible. Baited traps are often used to sample cray-
fish in Ontario (Guiasu et al. 1996; Somers and Reid
2010*). However, traps were not evaluated in this
study as they require repeat site visits, are vulnerable
to vandalism or theft (Bernardo et al. 2011), and their
deployment may be impractical in shallow or fast-
flowing water.

A removal-based sampling strategy (depletion) was
used to assess the effectiveness of each method in cap-
turing and characterizing the abundance of native cray-
fish and Rusty Crayfish and differences in probability
of capture related to size and reproductive form. Re-
moval-based strategies have been used successfully to
assess the efficiency of techniques for sampling stream-
dwelling crayfish (Rabeni et al. 1997; Alonzo 2001)
and fish (Amiro 1990; Reid et al. 2009). Unlike mark-
recapture methods, they also do not require multiple
sampling visits.

Study Area

The study was undertaken along six streams in the
Kawartha Lakes region (44°18'N, 78°19'W) — Fleet-
wood, Jackson, Jennings, Meade, Riverview, and
Thompson creeks — and five streams in the Muskoka/
Haliburton Lakes region (45°9'N, 79°4'W) — Dickie
Lake outlet and Blairhampton, Cinder, Coca-Cola, and
Moot Lake creeks.

Sampling locations — ten sites in the Kawartha
region and five in the Muskoka/Haliburton region —
were selected using recent Rusty Crayfish collection
records (EDDMapSOntario 2014*). They represent a
range of water temperatures (mean 16.2°C, range 6°—
25°), conductivity (mean 353 us/em, range 8-990 is/
cm), channel width (mean 5.3 m, range 1—13.6 m),
and streambed materials. Seven of the ten crayfish
species reported to occur in Ontario are found in these
regions. Of those not found, two (Devil Crayfish, Cam-
barus diogenes, and Digger Crayfish, Fallicambarus
fodiens) are obligate burrowers (Crocker and Barr

2 THE CANADIAN FIELD-NATURALIST

Vol. 128

1968); thus, the effectiveness of sampling methods for
these species must be tested separately (e.g., Ridge e/
al, 2008).

Methods

Fifteen removal-based samples were acquired for
each sampling method. Electrofishing and hand-capture
were undertaken along reaches of riffle and shallow-run
habitat. For these two methods, unit sampling distance
was set at 20 m. Seining was undertaken in deeper-run
and pool habitats, where this method is more suitable.
A 10-m unit sampling distance was set for seining.
This shorter distance was chosen because of the limited
size of pool and deeper-run habitats in these streams
and the typical extent of habitat sampled with a seine
during stream fish inventories (Bonar ef al. 2009).
The mean area of stream sampled was: electrofishing
110 m?* (range 31-228 m2); hand capture 79 m? (range
22-211 m’); and seining 65 m? (range 24-112 m2). The
dominant material in streambeds at sites where sein-
ing was carried out was typically finer (clay and sand)
than at electrofishing and hand-capture sites (gravel,
cobble, and bedrock). Sampling occurred between 29
July and 19 October 2010 and was completed before
Rusty Crayfish initiated burrowing activity associated
with winter hibernation (Hamr 2010).

At each unit, the sampling area was isolated with
3.2-mm mesh block nets. Electrofishing was under-
taken with a Smith-Root Type 12A backpack electro-
fisher (pulsed DC settings: 300-400 V, 50-60 Hz.
4-6 ms; Smith-Root Inc., Vancouver, B.C.); one or two
people, depending on channel width, used nets to pick
up crayfish; and sampling rate was 10 s/m2. Hand cap-
ture involved two or three people moving upstream,
overturning rocks, and collecting crayfish by hand or
using an aquarium net (Hamr 2007*). Seining was car-
ried out by two or three people pulling a bag seine (3.2-
mm mesh bag with 4.8-mm mesh wings) upstream. A
minimum of three passes were made for each sampling
unit. At each unit, effort was standardized among sam-
pling passes. Additional sampling passes were made if
a decline in crayfish catch (i.e., depletion) was not ob-
served. Overall, the mean number of passes for each
sampling method was similar (electrofishing 4.0; hand
capture 4.1; and seine 4.3).

Crayfish captured at each pass were held in separate
bins until processed. Individual crayfish were identi-
fied to species (Crocker and Barr 1968) and reproduc-
tive form: female, male form | (sexually active), and
male form II (sexually inactive). For all crayfish, cara-
pace length (CL) was measured to the nearest 0.1 mm
and injuries (e.g., missing chelae) were noted. Batch
weight of each species was measured to the nearest
0.01 g for each pass. Photographs were taken and
voucher specimens were preserved in 70% ethanol for
later confirmation of field identification. Voucher spec-
imens were not retained after species identification
was confirmed.

2014

Data analysis

Population size (number of individuals) and capture
probability were estimated for all crayfish species cap-
tured as well as for Rusty Crayfish using multiple-pass
data and the maximum weighted likelihood method
(Carle and Strub 1978). Catches of individual native
species were low and inconsistent and, therefore, not
suitable for analysis. The efficiency of sampling equip-
ment is affected by crayfish size and sex (Alonzo 2001;
Ogle and Kret 2008); therefore, estimates of capture
probability were derived for each reproductive form
of Rusty Crayfish and for four classes of CL (< 10 mm,
11-20 mm, 21—30 mm, and > 30 mm). Although small
mature individuals have been reported, Rusty Crayfish
with CL less than 20 mm are generally considered im-
mature (Hamr and Berrill 1985; Hamr 2010). Multiple-
pass data were analyzed with Removal Sampling (ver-
sion 2) software (Seaby and Henderson 2007*). The
constant probability of capture assumption was tested
using a y°-based statistic (Seber 1982). Using the De-
Lury method (Ricker 1958*), capture probability was
also estimated from the total biomass of all species of
crayfish collected and the biomass of Rusty Crayfish.

Sampling events were assessed based on whether a
decline in catch was observed with successive passes
(i.e., depletion); capture probability; and e%, a meas-
ure of the precision of population estimates (calculat-
ed as 95% confidence interval*100/N) (Penczak and
Romero 1990). Penczak and Romero (1990) proposed
the following four-point scale to assess e%: 1, very
good estimates (< 10%); 2, good estimates (11-25%);
3, adequate estimates (26-50%); and 4, poor estimates
(> 51%).

As seining was used in a different habitat type, sta-
tistical comparisons among methods were limited to
electrofishing and hand capture. Between-method dif-
ferences in frequency of depletion were tested using
Fisher’s exact test (Sokal and Rohlf 1995). Differences
in capture probability and the representation of each sex
(proportion of Rusty Crayfish males in each sample;
Alonzo 2001) were tested with the unpaired ¢test.
Percentage data were arc-transformed before analysis.
Graphic inspection of length frequency distributions

REID AND DEVLIN: SAMPLING RUSTY CRAYFISH IN ONTARIO STREAMS

113

and the Kolmogorov-Smirnov (K-S) test (based on sep-
arate pooled datasets for all crayfish species and for
Rusty Crayfish) were used to assess differences in the
size of crayfish captured (Zar 1984; Dorn ef al. 2005).

Results

Six species of crayfish were collected from stream
sites: Appalachian Brook Crayfish, Cambarus bar-
tonii (n = 131); Calico Crayfish, O. immunis (n = 2);
Allegheny Crayfish (7 = 1); Northern Clearwater Cray-
fish, O. propinquus (n= 32); Rusty Crayfish (1 = 2082);
and Virile Crayfish, O. virilis (n = 1). The Rusty Cray-
fish was collected from 65% of sampling units, com-
prised 90% of all crayfish captured and 85% of the
crayfish biomass, and was the only species collected
from four of the 11 streams sampled (Table 1). It was
found in all Kawartha Lakes region streams except
for Fleetwood Creek, where only a single Allegheny
Crayfish was captured. Despite its presence in near-
by lakes, the Rusty Crayfish was not collected from
any Muskoka/Haliburton Lakes region streams. Only
native crayfish species (Appalachian Brook Crayfish,
Northern Clearwater Crayfish, and Virile Crayfish)
were collected from these streams.

For all sampling methods, the Rusty Crayfish was
typically collected during the first sampling pass (>
88% of sites where present). Overall, electrofishing
resulted in the capture of all six species, whereas Calico
Crayfish and Allegheny Crayfish were absent from
both hand-capture and seining samples. The Appalachi-
an Brook Crayfish was not collected during seining,
and capture of the Northern Clearwater Crayfish and
the Virile Crayfish required multiple hauls.

Seining was the least time-intensive method, requir-
ing on average 1.2 min/m? for sampling and process-
ing. Hand capture was the most time intensive at 3.9
min/m2, while electrofishing required 2.6 min/m”.

Population size

Electrofishing always resulted in crayfish capture.
In contrast, no crayfish were collected at three hand-
capture sites and five seining sites. For 53% of sam-
ples, we were able to estimate total crayfish population
(all species) and capture probability. In most other cas-

TABLE 1. Comparison of catch data for Rusty Crayfish, Orconectes rusticus, using three sampling methods in 11 Ontario

creeks.

Variable Electrofishing Hand capture Seining

No. sites yielding Rusty Crayfish 10 9 7

Total number captured 918 109] 73

Females (%) 458 (49.9) 548 (50.2) 35 (48.0)

Form I males* (%) 174 (19.0) 158 (14.5) 22 (30.1)

Form II males* (%) 258 (28.1) 380 (34.8) 16 (21.9)
‘ SS5 9]

Total biomass (g) ' 2739 2555 332

an carapace length (mm [SE])
fae ‘ 19.6 (0.30) 17.6 (0.24) 24.7 (1.16)
Form I males 23.0 (0.37) 24.3 (0.32) 25.8 (1.05)
1731027) 16.0 (0.19) 16.6 (1.15)

Form II males

*Form I male = sexually active, form II male = sexually inactive.

0.2

05 7 —— — 4
All Crayfish er
—s—HC
04 ] ‘A SN
034 ,
i 02 a YY
Bay i ‘
Tv
c
o 014
c
:
E of
5 1 2 3
2
5
= 05
i] ——OHa=
° O. rusticus a
= —a— HC
° 04 *&*: SN
c
5 I.
5 a
03 sak
Qa *.
: attas vail cee Sac

0.1

|
|
|
|
ee

0

Pass

FIGURE |. Proportion of total catch of crayfish of all species
(above) and Rusty Crayfish, Orconectes rusticus
(below) captured during the first three passes using
three removal-based sampling methods in 11 Ontario
creeks: electrofishing (EF), hand capture (HC), and
seining (SN). Symbols indicate mean values and ver-
tical lines represent standard errors of the mean.

es, either too few individuals were captured (42% of
failed estimates), or there was no decline in catch with
successive passes (46% of failed estimates) (Figure 1).
There was no significant difference in the percentage
of samples in which depletion occurred with succes-
sive passes between electrofishing (60%) and hand cap-
ture (66%) (exact test, P = 1.0). Depletion occurred
less frequently (33%) in seined samples.

Estimates of total crayfish population were highest
for hand-captured samples and lowest for seining
(Table 2). In contrast, mean capture probability was
highest for seining (Table 2). There were no signifi-

TABLE 2. Estimates of population size, in terms of abundance

THE CANADIAN FIELD-NATURALIST

Vol. 128

cant differences in capture probability between elec-
trofishing and hand capture (¢ = 0.23, P = 0.82). All
estimates resulted in capture probability > 0.2, and,
therefore, are considered valid (White ef al. 1982*).
However, capture probabilities for electrofishing and
hand capture were below the level considered to pro-
vide consistently reliable results (= 0.4, Seber 1982).
Based on e%, most electrofishing (83%) and hand-
capture (86%) estimates were considered at least ade-
quate; 60% of seining estimates were poor.

For 63% of sampling events at Rusty Crayfish sites,
we were able to estimate population size and proba-
bility of capture. There was no significant difference
in the frequency of depletion between electrofishing
(60%) and hand-capture samples (70%) (P = 0.63),
and the frequency of depletion (60%) was similar at
seining sites. Differences among methods and estimates
of population size and capture probability were simi-
lar for Rusty Crayfish and all crayfish species (Table
2). There were no significant differences in capture
probability between electrofishing and hand capture
(t = —0.57, P = 0.58). Based on e%, most estimates
based on electrofishing (78%), hand capture (80%),
and seining (60%) were considered at least adequate.

Biomass estimates were less suitable for compar-
ing sampling methods than abundance data. For both
Rusty Crayfish and all species, only 30% of sampling
events permitted estimates of population size and cap-
ture probability based on biomass. However, capture
probabilities for electrofishing and hand capture were
similar to those derived from abundance estimates
(Table 2).

Reproductive form

For Rusty Crayfish in all samples, the sex ratio was
close to 1:1. Males captured by electrofishing and hand
capture were more often form II, whereas seining
resulted in the capture of more form I males (Table 1),
There was little difference in the overall proportion of
females captured by the three methods. No si gnificant
difference was detected in the proportion of males col-
lected by electrofishing and hand capture (t = 0.06, P
=):

For 40% of samples, we were able to estimate cap-
ture probability by reproductive form (Table 3). For

and biomass, and capture probabilities for all crayfish species

and for Rusty Crayfish, Orconectes rusticus, based on three sampling methods used in 11 Ontario creeks: electrofishing

(EF), hand capture (HC), and seining (SN).

Population estimate (mean [SE])

lee HC

Capture probability (mean [SE])

SN EF HC SN
Abundance (no./m?)
Allcrayfish species 1.04 (0.31) 2.48 (1.12) 0.11 (0.05) 0.30(0.04) 0.31 0.04) 0.56 (0.13)
Rusty Crayfish 1.30 (0.41) 4.20 (1.80) 0.11 (0.04) 0.29 (0.03) 0.26 (0.04) 0.47 (0.05)
Biomass (g/m?)
All crayfish species 5.08 (1.65) 5.16 (2.02) 0.65 (0.48) 0.34(0.08) 0.33 (0.07) 0.74 (0.26)
Rusty Crayfish 4.54 (1.51) 3.92 (2.03) 0.17 (n/a)* 0.33 (0.05) 0.40(0.06) ‘1.0 (/ay*

Wek Li 2
*Standard error of the mean (SE) not applicable as only one sample obtained.

2014

REID AND DEVLIN: SAMPLING RUSTY CRAYFISH IN ONTARIO STREAMS

TABLE 3. Probability of capture by reproductive form and carapace length of Rusty Crayfish, Orconectes rusticus, associat-
ed with three stream sampling methods in 11 Ontario creeks: electrofishing, hand capture, and seining.

Capture probability (mean [range])

Electrofishing

Reproductive form

Form I male*

Form II male*

Female
Carapace length

< 10.5 mm

10.5—20.4 mm

20.5-30 mm

> 30 mm

0.34 (0.26—0.54)
0.46 (0.25—0.66)
0.16 (0.06—0.28)

0.42 (0.38-0.45)
0.10 (0.01—0.19)
0.45 (0.27—0.62)
0.33 (0.10—0.49)

Hand capture Seining

0.36 (0.33-0.39)
0.36 (0.10—0.38)
0.43 (0.14—-0.57)

0.267
0.20 (0.02—0.39)
0.34 (0.32—0.35)

0.537 n/a
0.45 (0.19-0.62) 0.02*
0.32 (0.09-—0.44) ().42 (0.29-—0.54)
0.70 (0.68—0.71) 0.62+

*Form | male = sexually active, form I] male = sexually inactive.

+Single sample.

electrofishing and hand capture, a decline in catch was
most frequently associated with form I males (63% and
50% of cases). Alternatively, seining declines occurred
least often for form I males (17% of cases). Although

50 Electrofishing

§ 10 15 20 2 30 35 40 4

50 Hand capture

5 10 15 20 25 30 35 40 45

Percentage of crayfish collected
Ww
oO

5 10 15 20 25 30 35 40 45
Carapace length (mm)

FIGURE 2. Distribution by carapace length of all crayfish
species (black bars) and Rusty Crayfish, Orconectes
rusticus (grey bars) captured by electrofishing, hand
capture, and seining in 11 Ontario creeks.

based on a small number of samples, capture probabili-
ties for male Rusty Crayfish were lower during seining
than the other two methods, and for females lower dur-
ing electrofishing (Table 3). There were no obvious dif-
ferences among reproductive forms in capture proba-
bility by hand capture or seine. For electrofishing,
capture probabilities tended to be greater for males
(Table 3).

Carapace length

A broad range of sizes of all crayfish species (CL
range 543 mm) was collected during stream sampling
(Figure 2). Electrofishing and hand capture resulted in
a similar range of sizes, which included a greater pro-
portion of small individuals (< 25 mm) than seining.
However, CL distributions for samples collected by
electrofishing and hand capture were significantly dif-
ferent (all species: D = 0.15; P < 0.001; Rusty Crayfish:
D = 0.13; P < 0.001). Overall, hand-capture samples
included a greater percentage of small individuals (CL
15-25 mm) than electrofishing.

For a third of samples, we were able to estimate cap-
ture probability for individual length classes. For hand
capture and seining, a decline in catch was most fre-
quently associated with larger Rusty Crayfish (CL
20-30 mm and > 30 mm). For electrofishing, declines
occurred least often for the largest crayfish (CL >
30 mm), in only 18% of cases. Although based on a
small number of samples, capture probabilities for most
Rusty Crayfish length classes during hand capture were
higher than other methods (Table 3). For electrofish-
ing, capture probabilities were greatest for the smallest
(< 10.5 mm) and the 20.5-30 mm length classes of
Rusty Crayfish (Table 3). For the other two methods,
differences in capture probability among length classes
were also apparent. However, small sample sizes pre-
vent meaningful comparisons.

Sampling injury

Cheliped loss was recorded for 8.8% of all crayfish
collected by electrofishing, 6.3% by seine, and 5.4% by
hand capture. Observed mortality was 1.4% for elec-
trofishing and 2.2% for hand capture. No dead crayfish

116

of any species was observed during seining. Most dead
crayfish were found either crushed or with a cracked
carapace.

Discussion

All three sampling methods assessed in this study
were effective at detecting Rusty Crayfish. Although
multiple-pass sampling may improve detection of na-
tive species, it was not a reliable approach for estimat-
ing capture probabilities or population sizes of Rusty
Crayfish and other crayfish species. In most cases, fail-
ure of the removal model was observed as an increase
in the number of crayfish captured after the first pass
or too few individuals collected. The first sampling pass
may disturb or draw out crayfish from areas of cover
making them more vulnerable to capture in subsequent
passes (Gladman ef al. 2010). Removal sampling by
both electrofishing and hand capture has been suitable
for generating population estimates of other stream-
dwelling crayfish (Rabeni et a/. 1997; Alonzo 2001);
however, in those studies, sampling was carried out in
much smaller streams and channel width can have a
strong influence on efficiency of a method (Zalewski
and Cowx 1990). To improve catch efficiency, Alonzo
(2001) also used low voltage and activated the electrode
for only 1-2 seconds at a time. In our study, we used
voltage output and sampling strategies typical of stream
fish inventories in Ontario. Although affected by elec-
trical current, crayfish do not display the same degree
of galvanotaxis as many stream fish. Experimentation
with different settings could improve our capture effi-
ciency.

Electrofishing and visual sampling methods are often
biased toward capture of large individuals and against
cryptic taxa or life stages (Zalewski and Cowx 1990).
Activity levels and use of cover and deeper habitats
vary among crayfish species, sizes, and reproductive
forms (Berrill and Arsenault 1982; Guiasu 1997; Davis
and Huber 2007) and, therefore, can also be expected
to affect capture probabilities. For electrofishing, Alon-
zo (2001) reported a difference in the probability of
capturing small and large crayfish in small streams, but
none between sexes. In our study, the low number of
removal estimates prevented robust comparisons of
capture probability among reproductive forms and
length classes. However, capture probabilities were
generally higher for males during electrofishing and
for the smallest and largest crayfish during hand cap-
ture. All three methods collected crayfish of different
sizes, with seining providing a greater representation
of large individuals. The large size of Rusty Crayfish
associated with seine hauls from pools is consistent
with Davis and Huber (2007) who observed that large
Rusty Crayfish prefer deeper water than smaller indi-
viduals.

Although changes in the size of native crayfish pop-
ulations after Rusty Crayfish introduction into lakes

THE CANADIAN FIELD-NATURALIST

Vol. 128

have been thoroughly documented, research on such
declines in streams and rivers has been less intensive
(Jezerinac 1982, 1991; Daniels 1998). Recent stream
surveys in the mid-west and eastern regions of the Unit-
ed States indicate the continuing spread of Rusty Cray-
fish and concurrent declines in native Orconectes spe-
cies (Kuhlmann and Hazelton 2007; Kilian e¢ al. 2010;
Lieb et al. 2011b; Olden et a/. 2011). Based on a broad-
scale survey of the Kawartha Lakes region, Berrill
(1978) found the Rusty Crayfish to be widespread and
common and indicated that it was likely replacing
Northern Clearwater Crayfish. At our Kawartha Lakes
stream sites, the Rusty Crayfish was the dominant (and
often only) crayfish species present, indicating that the
shift in species composition in this region has per-
sisted for several decades. Jezerinac (1982) reported
that location within the watershed influenced the effect
of the Rusty Crayfish, with Northern Clearwater Cray-
fish abundant in headwaters and small tributaries (where
few Rusty Crayfish were present) and absent or in low
numbers in the main stream channel (where Rusty
Crayfish were abundant). Although the expanding dis-
tribution of the Rusty Crayfish in southern Ontario has
begun to be tracked, factors influencing its effect on
native crayfishes in flowing waters are not well under-
stood and require greater attention.

As the likelihood of species detection is improved,
the use of multiple sampling methods has been recom-
mended for crayfish surveys (Holdich er al. 2002).
Backpack electrofishing and seining provide the best
combination of methods to detect the Rusty Crayfish,
native species, and a broad range of sizes of crayfish
in Ontario streams. Although electrofishing was most
effective at detecting both native and non-native spe-
cies, seining is more suitable for sites that are turbid,
deep, or have substrates too soft for effective electro-
fishing. Hand capture (or hand searching) does not re-
quire expensive equipment and is associated with few-
er safety concerns than electrofishing. However, it was
the most labour-intensive method, resulted in the high-
est rate of mortality, and is less readily adaptable to
current stream fish monitoring programs. Gladman ef
al. (2010) and United States National Park Service
(2007*) both reported hand capture to be less efficient
than electrofishing for sampling stream crayfish.

As applied in this study, multiple-pass sampling was
not a reliable strategy to estimate crayfish population
size. However, multiple passes are still preferable to
single-pass sampling as this improves native species
detection (Gladman et al. 2010), and a large percent-
age of individuals will be vulnerable to capture only
after being disturbed by initial sampling efforts. If esti-
mates of population size are required for stream reach-
es, the likelihood of failure could be reduced by pooling
data from randomly distributed sampling sites and ap-
plying unbiased removal-type estimators (Heimbuch
et al. 1997). In this study, we did not estimate detection

2014

probabilities for native crayfish. The design of stream
monitoring efforts would benefit from additional stud-
ies that apply repeat survey designs (MacKenzie et al.
2002) to estimate method-specific detection probabil-
ities.

Acknowledgements

This research was supported by the Ontario Ministry
of Natural Resources and the Invasive Species Partner-
ship Fund. J. Ciampa, S. Hogg, and M. Parna assisted
with sampling. J. Rusak and K. Somers assisted with
crayfish identification. The manuscript was improved
with suggestions from D. McAlpine, K. Somers, E.
Snyder, and two anonymous reviewers.

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Supplementary material available at:
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Received 12 September 2013
Accepted 9 February 2014

TABLE S1. Crayfish catch data from each south-central Ontario stream sampling unit. For each method, electrofishing (EF).
hand capture (HC), and seining (SN), the number of sampling passes is provided in parentheses. Note: Cb = C bartonii
(Appalachian Brook Crayfish), O1 = O. immunis (Calico Crayfish), Ob = O. obscurus (Allegheny Crayfish), Op = O
propinquus (Northern Clearwater Crayfish), Or = O. rusticus (Rusty Crayfish), and Ov = él virilis (Virile Crayfish).

TABLE S2. Summary of catch data for Appalachian Brook Crayfish (Cambarus bartonii) and Northern Clearw
(Orconectes propinquus) captured from Ontario streams using three methods: electrofishing, hand capture

ater Crayfish
and seining.

Estimating Breeding Bird Survey Trends and Annual Indices for

Canada: How Do the New Hierarchical Bayesian Estimates Differ
from Previous Estimates?

eed _ * a
ADAM C, SmituH!:?, MARIE-ANNE R. HUDSON!, CONSTANCE Downes!, and CHARLES M. FRANCIs!

Canadian Wildlife Service, Environment Canada, National Wildlife Research Centre, 1125 Colonel By Drive, Ottawa, Ontario
K1A 0OH3 Canada
“Corresponding author: Adam.Smith@ec.ge.ca

Smith, Adam C., Marie-Anne R. Hudson, Constance Downes, and Charles M. Francis. 2014. Estimating breeding bird survey
trends and annual indices for Canada: how do the new hierarchical Bayesian estimates differ from previous estimates?
Canadian Field-Naturalist 128(2); 119-134.

Canadian data from the North American Breeding Bird Survey (BBS) provide information on the population status and trends
for over 300 species that regularly breed in Canada. Since the first assessments were made in the mid-1970s, both the dataset
and the suite of statistical tools and techniques available to researchers have grown. As a result, Canadian BBS trend esti-
mates have been derived from numerous statistical models. Because the BBS data are relatively complex, different statistical
models can generate different trend and status estimates from the same data. In 2013, Environment Canada’s Canadian
Wildlife Service began producing BBS status and trend estimates using a hierarchical Bayesian model. To give users of BBS
trends and annual indices of abundance a better understanding of these estimates, we demonstrate and explain some of the
similarities and differences between the new hierarchical Bayesian estimates and those from the previous model; discuss the
philosophical and inferential consequences of estimating trends with the new model; and describe how the hierarchical Bayesian
model differs from the model currently used in the United States. Overall, trends and annual indices from the new model are
generally similar to estimates from the previous model; however, they are more precise, less variable among years, better represent
the spatial variation across Canada in population status, and allow for more intuitive and useful assessments of uncertainty.

Key Words: breeding bird survey; hierarchical Bayesian model; maximum likelihood model; population trend; population

monitoring

Introduction

The North American Breeding Bird Survey (BBS)
provides information on the population status (trends
and annual indices of abundance) of almost 300 bird
species in Canada. Indeed, it is the primary source of
information for over 200 of those species. Due, in part,
to its long history and continental coverage, the BBS
is considered to be the backbone of land bird conser-
vation in North America (Rich et a/. 2004*). It is a vital
component of a wide variety of high-level conservation
documents for Canada, e.g., the State of Canada’s Birds
(North American Bird Conservation Initiative Canada
2012*) and the Status of Birds in Canada website
(Environment Canada 2011*). Data from the survey
have also been used in hundreds of scientific publica-
tions on topics ranging from assessing population
trends to modelling habitat associations or impacts of
climate change on range shifts (Pardieck er a/. 2008).

The approaches used to analyze BBS data have
evolved over time as new statistical methods have be-
come available and as the dataset has grown to support
increasingly complex models. The first Canadian BBS
routes were conducted in 1966, and the first national
trend estimates included the data from the first 10 years
(Erskine 1978*). Population trends in that report were
estimated using a chaining-style analysis of averaged
year-to-year changes in species counts on routes run in
successive years, with the chain of year-to-year changes

indexed to a value of 100 in a base year (1973). Sub-
sequently, both the Canadian Wildlife Service (CWS)
and the United States Geological Service (USGS) began
to analyze the BBS data annually, but using different
methods. Early analyses by the United States’ agencies
(at the time, the Fish and Wildlife Service) calculated
annual indices using weighted average counts across
years and trends using route-regression models (Rob-
bins et al. 1986). Through the 1980s and 1990s, various
route-regression models (Geissler and Noon 1981) were
used by both the CWS and USGS, although variation in
approaches to weighting routes led to different estimates
(Thomas and Martin 1996). The USGS analyses con-
tinued to use route-regression models through 2008,
while, from 2002 to 2009, Canadian BBS data were
analyzed using a maximum likelihood (ML) model,
which estimated an annual index. Starting with the 2011
BBS data, Canadian trends and annual indices have
been estimated using a hierarchical Bayesian (HB) mod-
el, which is very similar to the model adopted by the
USGS in 2009 and described in Link and Sauer (2002)
and Sauer and Link (2011).

HB methods suit the BBS data’s complex structure,
because the Bayesian framework provides a coherent
and flexible approach for modelling the effects of sam-
pling variation (e.g., not all routes are surveyed each
year; not all birds present on a route are detected on any
given survey; and detection probabilities vary among

he,

observers) separately from temporal and geographic
variation in the underlying populations (e.g., popula-
tion change due to changing weather, resource abun-
dance, and human activity). In addition, the hierar-
chical structure is particularly efficient and powerful
because the distributional assumptions of the random
effects greatly reduce the number of parameters that
need to be estimated to model observer effects and
other nuisance parameters (Clark 2005). Similarly, the
hierarchical structure of the model mirrors the hierar-
chical structure of the data and allows effects to be
modelled at the appropriate scale, e.g., nuisance par-
ameters, such as overdispersion and the effects of
changes in observers over time, can be efficiently mod-
elled across all counts, while trends and annual varia-
tion around the trend can be modelled across routes
within strata (Link and Sauer 2002).

Bayesian methods also provide intuitive and direct
estimates of the uncertainty around the population
parameters (i.e., population trends and indices of annu-
al abundance) that are of primary interest to most users.
Bayesian methods are based on the idea that our under-
standing of an imperfectly known parameter, such as a
population trend, has a probability distribution. That is,
the mean, median, or mode of the distribution is our
best estimate of the parameter, and the variance, stan-
dard error, or percentiles (1.e., the credible interval) of
the distribution represent the uncertainty around our
best estimate. Our initial understanding of that distri-
bution is termed the prior probability distribution. The
data are used to update the prior probability distribu-
tion, through Bayes’ rule, to produce the posterior prob-
ability distribution, which expresses our final under-
standing of the parameter (Link and Barker, 2009).
For example, with a Bayesian framework, it is relative-
ly simple to estimate the posterior distribution of the
total change in a population since 1970, and from that
distribution, make concise and direct statements about
the probability that a population has declined; where-
as with previous analyses, uncertainty estimates were
largely limited to probabilities of observing the data
(or more extreme data) if the true population had not
changed.

The differences between Bayesian and frequentist
(i.e., traditional) estimates of uncertainty are exempli-
fied in comparing Bayesian credible intervals with fre-
quentist confidence intervals. Credible intervals pro-
vide a probable range of values for a parameter (e.g.,
in the case of a 95% credible interval, the range of val-
ues that contains, with a 95% probability, the true val-
ue of a population trend). Many readers will be more
familiar with confidence intervals and likely with inter-
preting them in the same way. However, for confidence
intervals this interpretation is incorrect, and the con-
fusion largely stems from the unintuitive nature of the
true definition of confidence interval, which relates to
an infinite number of hypothetical realisations of the

THE CANADIAN FIELD-NATURALIST

Vol. 128

data (Clark 2005). In essence, credible intervals, and
Bayesian methods in general, provide intuitive and
useful measures of uncertainty. For example, the cur-
rent BBS results website maintained by Environment
Canada (www.ec.ge.ca/ron-bbs) provides, for every
trend estimate, associated estimates of the probabilities
of eight thresholds of population change (¢.g.. proba-
bility that the population has decreased, increased by
> 100%, etc.). Species-at-risk status assessments can
now easily consider an explicit measure of the proba-
bility that a population has reached one of the thresh-
olds used to categorize a species as threatened or
endangered (e.g., COSEWIC 2011*).

The application of HB methods to the BBS repre-
sents the most recent development in estimating the
status and trends of hundreds of bird species across
North America. Three closely related HB models are
described in the literature or are in use for analyzing
trend and annual indices from the BBS data. The first
model (HB,) was described by Link and Sauer (2002).
It was subsequently refined to account for the bias in
retransforming annual index estimates from the log-
scale to the scale of average counts (second equation
in Sauer and Link 2011). The second model (HB,) is
currently used in annual analyses by the USGS. The
third model (HB,) includes an additional refinement
that has the effect of scaling the annual indices more
closely to the average observed count on routes within
a stratum and improving estimates of trends for larger
regions. HB, is the model currently used in the CWS
annual analyses. The distinction between models HB,
and HB, is important because trend and annual index
estimates from the USGS include Canada and Canadi-
an provinces. Therefore, anyone interested in a species
trend for Canada must choose between the estimates
derived by the USGS (using model HB,) and those
derived by the CWS (using model HB,). -

Because the BBS data and the statistical models
used to analyze them are complex, different analytical
approaches can result in different estimates of popu-
lation parameters, particularly if the underlying signals
in the data are relatively weak or variable or if the
sampling is imbalanced. The BBS sampling method is
a stratified random design that aims to generate a bal-
anced and approximately representative sample of bird
populations across North America (Erskine 1978*).
Of course, as with any field survey, practical constraints
introduce some variation and bias into the data: uneven
spatial distribution of active routes, changes in the num-
ber and spatial coverage of routes surveyed over time,
and variation among and changes over time in ob-
servers. The nested structure of the data (i.e., stops nest-
ed within routes, routes selected within degree-blocks
and aggregated into analytical strata) provides added
challenges. Analyses of BBS data have different ways
of accounting for all or some of these sources of vari-
ation and bias, and, as a result, different analyses can

2014

sometimes generate different estimates of population
trends and annual indices (e.g., Thomas 1996: Thomas
and Martin 1996; Sauer and Link 2011).

For many species and regions, the underlying sig-
nals in the data are clear and strong, the sample sizes
are large, or the BBS sampling is very well balanced in
space and time. In these cases, even very different mod-
els will generate similar estimates of population status
and trends. In addition, although point estimates of
trend for a given species and region may differ among
models, most conservation or management decisions
and actions are based on relatively broad overall pat-
terns in population status. For example, the Status of
Birds in Canada website classifies species based on six
categories of population status, such as large decrease,
moderate increase, little change, and data deficient
(Environment Canada 2011*). Each of these categories
encompasses a relatively wide range of trend values;
thus, even when point estimates of trend or trajectory
differ among models, the broad population inferences
may be similar, particularly for populations with rela-
tively extreme status (i.e., large decrease or large in-
crease), which are likely to be the primary focus of
management.

Our three goals in this paper are to describe in detail
the HB, model we are now using in Canada to estimate
population trends and how it differs from the HB, mod-
el used by the USGS; to demonstrate and explain some
of the similarities and differences in the estimates pro-
vided by the HB, model and the previous Canadian
model (the ML model); and to discuss the most impor-
tant philosophical and inferential consequences of esti-
mating trends and annual indices of abundance with
the HB, model.

Methods
Maximum likelihood model

The model used to generate Canadian BBS trends
between 2002 and 2009 is a weighted ML model (see
Appendix 1). It uses a series of equations to derive ML
estimates of stratum and year-specific annual indices

(A;;):
Aj = exp(u +a; t+ Bi + Vj x)

The model assumes counts of birds on BBS routes
are Poisson distributed, and estimates terms to account
for the effects of each year (/3,), variation among routes
(a), and variation among observers and observation
conditions in each year (7, ,). The means of the esti-
mated effects of year, route, and observer are fixed at
0, and x is not estimated, but held constant as the ob-
served average count of birds across all years and
routes. The analysis is conducted using the sampling
strata of the survey: degree blocks (regions defined
by one degree latitude and one degree longitude, Fig-
ure 1 inset). The ML estimation of annual indices is
weighted so that each degree block with data influ-
ences the estimate in proportion to its area, regardless

SMITH ET AL.: NEW BREEDING BIRD SURVEY TRENDS AND INDICES 12]

of the number of routes with data. Weighting limits the
influence of degree blocks with more routes, which,
because the BBS is volunteer-based, tend to occur in
areas with larger human populations. The weight for
each route from degree block i used in the analysis is
n-| * Ai, where n is the number of routes in the degree
block and 4 is the degree block’s area. The model was
run through a custom C++ program.

The ML model estimates the annual indices direct-
ly, and trends are derived from log-linear regressions
of the annual indices of abundance. A small constant
(0.001) is added to the annual indices before the log-
linear regression, because in years when the species is
not observed the annual index is set to 0. The standard
errors of the abundance indices and trends are esti-
mated using a “jackknife” procedure, which discards
one route at a time. A full description of the model
has never been published; however further details are
provided in Appendix 1.

Hierarchical Bayesian model, HB,

The CWS is now generating Canadian BBS trends
using the HB, model. It assumes that observed counts
on a BBS route / in year ¢ and stratum i have a Pois-
son distribution with mean (),;,). The strata used are
areas created by the intersection of provincial and ter-
ritorial boundaries with those of the North American
Bird Conservation Regions (BCRs; North American
Bird Conservation Initiative International 2013*;
Figure 1). On the log scale, the As are modelled by
fixed effects for first-year observer effects (7, where
the value of I(j,f) is 1 if a route-year combination
represents an observer’s first year on that route and 0
otherwise) and stratum-specific fixed effects for abun-
dance (S;), trend over time (f;), and variance of the
year effects within each stratum (0; ). The model also
has random effects for overdispersion (¢; ;,) of counts
and stratum-specific random terms for year effects (); ,)
and observer—route effects (,; ;).

log (Ai) = Sj oe B; *t+ Vint ar Wij + nl, t) + Ei j
[equation 1]

All prior distributions are standard, diffuse (non-
informative) conjugate priors; that is, the priors are
chosen so that the estimates are not influenced by any
prior knowledge—non-informative—and the specific
distributions, such as the inverse gamma for variances,
ensure proper posterior distributions—conjugate pri-
ors. Specifically, all variances are assigned diffuse
inverse gamma prior distributions (scale and shape
parameters set to 0.001) and the parameters S, /, y,,
and 7 are given diffuse normal distributions (mean 0,
variance 10°).

The annual indices of abundance for stratum i and
year ¢ are exponentiated sums of the year, stratum,
and trend effects estimated in equation 1, scaled by
the proportion of routes in the stratum on which the
species was observed (z;).

= mt?
[22 THE CANADIAN FIELD-NATURALIST Vol. 128

0 250 500 1,000 Km

AS
V/s
\

|

_

~Raris

A

Ir

fs fee le eg |

= See

i

71°W 70°W 69°W 68°WSO67°W~séGS&G°W_Ss«é6 SW

FiGureE 1. Geographic strata for analysis of Breeding Bird Survey (BBS) data using the Canadian hierarchical Bayesian

model. The strata are the areas created by the intersection of provincial and territorial boundaries with those of Bird
Conservation Regions (BCRs). Exceptions are the strata consisting of all of BCR 7, which crosses several provinces
and territories, and all of BCR 14, which includes Nova Scotia and Prince Edward Island. BCR borders are indicated
by heavy black lines: BCR 3, Arctic Plains and Mountains; BCR 4, Northwestern Interior Forest: BCR 5. Northern
Pacific Rainforest; BCR 6, Boreal Taiga Plains; BCR 7, Taiga Shield and Hudson Plains: BCR 8. Boreal Softwood
Shield; BCR 9, Great Basin; BCR 10, Northern Rockies; BCR 11, Prairie Petholes;: BCR 12. Boreal Hardwood
Transition; BCR 13, Lower Great Lakes/St. Lawrence Plain; and BCR 14, Atlantic Northern Forest. Within provinces
and territories, strata are indicated by different shades of grey. Regions filled with diagonal lines have insufficient
BBS data to be included in any analyses. The inset shows the original sampling strata for the BBS, i.e., degree-blocks,
overlaid on the province of New Brunswick.

2014

Nae Says editBitt+Vict0.5+o% +0.5¥a2

[equation 2]
The variance components (0.5 * 63.+ 0.5 * 62) are
added to correct for retransformation bias: the expo-
nent of the average of a normal distribution — the
sum of S; + fi * t+ Y,, — underestimates the average
of the log-normal distribution (Newman 1993), The
variance components ensure that the indices are scaled
to an average number of birds per route. Annual indices
for larger regions, i.e., provinces, territories, BCRs, or
country, are area-weighted sums of the stratum-level

estimates:

N = LAENi
A

[equation 3]

Trends in the HB, model are estimated as geomet-
ric mean annual changes in population size over spe-
cific intervals, expressed as a percentage. That is, the
trend B from year ¢, to year ¢, for a given region is:

1

B= 100+ (Se) 4
N

ta

[equation 4]

The HB, model described here is very similar to
the HB, model used by the USGS (described in Sauer
and Link 2011) with one modification: the HB, mod-
el estimates a common observer-route variance across

>

all strata (62 instead of o3,). In the HB, model, the
stratum-specific observer—route variances (Gj,) are
drawn from a hyperdistribution that allows the vari-
ance of observer—route effects to differ among strata
and yet be estimated relatively efficiently, even for
strata with relatively few observer—route combinations.
The hyperdistribution is defined such that the preci-
sion (the inverse of the variance) of stratum-specific
observer—route effects (1/o2,), is assumed to come
from a log-normal distribution with a common mean
and variance, i.e., log(1/o2,) = Mu,),o2). In compari-
son, the HB, model estimates a single observer—route
variance term (c2) across all observer—route combi-
nations, which is given a diffuse inverse gamma prior
distribution (scale = shape = 0.001). The WinBUGS

code for the HB, model is available in Appendix 2A.

Comparisons between models

We used the HB, and ML models to estimate trends
and annual indices for all species using a common
dataset. The dataset included all observations from
Canadian BBS routes, run from 1970 through 201 IF
under acceptable observation conditions (1.¢., coded as
RunType = 1; USGS 2012*). Although the original
datasets were identical, the models have slightly dif-
ferent inclusion criteria for a given route-year com-
bination. For example, the ML model estimates a route-

SMITH ET AL.: NEW BREEDING BIRD SURVEY TRENDS AND INDICES |

i)
Ww

specific trend parameter that requires a minimum of
2 years of observations on each route, whereas the HB,
model estimates a stratum-specific trend parameter
and, therefore, can include data from routes with only
| year of data. These differences were relevant to very
few route-year combinations (< 1%), so they resulted
in only very small differences in the final estimates.
All species names (English, French, and scientific), as
well as all trend estimates for each species, region, and
trend period, are available in Appendix 3.

For most comparisons, we contrasted estimates from
the two models at three scales. First, the national scale
provides a comprehensive comparison that has rele-
vance to many conservation and management deci-
sions. Second, the BCR-scale comparisons highlight
some of the important spatial characteristics of each
model, e.g., differences in area weights, spatial scope
of inference, and minimum data criteria (Canada’s 11]
BCRs are shown in Figure 1). Finally, a comparison for
one province (New Brunswick; Figure | inset), which
is an identical stratum in both analyses, provides the
most direct comparison. This last comparison highlights
differences between the overall frameworks of the ML
and HB, models, because the data, spatial extent, and
relative area weights are as similar as possible.

We made quantitative comparisons of five charac-
teristics of the two models: (1) the number of species
in each region with trend estimates, (2) the magnitude
of trends, both long term (> 40 years) and short term
(10 years), (3) the precision of trend estimates, (4) the
inter-annual variability of short-term trends, and (5) the
scale of the annual indices. We did not include a com-
parison of the number of statistically significant trends
in the two modeis (i.e., a trend estimate for which the
confidence or credible interval excludes zero). Statis-
tical significance is a synthetic metric that confounds
trend magnitude and precision; therefore, making such
a comparison would add nothing new to the compar-
isons made here.

Magnitude of trends: To compare the magnitude of
the long- and short-term trends, we used an HB mod-
el analogous to a weighted paired f-test that accounts
for the uncertainty of each individual trend estimate.
Extreme trend estimates (i.e., estimates that are larger
in absolute magnitude) are more likely to be imprecise
and would strongly influence a direct comparison of
trends if not weighted by precision. Our comparison
approach was derived from one originally described in
Sauer and Link (2002), which assumes that estimated
trends are part of a collection of trends that share a
common mean and variance. However, we did not
assume a common average trend for each model (as
suggested in Link and Barker 2009) and did not, there-
fore, shrink imprecise estimates toward a common
mean. The comparison model uses trend estimates
(8,,,,, trend estimates for species s from model m) as
data and accounts for uncertainty in the estimation of
those trends using estimates of the variance of each

trend (62,,). It makes pairwise, within-species compar-
isons between the posterior estimates of trend from

each model
(Bs up vai Bs mt)

and compares the overall average species-
differences

specific

a EEE ~Bsmty

The comparison model estimates the average, species-
level difference (1.e., the result of subtracting one esti-
mate from another) in trends between the two models,
weighted by the relative precision of each estimate.
The average of the species-level differences is an esti-
mate of the relative bias in the two models: positive
differences indicate that the HB, trend estimates tend
to be more positive (less negative) than the ML trend
estimates for a given species. In addition to evaluating
bias, we also estimated the correlation coefficient of
species-level posterior trend estimates from this model,
as a measure of overall agreement between trends
estimated by the two models. The WinBUGS code for
the comparison model is provided in Appendix 2B.

To compare the tendency for each model to pro-
duce extreme trend estimates (i.e., larger in absolute
magnitude), we estimated the slope of a regression of
ML trends on HB trends, using major axis, model II
regression methods (Legendre and Legendre 1998).
Slope values > 1 would indicate that, on average, trends
from the ML model are more extreme for a given
species. Values < | indicate that HB trends are more
extreme. We used an unweighted regression, because
precision-weighted comparisons would greatly reduce
the influence of the extreme trends, which are often
imprecisely estimated.

Precision of trends: To compare the precision of
trend estimates between the two models, we treated
credible intervals for the HB, model and confidence
intervals for the ML model as comparable estimates
of variance (Sauer and Link 2011). Despite their fun-
damental mathematical and philosophical differences,
95% credible intervals from simple Bayesian models
using diffuse priors are generally very similar to 95%
confidence intervals from comparable frequentist mod-
els and, in practice, they are used in similar ways to
assess uncertainty in model estimates (Clark 2005).
We used the model described for comparing the mag-
nitude of trends to simultaneously compare the preci-
sion of trend estimates

1

(Care

osm

This model accounts for both the imprecision of each
trend estimate (through the estimated variance of
trends) and the imprecision around the estimate of the
variance. Specifically, the model assumes that the esti-
mated variances (62,,) of each trend estimate are mutu-

THE CANADIAN FIELD-NATURALIST

Vol. 128

ally independent (both within species [s] and within
models [m]) and independent of their associated trend
estimates (Sauer and Link 2002). The true variance of
each trend (o2,,) was estimated using a chi-squared
distribution with n degrees of freedom, such that
n*6Em m
osm ~n

where v is the number of routes on 1 which the species
was observed (i.e., the number of routes contributing
data to the trend analysis). We then estimated the
proportion of species for which the HB, trend was
more precise (1.€., 6? ;,< 624,,) and the average pair-
wise differences in the standard errors of each species’
trend

( Ui 05 1B~75MLy

5 :

where S is the number of species.

Inter-annual variability in short-term trends: The
two models have very different definitions of trend
(Sauer and Link, 2011): the HB, model estimates trend
as an endpoint comparison of the annual indices in the
first and last years of a given period: the ML model
estimates a series of annual indices, then derives trend
as the slope of a log-linear regression of the indices
against time. Because the HB, model trend estimate
is an end-point comparison, which does not directly
include any of the indices in the intervening years,
trend estimates from the HB, model could be more
variable among consecutive years (i.e., greater inter-
annual variation). For example, consecutive 10-year
trends estimated using the HB, model (e.g., 2000-2010
and 2001-2011) are based on completely different pairs
of annual indices. Variability of this sort is particular-
ly important because these short-term, 10-year trends
are the basic estimates of population change used for
species assessments by COSEWIC. Undue inter-annual
variation in trends could introduce some uncertainty
and instability into important and potentially costly
conservation decisions.

To compare inter-annual variation in short-term
trends, we calculated all possible 10-year trends (Byear
x - year x+10) USING each model from the entire time-
series of BBS data, i.e., B1970-1980, B1971-1981,
B2001-2011. We then calculated the absolute value of the
change in estimates between all 32 consecutive € pairs
of trends, e.g., AB; = |Bi970-1980 — Bio71-1981|, AB2 =
|Bi971-1981-B 1972-1989], > AB32 = |Boo00-2010- Boo01-20111:
We averaged these éstimates of consecutive change
in trend across the full time series

aie ea vi=1..32 AB;
(AB = ‘T39)
and compared the averages between the ML and HB,
models for each species at the national scale.

Scale of annual indices: The annual indices from
the BBS reflect the relative abundance of a species

2014

among regions and influence the relative weight of
each region’s population trend in composite trends for
larger regions. In addition, the indices are used in many
conservation documents as a scale of the relative abun-
dance among species. In regions that are identical in
the two analyses, the two models produce annual in-
dices of abundance on the same scale — the average
expected count on an average route, conducted by an
average observer. However, annual indices from the
two models are derived in fundamentally different
ways. The ML model estimates the annual indices
directly, and trends are derived from log-linear regres-
sions of the indices on year. The HB, model estimates
trends directly and annual indices are derived esti-
mates of departures from a long-term trend line. We
compared the scaling of annual indices from the two
models for a region that is identical in the two analyses
— New Brunswick. For each species, we calculated
the average of the annual indices from each model
for New Brunswick for 1985-1995, which roughly
represents the middle of the time series for the BBS
in Canada. We limited our quantitative comparison to
this one region and window of time because it provid-
ed the most direct comparison. It was largely unaffect-
ed by differences among strata in area weights, the
inclusion or exclusion of particular routes, or in the
indices at either end of the time series, which are more
likely to be influenced by the way in which the ML
model handles years with no observations or by strong
underlying trends estimated using the HB, model.

To quantify differences in the scale of annual in-
dices, we used a generalized linear mixed model with
a Gaussian error distribution, a log-link, a fixed effect
of model type (HB, vs. ML), and a random effect for
species. We used the log-link because the models that
produce the annual indices estimate the variances of
the indices on the log scale and so that we could pro-
vide each species with an approximately equal weight
in the analysis (1.e., comparing estimates on the linear
scale would give abundant species undue weight). We
report the retransformed parameter estimate and con-
fidence intervals for the fixed effect of model type as
the average difference in annual indices between the
two models.

The HB, model used here produces annual indices
ona slightly different scale than the HB, model, which
estimates a single observer—route variance term for
all strata (o2). Therefore, we also compared annual
indices from the HB, model with indices from the
ML model. Observer—route variance influences the
scaling of annual indices in both HB models in two
ways. First, the structure of the observer—route vari-
ance (62 vs. 6,) affects the estimates of the average
stratum-level abundance (5;). Second, the estimates
of observer-route variance are used as a retransfor-
mation factor that scales the annual indices to an esti-
mate of average counts per route.

SMITH ET AL.: NEW BREEDING BIRD SURVEY TRENDS AND INDICES |

tO
n

If a common observer-route variance across all
strata is estimated, as in the HB, model, some of the
ecologically relevant variation in abundance among
strata is modelled as observer—route error and not in-
corporated into the estimates of S,, which, as a result,
become more similar among strata, i.e., they approach
the survey-wide average abundance. The observer
route variance estimate is composed of the variation
in abundance among individual BBS routes as well
as the variation among observers. Although the vari-
ance among observers is entirely a source of error, the
variance among routes represents, at least partly, eco-
logically-relevant spatial variation in abundance. This
spatial variation in abundance among routes is also
nested within the variation in abundance among strata
— routes are uniquely associated with a given strata.
As a result, when a common observer—route variance

>

is estimated (o2 in the HB, model), some of the vari-

ation in abundance among routes is modelled as error.
By contrast, the stratum-specific estimates of observer—

5

route variance (63, in the HB, model) allocate more of
the variation in abundance among strata to the esti-
mates of S..

The stratum-specific variance estimates in the HB,
model also retain greater variation in abundance among
strata, because they provide stratum-specific retrans-
formation factors to rescale the annual indices. The
observer—route variance is one of two variance com-
ponents (0.5 * 6%, in equation 2) that are added to the
parameter estimates on the log-scale to adjust for the
bias in transforming from the log scale to the linear
scale (i.e., count of birds). Annual indices in the HB,
model are scaled by a multiplicative function of e°5*62.,
which varies among strata; the analogous function
(e°5"oZ) is the same for all strata in the HB, model.

All figures that include comparisons of the ML
and HB, models at the BCR scale have been arranged
so that the BCRs are sorted in descending order of an
approximate estimate of data quality, i.e., BCRs are
sorted in descending order of

fits

Ab
for each BCR, b, where 7, is the number of routes and
A, is the area of the BCR. Some additional compar-
isons are included in Appendix 4.

Results
Magnitude of trends

Overall, trends estimated from the ML and HB,
models were generally similar. Estimates from the two
models were correlated for all regions and time periods
(Figure 2), with higher correlations at the level of indi-
vidual strata and BCRs than at the national level (com-
pare plots for Canada and New Brunswick in Figure 2
and Figure 3A). Long-term trends were generally more
correlated than short-term trends (Figure 3A). Trends

126 THE CANADIAN FIELD-NATURALIST Vol. 128
Canada New Brunswick
a &
° gad
§ §
2 <0 a
ie 2
% oO
i=
2 ®
z no}
oD oes Bits
5 E
& ®
oly -_
5 Fi
S ° = °
1 1
wo
.

-15 -10 -5 0 5 10 15 20

HB ; long-term trend estimate (%/year)

HB ; long-term trend estimate (%/year)

FIGURE 2. Point estimates (circles) and associated uncertainty (semi-transparent ellipses) of long-term Breeding Bird Survey
trends for Canada and New Brunswick calculated from the previously used maximum likelihood (ML) model and
the new hierarchical Bayesian (HB,) model. The diameters of the ellipses represent the central 67th percentile of the
posterior distribution of trend estimates from the Bayesian trend comparison model (vertical diameter = ML trend
estimates and horizontal diameter = HB, trend estimates). The diagonal line indicates a 1:1 relationship. The dotted
black lines divide the plot into quadrants representing combinations of trend—signs from the two models (e.g., points
and posterior mass plotted in the upper right quadrant represent species with positive trend estimates from both
models). Differences in trends for Canada incorporate differences between the two models in terms of model struc-
ture and assumptions, as well as differences in area weights, routes, and regions of the country included in the esti-
mates. Differences in trend estimates for New Brunswick are almost entirely due to differences in model structure
and assumptions. One species was omitted from both graphs because its trend estimates were extreme (> 25%/year)

and including it changed the scale such that it became difficult to discern relations among the remaining species.

from the two models also tended to be more correlat-
ed in regions with more data (i.e., BCRs closer to the
left side of Figure 3A, which have more routes in rela-
tion to their area).

At the national level, 54% of species had HB, trends
that were more negative than their ML trends, and
HB, trends were on average slightly more negative:
the average species-level difference in long-term trends
was —0.26%/year, although the 95% credible inter-
val—the interval between the 2.5 and 97.5 percentiles
of the posterior distribution—included zero (—0.68 to
0.16; Figure 3B). In contrast, national short-term trends
from the HB, model were on average more positive:
64% of species had HB, trends that were more posi-
tive, and the average difference was 1.1%/year, (95%
credible interval —0.41 to 1.7; Figure 3B). For most
BCRs, the average differences between trend estimates
from the two models followed the same pattern ob-
served in the national estimates, i.e., the differences in
long-term trends were more often negative and differ-
ences in short-term trends were more often positive
(open circles below the dotted line and closed circles
above in Figure 3B).

All trends from the HB, model tended to be less
extreme (i.€., less in absolute magnitude, points above

the dotted line in Figure 4), except for the national
long-term trends, where the opposite was true.

Precision of trends

Trend estimates from the HB, model, both long- and
short-term, were more precise than estimates from the
ML model in all regions. For national trends, 63% of
species had more precise trends with the HB, model,
and the standard errors of HB, trends averaged approx-
imately 0.5%/year smaller than the standard errors of
the ML trends (Figure 5). Within the BCRs, HB, trends
were more precise for almost all species (73-98%)
and their standard errors averaged approximately 2%/
year smaller (Figure 5).

Inter-annual variability of short-term trends
Ten-year trend estimates from the HB, model were
generally less variable among years than those from
the ML model (Figure 6). Across species, the mean
absolute year-to-year change in 10-year trend estimates
was |.12%/year smaller using the HB, model than the
ML model for a given species. The average species-
level difference (HB,_ ML) in the mean absolute year-
to-year change in national 10-year BBS trends was
=L124SE- O07) = in Figure 6A, most points are to the
right of the 1:1 diagonal line. In general, 10-year trends

2014

SMITH ET AL.: NEW BREEDING BIRD SURVEY TRENDS AND INDICES 127
1.0 A
| © Long-term
} 4 ; © Short-term
505) } ; } ; ;
aS)
©
8 oo 4
-0.5 4
47) B
voral
=
ea
_ ;
i}
E ,
oO 0
a,
S @
: \
ee
(a)
44
tS Sf 1 —=41- SS 1 = 1 eo
Ce a Tee ee ee eee Oe ee ee:
o = | ao | foal | | \ o o o
=
()
z

FiGuRE 3. Correlation and comparison of the magnitude of long- and short-term Canadian Breeding Bird Survey trends esti-

FiGurE 4. Slopes of major axis, mod

mated using the earlier maximum likelihood (ML) model and the new hierarchical Bayesian (HB,) model. Bird
Conservation Regions (BCRs) are arranged in descending order of the ratio of the square root of sample size to area.
The correlations plotted in the upper graph show the overall similarity of trends estimated using the two models. The
differences plotted in the lower graph show the relative bias in the two models: negative values indicate that the HB,
trend estimates were smaller than the ML estimates (i.e., the population change was more negative and/or less posi-
tive); positive values indicate the opposite. All comparisons were made on the posterior estimates of trends, output
from a model that accounts for the relative precision of the original estimates. The circles represent the means of the
posterior distribution, with error bars indicating 95% credible intervals. See Figure | for BCR definitions.

10.0 + 0 Long-term ’
Bo =e Short-term {

a ¢
ee. a ia cals o 4
ee a. Ae oF en
= 1.0“; 9
®
om O18
re)
a |
2 4
0.1 >
a 2 Co te SS io Ny tk Sas, 2
pee oe Oe ee ee
os 2 Oo
§8E 8 2 8 Ba tok ay 8
oO
=
oO
z

el II regressions of Breeding Bird Survey trend estimates from the previously used
ML) model regressed on estimates from the new hierarchical Bayesian (HB,) model. Bird
Conservation Regions (BCRs) are arranged in descending order of the ratio of the square root of sample size to area.
These comparisons do not consider the precision of the estimated trends, for reasons explained in Methods. Values > 1.0
indicate that the absolute values of the ML trends are, on average, more extreme than the HB, trends for each species
(i.e., the ML trends are larger in absolute magnitude) and values < 1.0 indicate that the HB, trends are more extreme.
Error bars indicate 95% confidence intervals for the slope estimates. Slope values are plotted on a logarithmic scale
so that equal deviations from the 1:1 expectation appear equal above and below the dotted line (e.g., a slope of 2.0
de of deviation from the expectation as a slope 0.5). See Figure | for BCR definitions.

maximum likelihood (

represents the same magnitu

THE CANADIAN FIELD-NATURALIST

Proportion of species

ao % 6 © o) oe

HB 3 trends more precise

Mean pairwise differences (HB ;—-ML)
in SE of trend estimates (%/year)
—e
~e

-6 4

© Long-term
@ Short-term

—

Canada
BCR-13
BCR-9
BCR-14

New Brunswick

BCR-10
BCR-4
BCR-6
BCR-8

BCR-5
BCR-12
BCR-11

Vol.

128

FIGURE 5. Comparisons of precision of Breeding Bird Survey trend estimates between the previously used maximum likeli-
hood (ML) model and the new hierarchical Bayesian model (HB,) in various regions of Canada. Bird Conservation
Regions (BCRs) are arranged in descending order of the ratio of the square root of sample size to area. The upper
graph shows the proportion of species in each region for which the HB, trend estimates were more precise than the
ML trend estimates. The lower graph shows the mean pairwise differences in trend estimates’ standard errors (SE);
negative values indicate the HB, estimates were more precise, i.e., have smaller SEs. Circles and associated error
bars represent the means and 95% credible intervals of the posterior distribution of estimates from a trend compari-
son analysis that accounts for the uncertainty in both the estimates of trend and the estimates of the variance of the

trends. See Figure | for BCR definitions.

from the ML model were much more variable for
species with relatively imprecise annual index estimates
(Figure 6B, lower row). For some species, 10-year
trends from the HB, model were more variable than
those from the ML model (points to the left of the diag-
onal line in Figure 6A); these are species with relative-
ly precise estimates of highly variable annual indices
(Figure 6B, upper row).

Scale of annual indices

On average, annual indices from the HB, model
were somewhat larger than those from the ML model
(mean difference = 1.29, 95% credible interval 1.23—
1.36), however the log-log relationship closely follows
the 1:1 line (Figure 7). Mean estimates from the HB,
model were more similar to the annual indices from
the ML model than those generated by the HB, model
with a common observer-route effect (mean difference
2.51, 95% credible interval 2.32-2.71). In addition, the
differences between the HB, and ML model depended
on the species’ abundance; almost all species with ML
indices > 2 were above the 1:1 line and the slope of the

Pine Siskin o g
e
American Coot
e

°

Mountain Chickadee
e °

/

o 4

Greater Yellowlegs
.

fo)

Do e
Northefh Pygmy-Owl
[e}

Mean absolute year to year change HB , ten-year trends

1}
Brewer's Sparrow e

T T
6 8

Mean absolute year to year change ML ten-year trends

FIGURE 6A.

2014 SMITH ET 4L.: NEW BREEDING BIRD SURVEY TRENDS AND INDICES 129

American Coot Pine Siskin

1:5: 92:0: 22.6

1.0

0.0 0.5

1970 1980 1990 2000 2010 1970 1980 1990 2000 2010 1970 1980 1990 2000 2010

Greater Yellowlegs > | Northern Pygmy—Owl Brewer’s Sparrow

Annual index (HB 3 or ML)

1970 1980 1990 2000 2010 1970 1980 1990 2000 2010 1970 1980 1990 2000 2010

FiGuRE 6. Comparison of the year-to-year variability in 10-year Breeding Bird Survey trends for Canada estimated using the
maximum likelihood (ML) model and the new hierarchical Bayesian (HB,) model. A shows the species-level esti-
mates of the absolute year-to-year change in trend values for the two models, from a series of consecutive 10-year
frends SOM 2,6 paonne Megiiosi-<->toooraont Loe illed circles indicate the species trends plotted in B. B shows over-
lays of annual indices from the HB, model (solid lines) and the ML model (dashed lines). American Coot (Fulica
americana), Pine Siskin (Spinus pinus) and Mountain Chickadee (Poecile gambeli) are examples of species for which
the HB, trends are more variable, i.e., points above the diagonal in A. Greater Yellowlegs (Tringa melanoleuca),
Northern Pygmy-Owl (Glaucidium gnoma), and Brewer’s Sparrow (Spizella breweri) are species for which the ML
trends are more variable, i.e., points below the diagonal in A.

HB 3 (2011 CWS model) HB > (2011 USGS model)
7: 7
S S | -- log-log sl te
3 8 2 aisles og log slope ary
gs 8 R -
a
o
o w wo 4
D
Sic N
g
S wo | o
m oO foo) *
a 7 |
fo) = ies e te |
4 oO o | 7%’* F
= N N ol
oO " 4 |
3 = l eae T ips

0.02 0.1 OS 2.5 20; 100 0.02 «0.1 O5 2 6 20 100

ML average annual index

FIGURE 7. Log-scale comparisons of the average of Breeding Bird Survey annual indices for New Brunswick (for
1985-1995) from the previously used maximum likelihood (ML) model, plotted against annual indices from two
hierarchical Bayesian models: the new Canadian model (HB,) and the HB, model described in Sauer and Link
(2011). Each dot represents one species, and the black dashed lines are predictions from linear models regressing the

ual indices from the two HB models on the log of the average annual indices from the ML

f the average ann Seale ;
eRe : line represents a 1:1 relationship. Note: CWS = Canadian Wildlife Service; USGS = United

model. The solid grey
States Geological Survey

130

log-log relationship diverged from the 1:1 line (Fig-
une 7).

Discussion

Canadian BBS trends estimated using the HB, model
are generally similar in magnitude to those estimated
from the ML model that was in use between 2002 and
2009. This suggests that for most species, our under-
standing of population status has not been drastically
changed by the adoption of the HB, model.

Overall, the trends from the new HB, model are
more precise, less likely to have extreme values, and
less variable among years than those from the previ-
ous ML model. A similar hierarchical model (HB,)
showed similar improvements compared with the route-
regression model previously used to estimate BBS
trends in the United States (Sauer and Link 2011).
Although no single model is ideal for all species in
all situations, the differences demonstrated here sug-
gest that the HB, model provides better information
on the population status of approximately 300 bird
species breeding in Canada. The broad similarities in
trend estimates from the two models are not surpris-
ing; the BBS represents a large dataset, and many
temporal trends are clear and strong regardless of the
analytical models used to estimate them (as noted in
similar cross-model comparisons in Thomas and Mar-
tin 1996 and Sauer and Link 2011). For some species,
the two models generate strikingly different trend esti-

THE CANADIAN FIELD-NATURALIST

Vol. 128

mates, but, in most cases, the estimates are extremely
imprecise for both models. For example, in Figure 2,
the points that fall far from the 1:1 line are generally
very imprecise (1.e., have large credibility ellipses),
and the credibility ellipses overlap the 1:1 line in at
least one dimension for most of these species.

Magnitude

In some BCRs, long-term trends derived from the
HB, model are generally more negative/less positive
than those from the ML model. This difference is par-
tially the result of a statistical artefact in the ML mod-
el; thus, the HB, trend estimates are less biased. In the
ML model, for years in which a species was not ob-
served in a particular region, the annual index is set to
a small arbitrary value. That is, the annual index itself
is estimated at zero, but when the trend is calculated, a
small constant is added (0.001) so that the log-linear
regression can be calculated. For species and regions
with these zero counts near either end of a time series,
any estimated long-term trend will be strongly influ-
enced by this arbitrary value (O’Hara and Kotze 2010).
Because of the smaller number of BBS routes in ear-
lier years, species are more likely to have annual index
estimates of zero for those years, and, therefore, their
abundance is more likely to be represented by the arbi-
trary value leading to positive bias in the ML model
(Figure 8). In contrast, in the HB, model, the annual
index for those years is estimated to be a non-zero val-

st
~
fo)
NA
=
io)
(S}
i
< ©
=
2 6
=I
= (oe)
=£ o-°
= oO
o
a8
te | fo) ©
is)
5
2 3 ;
S 3 = fe) . © fe)
fe)
fe) fe) (oxexe) 25
o fe) fo) fo)
SS °
= 7 fe) fe)
fo) [exe) fe) fe)
9 G6 fe) fexe) fo)
s (oxo) Gio “6
eo 4 i
ro)
T T TI T all
1970 1980 1990 2000 2010

FIGURE 8. Proportion of species with annual index estimates equal to zero when calculated with the maximum likelihood

model, previously used to estimate Breeding Bird Survey trends in C
izontal lines indicate proportions of species with at least one annu
(1970s, 1980s, 1990s, and 2000s).

anada. Circles indicate annual proportions: hor-
al index equal to zero in each of four decades

2014 SMITH ET AL.: NEW BREEDING BIRD SURVEY TRENDS AND INDICES 131]

ue that reflects the best estimate of the abundance in
that year, given the number of routes with zero counts
in that year, and the estimates of all other parameters
in the model.

Precision

Canadian BBS trends estimated using the HB, mod-
el were more precise than those from the ML model,
and this difference was more pronounced for smaller
regions (i.e., New Brunswick and the BCRs). At the
national level, the HB, model’s hierarchical structure
makes efficient use of the data and gives moderately
more precise estimates than the ML model. For smaller
regions, the hierarchical structure of the model results
in additional efficiencies. Because the stratum-level
parameters are estimated within a single national mod-
el, estimates for the error, over-dispersion, first-year
observer effects, and the observer—route variance (1.e.,
the nuisance parameters) share information among
strata. This sharing of information makes for more pre-
cise stratum-level estimates of the nuisance parameters
than would be possible if they were estimated solely
from the much smaller set of routes within each stra-
tum. In contrast, the ML model accounts for the observ-
er and route-level nuisance parameters separately with-
in each region.

The HB, model’s increased precision has important
benefits for conservation in that it decreases uncer-
tainty around the assessment of species’ status. Steep
rates of change or abrupt changes in populations (e.g.,
30% decreases in populations that may warrant species
at risk status; COSEWIC 2011*) can be identified with
greater certainty and over shorter periods of time. Sim-
ilarly, species can be more confidently classified into
status categories, such as those used in Environment
Canada’s Status of Birds in Canada website (www.ec
.gc.ca/soc-sbc).

Variability among years

For most species, the HB, short-term trend estimates
are less variable among years than the ML short-term
trend estimates, because the hierarchical structure of
the year-effect parameters makes annual fluctuations
much less sensitive to sampling error and annual route
coverage. For a few species, short-term HB, trend esti-
mates are more variable among years than ML trends,
but only when there is relatively strong evidence sup-
porting large annual fluctuations in the population’s
status. In essence, although short-term HB, trend esti-
mates will fluctuate more for populations whose status
is well estimated, those fluctuations are likely appropri-
ate because they are more likely to reflect real changes
in the populations and not sampling error. In these cas-
es, management decisions that rely on short-term trend
estimates, such as COSEWIC status assessments,
should also examine the population’s recent trajectory
to give some context to the trend estimate in any given
year. For data-poor populations, the HB, model’s short-
term trends will be more stable across years, So man-

agement decisions can be made with some confidence
that a species’ status assessment will not strongly
depend on the year in which it was assessed.

Annual indices

At the stratum level, annual indices from the HB,
model are very similar to those from the ML model.
They are directly interpretable as the expected count
by an average observer on an average route in the stra-
tum. They are also more similar in scale to the indices
from the ML model than annual indices from the HB,
model. Estimating observer-route variance at the stra-
tum level in the HB, model brings the scaling of the
annual indices closer to a scale familiar to users of the
ML model results, which is approximately equal to the
observed average count across the routes run in any
given year.

The improvement in scaling of the annual indices in
the HB, model over the HB, model also has an influ-
ence on regional trends, because it better reflects the
variation in abundance among strata. Because region-
al annual indices are sums of the stratum-level indices,
the relative contribution of each stratum’s trend to a
regional trend is partly mediated by the relative abun-
dance of birds in each stratum. In relatively rare but
potentially important cases, regional trends from the
HB, and HB, models can be strikingly different: where
the average abundance, trend, and observer—route vari-
ance are highly variable among strata. For example,
the HB, model applied to Canadian data for Wood
Thrush (Hylocichla mustelina) estimates the national
trend at —1.8 (95% credible interval —2.6 to —0.9),
while the HB, model estimates the national trend at
4.4 (95% credible interval —5.3 to —3.6). This large
difference — the credible intervals do not overlap —
is due to the relative weight in the two analyses of the
largely stable Wood Thrush population in southern On-
tario (Ontario BCR 13). In this region, the observer—
route variance is much lower than in any other stratum
in the analysis. As a result, the observer—route variance
retransformation factors are very different in the two
models (62 >> O2rcp);). The larger retransformation
factor in the HB, model creates annual indices that
overestimate the observed abundance of Wood Thrush
in southern Ontario by a factor of 3 and, similarly,
overestimate the proportion of the national population
that occurs in the region and, therefore, its influence
on the national trend.

Users of published BBS estimates should be aware
that the scale of annual indices from the HB, model
and the ML model may differ because the models ac-
count differently for routes in regions not included in
each species analysis. The HB, model estimates an
annual index scaled to the expected count averaged
across routes within the strata included in the analysis,
i.e., it excludes routes where the species data are too
sparse or that are outside the species’ breeding range.
The ML model’s annual indices are scaled to the ex-
pected count averaged across all routes within the re-

132

gion being analyzed, e.g., for national trends, all routes
run in the country are included, regardless of whether
they fall within the species’ breeding range. For broad-
ly distributed and common species, for which all strata
and all routes are included in the HB, analysis, the
scales of the national estimates from each model will
be equal. For more locally distributed species, for which
the HB, analysis only includes a subset of the strata
(e.g., any species that breeds exclusively west of the
Rockies), the HB, annual indices will be scaled to a
much higher abundance because they ignore all of the
routes and strata outside of the species’ breeding range,
where the average counts are zero. It is important to
note that this difference does not affect our quantita-
tive comparisons of annual indices among the HB,,
ML, and HB, models (i.e., Figure 7), because we com-
pared them within a single stratum, nor does it affect
trend estimates in the two models; it only affects the
scale of the annual indices.

Improved population inference from BBS

Beyond the quantitative comparisons, some philo-
sophical arguments suggest that the HB, model rep-
resents an improvement over the ML model. First, the
Bayesian framework provides much more intuitive
and flexible inference than the frequentist framework
of the ML model. The correct inferential interpretation
of Bayesian credible intervals is almost certainly a more
natural expression of the type of inference that users
of BBS trends desire, i.e., there is a 95% probability
that the trend is greater than the lower bound and less
than the upper bound. In contrast, the correct interpre-
tation from frequentist analyses (usually confined to
tests of significance) do not relate to the estimates of
population change, but to the data that were sampled,
e.g., if the true rate of population change is zero, there
is less than a 5% chance of observing the data that
were collected. In addition, the ability to estimate the
full posterior distribution of a broad suite of derived
parameters, such as the probability that the population
has declined more than 50% in 10 years (i.e., one of
the COSEWIC criteria for “Endangered” status), pro-
vides a practical and flexible approach to assessing the
uncertainty around particular conclusions one might
draw from a BBS trend.

Second, the HB framework and the HB, model in
particular provide an efficient, flexible, coherent and
complete framework for the analysis, which the ML
model lacks. The hierarchical structure of the model
makes efficient use of the data and is less sensitive to
annual variation in sampling error. In addition, the ibs
model can be expanded easily to include covariables
and explicit spatial structure (e.g., Thogmartin ef al.
2004; Nielson et a/. 2008), and the HB framework
lends itself well to composite and comparative analy-
ses of the BBS and other surveys (e.g., Link and Sauer
2007 and the method used here to compare the mag-
nitude and precision of trends from the two models).
Finally, the HB, model estimates trends and annual in-

THE CANADIAN FIELD-NATURALIST

Vol. 128

dices within a single coherent model structure which
the ML model lacks. For example, the ML model re-
quires the addition of an arbitrarily chosen constant
to some years’ annual indices to estimate trends.

A third example of improved inference from results
of the HB, model is that the trends and estimates of
uncertainty around trends for multi-strata regions
(e.g., national trends), are a much clearer reflection of
the full uncertainty of the population status estimates
within the region. That is, regional and national esti-
mates are combinations of stratum-level estimates,
appropriately weighted by the proportion of the spe-
cies’ breeding population in each stratum (by summing
indices of relative abundance, weighted by the stra-
tum’s area). In addition, the strata in the HB, model are
more likely to reflect spatial variation in population
trends and trajectories than the degree-block strata in
the ML analysis. The HB, strata are structured on
BCRs, which are relevant to population processes, and
political units, which are relevant to management and
human activity. In contrast, inferences regarding trends
and uncertainty from the ML model only apply to the
degree blocks included in the analysis, i.e., degree-
blocks with data. Furthermore, the degree-block
weighting of the ML model means that the relative
influence of populations with potentially disparate
trends and trajectories among different regions (i.e.,
BCRs and political units) depends on the relative sam-
pling intensity of the BBS in those regions (i.e., the
number of degree-blocks with data). For example, the
White-throated Sparrow (Zonotrichia albicollis) occurs
on BBS routes in the southern BCRs (12, 13, and 14),
where there are many degree-blocks with data and
where trends are strongly negative. However, the bulk
of its population occurs in the more northern BCRs
(6, 7, and 8), where trends are relatively stable and
there are few degree-blocks with data. The degree-
block weighting of the ML analysis puts relatively
more weight on the decreasing southern populations
(national short-term ML trend —0.9%/year [95% con-
fidence interval —1.7% to —0.1%]), whereas the stra-
tum area weights of the HB, analysis put more weight
on the more positive and much less precisely estimat-
ed trends from the northern populations (national short-
term HB, trend 0.59%/year [95% credible interval
—1.64% to 3.87%]). The estimated average trend from
the HB, model better reflects the status of the bulk of
the Canadian population of the White-throated Spar-
row in the northern BCRs and. given that the north-
ern BCRs are relatively poorly surveyed by the BBS,
the wider credible interval of the national HB, trend is
a more appropriate measure of the uncertainty around
the population’s status. This more appropriate weight-
ing of imprecisely measured trends from regions that
are relatively large but poorly surveyed also explains
the modest improvements in precision at the national
level, relative to the greater improvements within indi-
vidual BCRs (Figure 3),

2014

Because of the flexibility of the HB, model frame-
work, the best short-term trend and the best long-term
trend do not necessarily include data from the same
spatial areas. Since 1966, spatial coverage by Canadi-
an BBS routes has expanded gradually into the more
northerly strata. The earliest BBS routes in Canada
were run in the Maritimes and Quebec in 1966. By the
early 1970s, much of southern Canada had adequate
coverage. However, some more northerly regions have
only had adequate coverage for the last 20 years. Be-
cause of the paucity of data from the Northwest Ter-
ritories and much of the Yukon before the late 1980s
or later, trends in these areas cannot reasonably be
extrapolated back to 1970. Long-term trend estimates
for large regions (e.g., national trends) can be estimat-
ed, but they must exclude some strata, for which we
have only recent data. In contrast, short-term estimates
for those regions should include all of the strata with
data. In the HB, model, deriving separate short- and
long-term trend estimates that include different strata
is a straightforward process of summarizing stratum-
level annual estimates, and it retains the entire poste-
rior distribution of all derived estimates. To get simi-
lar separate trends from the ML model would require
multiple runs using different subsets of the data, and
each subset would have a reduced sample size and,
therefore, reduced precision.

Limitations of the HB, model and future evolution
Along with the benefits of a complex hierarchical
model structure come complex consequences for pop-
ulation inference. For example, the trend term in the
HB, model combined with the distributional assump-
tions of the random year-effects terms means that, for
years with relatively sparse or highly variable data, an-
nual indices will tend to track a fitted estimate of the
species’ long-term trend. These assumptions and mod-
el structure are reasonable, given that the long-term
trend is estimated from the data. However, in some
cases, they could lead to an over-smoothing of the an-
nual indices. For example, the model does not account
for autocorrelation in the sequence of successive year-
effects and could, therefore, be insensitive to cyclical
population patterns, unless each individual year con-
tained relatively strong evidence of a departure from
the long-term trend. Similarly, the number of BBS
routes in Canada has more than doubled since 1980,
and, therefore, sparser data in the early years means
that those early annual indices are less likely to depart
from the long-term trend than annual indices in later
years. Although these assumptions greatly reduce the
influence of sampling errors on trend estimates (Link
and Sauer 2002), as evidenced by the reduced inter-
annual variability of HB, trend estimates (Figure 6),
there are species for which the smoothing of annual in-
dices could lead to bias in both the individual indices
and the trends. Future refinements of the model may
‘nclude terms to model the temporal autocorrelation
in the year-effects, so that the model will more closely

SMITH ET AL.: NEW BREEDING BIRD SURVEY TRENDS AND INDICES

133

reflect population fluctuations when successive years
show similar departures from the long-term trend.

Conclusion

Overall, the use of the HB, model to estimate trends
and annual indices of abundance provides improved
information on the population status of birds in Cana-
da. Estimates from the HB, model are generally sim-
ilar in direction and magnitude to estimates from the
previous model, but they offer greater precision, less
variation among years, a better representation of the
spatial variation in population status across Canada,
and a more intuitive and flexible assessment of uncer-
tainty. As statistical science evolves, the analysis of
BBS data will follow suit, continuing to improve as it
has in the past. This evolution and improvement hon-
ours the contributions of the thousands of volunteers
who have participated in the BBS since its inception,
and ensures that this survey will remain at the forefront
of our understanding of the status of Canada’s and
North America’s birds.

Acknowledgements

We thank the thousands of skilled volunteers who
have contributed to the Breeding Bird Survey over the
years, as well as those who have served as provincial
and territorial coordinators. Our thanks also go to Pete
Blancher, Brian Collins, Bill Link, David A. W. Miller,
John Sauer, and Joe Veech for their insightful com-
ments and review of the paper, model development
or both.

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Supplementary material available at:
http://www.canadianfieldnaturalist.ca

Received 23 September 2013
Accepted 22 November 2013

APPENDIX 1. Detailed description of the maximum likelihood (ML) model used to generate Canadian Breeding Bird Survey.

trends before 2010.

APPENDIX 2. WinBUGS (Bayesian inference Using Gibbs Sampling) language descriptions of the hierarchical Bayesian models.

APPENDIX 3. Microsoft Excel workbook showing differences in trend estimates between the maximum likelihood (ML) and
the hierarchical Bayesian (HB,) models, with separate worksheets for short- and long-term trends and for each Bird

Conservation Region, Canada, and New Brunswick.

APPENDIX 4. Additional comparisons of the hierarchical Bayesian (HB,) and maximum likelihood (ML) models, including
comparisons of the number of species with trend estimates and the classification of species trends into the categori-
cal assessments of population status used in Environment Canada’s Status of Birds in Canada website (Environment

Canada 2011).

Acoustic Monitoring of Migratory Birds over Western Lake Erie:
Avian Responses to Barriers and the Importance of Islands

CLAIRE E. SANDERS! and DANIEL J. MENNILL!

Department of Biological Sciences, University of Windsor, Windsor Ontario NOB 3P4 Canada
Corresponding authors: sandersc@uwindsor.ca and dmennill(@uwindsor.ca

Sanders, C laire E., and Daniel J. Mennill. 2014. Acoustic monitoring of migratory birds over western Lake Erie: avian responses
to barriers and the importance of islands. Canadian Field-Naturalist 128(2); 135-144.

Understanding the flight patterns of migrating birds is critical for informing conservation actions and management decisions.
We studied the geographic and temporal distribution of birds migrating through the southern Great Lakes using nocturnal
acoustic monitoring data and banding records from sites on Pelee Island in Lake Erie and on the mainland along the north
shore of Lake Erie. Given that Lake Erie may represent an ecological barrier to migratory birds, we predicted that mainland
and island sites would show different patterns in both the number of passage migrants and the timing of their migration.
Analysis of over 60 000 flight calls from 6200 h of recordings revealed significantly more migrants over the island than the
mainland in both spring and fall 2012. The acoustic data provide evidence that none of the species or species groups examined
avoided crossing the lake. Birds were detected significantly earlier on Pelee Island than on the north shore of Lake Erie in
spring, although they were not detected earlier on the mainland in fall. These results suggest that Lake Erie is not a major
barrier to migration. The large number of birds detected over the island suggest that birds may concentrate their flight over
islands in the middle of the lake, although recordings of migrants over open water will be required to support this suggestion.
Our results show that Pelee Island is an important part of the migratory route of North American birds and provide valuable
information on the movement of nocturnal migrants over the Great Lakes.

Key Words: acoustic monitoring; banding; bird migration; ecological barriers; geographic barriers; night-flight calls; Lake

Erie; Essex County; Ontario; Pelee Island

Introduction

Migratory birds can fly non-stop for thousands of
kilometres (Egevang ef al. 2010; Bairlein ef al. 2012;
Battley et al. 2012) and are likely to encounter ecolog-
ical barriers, such as bodies of water, mountain ranges,
or stretches of desert as they move between breeding
and wintering grounds. Numerous factors might lead
birds to select circuitous routes rather than cross over
ecological barriers (Alerstam 2001), including their
own body condition and external information such as
weather conditions (Schmaljohann and Naef-Daenzer
2011). The increased risk of crossing an ecological bar-
rier may be offset by the time or energy saved in using
the shortest migratory route (Bruderer 2001). This may
be particularly important during spring migration, when
urgency drives males to reach their breeding grounds to
compete for prime territory and attract mates (Francis
and Cooke 1986).

Migration strategies and the behaviour of migrating
birds are well-studied areas of ornithology (Moore and
Kerlinger 1987; Gauthreaux and Belser 1999). During
a journey from Africa to Europe, for example, migratory
birds encounter the Sahara Desert, the Mediterranean
Sea, and the Alps, all of which could pose a threat to the
long-distance migrant. Historically, researchers believed
that birds were distributed homogenously as they nav-
igated the Mediterranean (Bruderer 2001); however,
banding studies have demonstrated that birds exhibit
diverse migration strategies when encountering ecolog-
ical barriers (Spina and Pilastro 1999). In the Mediter-

ranean, radar studies have shown that the number of
migrants over water is often a third to half that over the
Iberian Peninsula, demonstrating that most birds prefer
to circumnavigate the sea (Bruderer and Liechti 1999).
Furthermore, islands and peninsulas appear to play an
important role for migrants that choose to cross the
Mediterranean Sea, although the use of island stopovers
varies among species and within species based on age
and sex (Barriocanal and Robson 2007).

The Gulf of Mexico is a major ecological barrier in
North America. Birds vary in their propensity to cross
the water of this gulf (Stutchbury ef a/. 2009). For ex-
ample, geolocator data from a Wood Thrush (Hylocicha
mustelina) suggest that it crossed the Gulf of Mexico
during its northward migration, but went around on its
southward journey; another thrush chose a land-based
route on its northward spring migration but then crossed
the gulf in the two subsequent spring migrations (Stan-
ley et al. 2012). Barrier islands along the Mississippi
coast play an important role for some migrants, provid-
ing habitat for foraging before or after the substantial
journey across the gulf (Moore er al. 1990).

North America’s Great Lakes are ecological obsta-
cles that influence the migration of songbirds (Diehl ef
al. 2003; Deutschlander and Muheim 2009). The geog-
raphy of the lakes channels millions of spring and fall
migrants through the region. The north and south shores
of Lake Erie, as well as the Lake Erie island archipel-
ago, are important stopover locations, and many parts
of this region have been designated Important Bird

S16)

Areas by Bird Studies Canada and Nature Canada be-
cause of their seasonal concentration of migratory birds
(BirdLife International n.d.*). However, because of the
technical challenges involved in monitoring nocturnal
migration of many bird species, we lack significant
information on the ecology and behaviour of migratory
birds that pass through the Great Lakes. Recent radar
studies in the region have explored ecological barrier
avoidance (Diehl et a/. 2003), as well as habitat use
by migrant birds and stopover ecology (Bonter ef al.
2009). Radar data confirm that many migrants cross the
Great Lakes in spring and fall, although higher densi-
ties of birds were always detected over land than over
water during migration (Diehl e¢ a/. 2003). Little is
known about the importance of the islands in western
Lake Erie to migratory bird species.

Although general patterns of barrier avoidance can
be revealed through radar studies of migratory birds,
it is not possible to distinguish between bird species
using radar. In addition, if nocturnal migrants travel at
lower altitudes than diurnal migrants, as reported in the
Gulf of Mexico, they may not be detected by radar
(Farnsworth and Russell 2005). Acoustic monitoring
technology overcomes these limitations. Using species-
specific signatures of the flight calls of migratory birds,
we can collect detailed population and behavioural in-
formation on cryptic, aquatic, and nocturnal species
that are otherwise difficult to monitor (Marques et al.
2013). We recently showed that night-flight call data
can be used to determine the timing and magnitude of
spring and fall migration through the southern Great
Lakes (Sanders and Mennill in press). Therefore, night-
flight call data may improve our understanding of the
spatial and temporal distribution of avian migration pat-
terns in the western basin of Lake Erie.

In this investigation, we used night-flight call data
and banding station records to determine whether Lake
Erie acts as an ecological barrier to migratory birds by
comparing the number of birds detected on an island
mid-way across the lake versus the number detected
on the north shore. If Lake Erie acts as an ecological
barrier, we expected to find a smaller community of
migrants over the island than on the mainland. If Lake
Erie does not serve as an ecological barrier, but instead
concentrates birds over the island chain as they cross
the lake, then we expected to detect more migratory
birds over the island than at mainland sites. Finally, if
Lake Erie does not serve as a barrier, but an obstacle
that temporarily delays the migration of birds, we ex-
pected to see a lag in detection between the southerly
island and the northerly mainland sites in spring migra-
tion, and the reverse pattern in the fall.

Study Area

We studied bird migration in Essex County, Ontario,
during the 2012 spring migration from April 15 to June
15 and the 2012 fall migration from August 15 to No-
vember 10. We recorded night-flight calls at seven loca-

THE CANADIAN FIELD-NATURALIST

Vol. 128

tions (Figure |). Three recording sites were located on
Pelee Island mid-way across Lake Erie: two at a bird
banding station in Fish Point Provincial Nature Reserve
at the southern end of the island (41°44'N, 82°40'W)
and one 15 km north at Lighthouse Point Provincial Na-
ture Reserve at the northern end of the island (41°47'N,
82°38'W). Four recording sites were located on the
north shore of Lake Erie: Holiday Beach Migration
Observatory in the Holiday Beach Conservation Area
(42°02'N, 83°02'W); a private woodlot near McGregor,
Ontario (42°06'N, 82°59'W); Cedar Creek Conserva-
tion Area (41°00'N, 82°47'W); and Point Pelee Nation-
al Park (41°56'N, 82°30'W).

Methods
Acoustic recordings

At each of the seven recording sites, we deployed an
autonomous digital recorder (SM-2 Song Meter, Wild-
life Acoustics, Concord, Massachusetts) equipped with
a single night-flight call microphone (SMX-NFC, Wild-
life Acoustics), mounted on a 30 x 30 cm plastic plate
to minimize recording of sounds below the microphone.
We attached the microphone to a 5.8-m pole that we
lashed to a tree or post at the seven recording sites. To
minimize interference noise from leaves, insects, and
amphibians, we positioned the microphones just above
tree height. With seven identical microphones all mount-
ed in similar fashion above the trees, we assumed that
each of our seven recorders was equally capable of de-
tecting the night-flight calls of birds passing overhead.

As recommended in eBird’s Nocturnal Flight Call
Count Protocol (eBird n.d.*), we programmed recorders
to begin sampling at astronomical dusk and conclude
at astronomical dawn (approximately 70 minutes after
sunset to 70 minutes before sunrise), when the sun was
more than 18° below the horizon. Recordings were
collected at 44 100-Hz sampling frequency with 16-bit
accuracy in WAVE format in 1-h and 59-minute files
(1-minute silent intervals between files allowed the
recorders to write the recordings to memory cards).
Each recorder was visited every 3—5 days to change
batteries and collect recordings.

Recordings were processed manually to ensure the
greatest possible accuracy in detecting all flight calls,
as recommended by Swiston and Mennill (2009). We
processed recordings in two stages (for detailed descrip-
tion see Sanders and Mennill in press). Briefly, record-
ings were first visualized as sound spectrograms using
Syrinx-PC sound analysis software (J. Burt, Seattle,
Washington) and scanned 30 s at a time, by a team of
12 volunteer sound analysts (spectral settings: 1024
FFT size, Blackman window). We used the time and
frequency cursors in Syrinx-PC to annotate all night-
flight calls in our recordings.

During the second stage of processing, we classified
the species or species group of as many night flight
calls as possible. Using both time and fr equency char-
acteristics of the spectrograms of various flight calls

2014 SANDE ; : rT OF
ERS AND MENNILL: MONITORING OF MIGRATORY BIRDS OVER WESTERN LAKE ERIE 137

Mainland
recorder 2

Mainland

Lake
Erie

Canada

2—>

Ontario

Mainland
recorder 1 haceidanel s aealdiler ttt
Mainland banding site
bandin (spring)
site (fall

Mainland
recorder 4

Island
recorder 3
Island
recorders Island banding site

1land2

<4
i

(spring and fall)

FIGURE 1. Map of study area showing the seven acoustic recording locations (white circles) and the three banding stations
(black stars) used to study spring and fall migration in 2012 around the western basin of Lake Erie, Essex County,

Ontario.

to distinguish between species (Sanders and Mennill
in press, Appendix 1), we constructed a classification
chart modified from Evans and Rosenberg (2000). After
evaluating our own recorded flight calls and reference
recordings (Evans and O’Brien 2002*), we determined
that 67 species of nocturnal migrants could be identi-
fied to the species level (i.e., their night-flight calls
were distinctive) or into seven bioacoustic categories,
or species groups, each comprising multiple bird spe-
cies whose calls could not be distinguished from one
another (details in Sanders and Mennill in press, Ap-
pendix |). For example, the night-flight calls of North-
ern Parula (Setophaga americana) and Pine Warbler
(S. pinus) both appear as a single down-sweep on a

sound spectrogram; thus, we pooled the detections for
these species into the “single down-sweep” species-
group category (Sanders and Mennill in press, Appen-
dix 1). Furthermore, several species spanned multiple
bioacoustic categories or were classified as distinct at
the species level as well as a member of a broader cat-
egory. For example, Ovenbirds (Seiurus aurocapilla)
produce a distinctive checkmark-shaped flight call that
is species-specific, but also produce calls that consist
only of a frequency modulated upsweep (i.e. the “up”
complex; Sanders and Mennill in press).

Banding data

Banding data were collected at two migration mon-
itoring stations that operated mist nets concurrent with

138 THE CANADIAN FIELD-NATURALIST

our recordings. During spring 2012, birds were banded
at Fish Point Provincial Nature Reserve on Pelee Island
and at Hillman Marsh Conservation Area on the north
shore of Lake Erie from April 15 to June 10. During
fall 2012, birds were banded at Fish Point Provincial
Nature Reserve on Pelee Island and at Holiday Beach
Conservation Area on the north shore of Lake Erie
from August 15 to November 10.

The type of species captured in mist nets is heavily
influenced by local habitat; the nets at both Pelee Island
and Holiday Beach banding stations are located in sim-
ilar habitats of semi-mature deciduous forest dominat-
ed by Silver Maple (Acer saccharinum) and Red Maple
(Acer rubrum), with Eastern Cottonwood (Populus
deltoides), Black Ash (Fraxinus nigra), American
Elm (U/mus americana), Common Hackberry (Celtis
occidentalis), and Pin Oak (Quercus palustris). The
Hillman Marsh banding station operates in an early suc-
cessional habitat previously dominated by hawthorn
(Crataegus sp.), Rough-leaved Dogwood (Cornus
drummondii), Eastern White Pine (Pinus strobus), Black
Cherry (Prunus serotina), Northern Red Oak (Quercus
rubra), willow shrubs (Salix sp.), and some Pin Oak
(Quercus palustris). The banding station on Pelee Is-
land used 10 mist nets and followed the monitoring
protocol recommendations of Hussell and Ralph (2005).
The nets at this station were open half an hour before
sunrise and ran for 6 h, excluding down time when nets
were closed due to bad weather (rain or wind) or high
bird volume when nets were closed to ensure safe and
efficient bird handling. The Holiday Beach Migration
Observatory operated both the banding stations on the
mainland at Hillman Marsh and Holiday Beach. As a
volunteer organization, this observatory’s research ef-
forts varied with the availability of volunteers; seven
nets were in use at Hillman Marsh in spring 2012 and
16 at Holiday Beach in fall.

To facilitate comparisons with our acoustic data, we
pooled banding records for the species that made up
each of our acoustic species groups (e.g. given their
similar night-flight calls, we pooled together all Fox
Sparrows and Song Sparrows captured in mist nets in
a “FOSP/SOSP” species group of bird captures).

Statistical analysis and sample size

To determine whether there were more detections
over the island or the mainland, we combined the detec-
tions from all mainland recorders, adjusted for the num-
ber of hours per recorder, and calculated the average
number of calls per recording station. We repeated this
process for the island recorders to produce a relative
number of calls per recording station for each species
or species group (Table 1). We used sign tests to deter-
mine whether the number of acoustic detections over
the island was less than the number over the mainland
across 38 species and species groups. Given the well-
established problems of estimating abundance from
mist net records (Karr 1981) — especially due to the
strong influence of surrounding habitat — we did not
compare the total number of birds in mainland and
island mist nets.

TABLE |. Number of classified acoustic detections (per 1000 h of recording) and mist net detections (per 1000 h of netting) of migratory birds in spring and fall 2012 at mainland and

island locations in Essex County, Ontario.

Fall migration

Spring migration

Net detections

Mainland

Acoustic detections

Mainland

Net detections

Mainland

Acoustic detections

Mainland

Bioacoustic

Island Island Island
1236.09

Island
228.44.

Species or species group (7 = 38)

category

64.64

4.49

3)

8

82.07

62.11

75.49

Bay-breasted Warbler (Dendroica castanea)

Zeep

Blackburnian Warbler (Dendroica fusca)

Blackpoll Warbler (Dendroica striata)

Cerulean Warbler (Dendroica cerulea)

Connecticut Warbler (Oporornis agilis)

Louisiana Waterthrush (Seiurius motacilla)

Magnolia Warbler (Dendroica magnolia)

Worm-eating Warbler (Helmitheros vermivorum)

Yellow Warbler (Dendroica petechia)

148.76

27

@)
28.

Madi

3)

16

3.25

111

86.23 145.31

5

2275.0

741.08

Black-throated Green Warbler (Dendroica virens)
Mourning Warbler (Oporornis philadelphia)

Nashville Warbler (Vermivora ruficapilla)

Up

Vol. 128

Orange-crowned Warbler (Vermivora celata)

Ovenbird (Seiurus aurocapilla)

139

RIE

OF MIGRATORY BIRDS OVER WESTERN LAKE E

MONITORING

2014 SANDERS AND MENNILL

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140

TABLE |. (continued) Number of classified acoustic detections (per 1000 h of recording) and mist net detections (per 1000 h of netting) of migratory birds in spring and fall 2012 at mainland

and island locations in Essex County, Ontario.

Fall migration

Spring migration

Net detections

Acoustic detections

Mainland

Net detections

Mainland

Acoustic detections

Mainland

Bioacoustic

Island

Mainland

Island Island

Island

Species or species group (7 = 38)

category
PAWA
PISI

Palm Warbler (Dendroica palmarum)

Pine Siskin (Carduelis pinus)

12.78 34.34 104.07 OM

5), 12

138.10

30.04

5.81
11.05

0.36

3D
1.78

4.04
921.46

0.73
0

SUS

20.80

Rose-breasted Grosbeak (Pheucticus ludovicianus)

Scarlet Tanager (Piranga olivacea)

RBGR
SCTA
SWTH

10.38
1936.52

6.16
905.17

199.89

8.93
231

5

2

2430.3

ISS Il

QS
11.69

Swainson’s Thrush (Catharus ustulatus)

Veery (Catharus fuscescens)

21.66

78.49
17.44

38.80

7.4

363.42

167.94

VEER

4.46
0.45
D357)

3,12
16.61

6.93
DSM
3

Vesper Sparrow (Pooecetes gramineus)

Virginia Rail (Ra//us limicola)

VESP

THE CANADIAN FIELD-NATURALIST Vol. 128

0
6.11 0.71

14.54

5.38
5.38
0

0
4.38

2.08
75.80
148.48
2727.74

cl

Wilson’s Warbler (Wilsonia pusilla)

VIRA
WIWA
WOTH

0.17

ion

N

2
0
84.03

eS)
T3953
445.27

White-crowned Sparrow (Zonotrichia leucophrys)
White-throated Sparrow (Zonotrichia albicollis)

Wood Thrush (Hylocichla mustelina)

WCSP
WTSP

Va}

VES

1793.67

511.58

N

ras

4.

N

To determine whether some species were present at
the mainland sites, but not the island sites, we exam-
ined both the acoustic and banding data for the pres-
ence or absence of each species or species group in
both spring and fall.

To determine whether the date of first detection on
the island was different from that on the mainland, we
used Wilcoxon sign-rank tests to compare average ar-
rival dates for all species. We pooled the data for the
four mainland recorders and calculated the average first
detection date for all species or species groups at the
mainland sites, and repeated this process for island re-
corders and sites. We performed a parallel analysis using
banding data to compare first arrival dates at the island
and mainland banding stations in spring and fall.

Results

We collected 2261 h of recordings over 58 nights at
the seven recording locations during the spring migra-
tion and 3960 h of recordings over 63 nights at the
same locations during the fall migration, for a total of
6237 h of recordings over both seasons. In these record-
ings, we noted 60 013 nocturnal flight calls: 22 554 dur-
ing spring migration and 37 567 during fall migration.
We were able to classify, with confidence, 46 846 (78%)
calls to species or species group: 16 646 (74%) spring
recordings and 30 200 (81%) fall recordings. There was
substantial variation in the number of night-flight call
detections at the various recording locations in spring
and fall (Table 2).

During the spring migration, 834 birds were banded
at the island site and 910 birds were banded at the main-
land site. During the fall migration, 2079 birds were
banded at the island site and 2711 birds were banded at
the mainland site.

Number of migrants detected on mainland versus island

During spring migration, we detected more night-
flight calls on the island than on the mainland (Table 1):
35 species or species groups were more abundant on
the island; only two species — Vesper Sparrow (Po-
oecetes gramineus) and Wilson’s Warbler (Wilsonia
pusilla) — were more abundant on the mainland (sign
test, P < 0.0001, n = 37); two of the species indicated
in Table 1 were not detected on either the mainland or
island recorders in spring: Golden-crowned Kinglet
(Regulus satrapa) and Pine Siskin (Carduelis pinus).

Similarly, during fall migration, we also detected
more calls on the island than on the mainland (Table 1):
35 species or species groups were more abundant on
the island; only three species — Pine Siskin, Scarlet
Tanager (Piranga olivacea), and Virginia Rail (Rallus
limicola) — were more abundant on the mainland (sign
test, P< 0.0001, n = 38; cuckoos were not detected by
either mainland or island recorders in fall). :

Species detected on mainland versus island
Acoustic monitoring data revealed that the same
composition of species passed over the mainland and

2014 SANDERS AND MENNILI

f acoustic detections (including both classified and unclassified detections) per 1000 h recording time during spring and fall migration, 2012, at locations in

2. Total number o

TABLI

Essex County, Ontario (see Figure 1).

Detections/1000 h

Total number

No. of recording

recordings

of calls*

Spring

hours

Fall
3192.2

Spring

Fall

Fall
576.40

Spring

Location

Recorder

1828.8
5242.6

1840

600

328.08

Holiday Beach Conservation Area

Mainland |

10131.6
10350.5

4

1720
3633

521.34

328.08

, Ontario

Cedar Creek Conservation Area
Point Pelee National Park

=)

Private woodlot near McGregor

Mainland 2

11575.2
3410.8

5966
2568
6927

576.40

313.86

Mainland 3

~
fos]
va)

1119
5296

567-15

328.08

Mainland 4

| MONITORING OF MIGRATORY BIRDS OVER WESTERN LAKE ERIE 14]

12230.5
19826.5
6169.3

25529

566.37

306.91

Fish Point Provincial Nature Reserve, southern Pelee Island
Fish Point Provincial Nature Reserve, southern Pelee Island

Island |
Island 2
Island 3

Total

1990.4
29056.9

11428
3556
37567

653
9583

576.40

328.08

576.40

328.08

Lighthouse Point Provincial Nature Reserve, northern Pelee Island

3960.46 22554

2261.17

the island (Table 1). We found no evidence in the
acoustic recordings that any species avoided crossing
Lake Erie. In spring migration, all species or species
groups were detected in both mainland and island
recordings; in fall migration, American Robin (7urdus
migratorius) and Wood Thrush (/ylocichla mustelina)
were recorded only on the island, and a single Virginia
Rail (Rallus limicola) was recorded on the mainland,
with none on the island.

The banding data show a similar pattern in that most
species or species groups were captured in both main-
land and island mist nets (Table 1). In spring migra-
tion, the “double-down” species group and the Fox
Sparrow (Passerella iliaca) / Song Sparrow (Melospiza
melodia) species group were captured in mainland but
not island mist nets, whereas Hooded Warbler (Wilso-
nia citrina), Mourning Warbler (Oporornis philadel-
phia), and White-crowned Sparrow (Zonotrichia leu-
cophrys) were captured in island but not mainland mist
nets,

In fall migration, the “double down” species group
and the Fox Sparrow / Song Sparrow group was cap-
tured on the mainland, but not in island mist nets,
whereas a Rose-breasted Grosbeak (Pheucticus ludovi-
cianus) was captured on the island but not in the main-
land mist nets. Several species (American Robin, Wood
Thrush, and the Cuckoos) were netted in spring but
not in the fall.

Date of arrival

In the spring, species were first detected significant-
ly earlier at the island recorders than on the mainland
(average ordinal date of first detection on island:
114.1 + 1.6; mainland: 117.1 + 1.6; Wilcoxon sign-rank
test: W = 154.5, P = 0.0006, n = 36). This matched
our prediction that birds would be detected earlier at
the more southerly site during northward migration.
However, in the fall, there was no significant difference
between dates of arrival on the mainland and the island
(average ordinal date of first detection on island: 254.6
+ 1.8; mainland: 258.1 + 2.8; Wilcoxon sign-rank test:
W=73.5, F = 0.09 17 = 36).

Based on data from the banding stations, we found
no significant difference in date of arrival during spring
(average ordinal date of first detection on island: 119.8
+ 1.8; mainland: 121.2 + 1.9; Wilcoxon sign-rank test:
W = 0.5, P= 0.99, n = 22) or fall migration (average
ordinal date of first detection on island: 248.4 + 2.0;
mainland: 252.0 + 5.0; Wilcoxon sign-rank test: W =
36.0, P= 0.28, n = 25).

Discussion

Acoustic recordings of nocturnal migrants over west-
ern Lake Erie showed that migratory birds do not avoid
crossing the lake and suggest that birds may concen-
trate over Pelee Island mid-way across the lake on both
their northward and southward journeys. Island micro-
phones detected significantly more calls in both spring
and fall than the microphones located nearby on the

north shore of Lake Erie. Migrant communities on the
mainland and island were similar in composition, al-
though subtle seasonal differences in the relative abun-
dance of each species were evident. Species were detect-
ed by acoustic recorders earlier in the spring on the
island than on the mainland. Based on the differences
in the number of birds and the timing of migration, we
conclude that Lake Erie does not represent an ecolog-
ical barrier to migratory birds. Although our results
suggest that a migratory route across Pelee Island is
especially important in this region, acoustic record-
ings collected over the open water of Lake Erie will be
required to confirm whether birds concentrate over the
island during migration.

Thousands of night-flight call detections revealed a
substantially greater number of calls over Pelee Island
than over mainland Ontario during both spring and fall
migration. This implies that birds use the island archi-
pelago in large numbers during migration. Pelee Is-
land’s geography could act to concentrate birds as they
approach the north or south tip of the island (Farns-
worth 2005). Point Pelee National Park, a peninsula that
projects into Lake Erie, may also concentrate birds as
they migrate south in the fall (Lincoln ef al. 1998*),
although our acoustic recorder in the centre of this
peninsula (Figure 1) did not detect significantly more
migrants than the other mainland sites (Table 2). The
theory that peninsulas concentrate migratory birds is
supported by the fact that more calls were detected by
the microphone at the north end than those at the south
end of Pelee Island in spring and the reverse in the
fall (Table 2), although this is an anecdotal observation.

We interpret the higher number of acoustic detec-
tions over Pelee Island than over the mainland in both
seasons as evidence that more migrants pass over the
island, but this pattern could also arise if birds increase
their calling rate as they pass over islands. Although
the function of night-flight calls is still under debate,
they may be important in maintaining flock cohesion
and communicating directional information to flock-
mates (Hamilton 1962; Farnsworth 2005). As birds ap-
proach a shoreline and detect open water, this may be
an especially opportune time to communicate, perhaps
to aid orientation among flockmates, leading to higher
numbers of flight calls (Farnsworth 2005). However,
confirming this pattern requires further investigation at
multiple positions, such as microphones mounted in
the middle of the lake away from islands, for direct
comparison with island recordings. This is a challeng-
ing area for future research.

It is also noteworthy that radar research suggests that
migrating birds may decrease their altitude as much
as 9% when crossing bodies of water (Bruderer and
Liechti 1998); this might make them more likely to
be detected by our island microphones than mainland
microphones and explain the greater number of calls
detected on the island. However, radar data suggest that
Great Lake islands are important parts of the migratory

THE CANADIAN FIELD-NATURALIST

Vol. 128

flyway (Diehl et a/. 2003); thus, these alternative expla-
nations alone cannot account for the higher number of
migrants detected on the island.

All songbird species or species groups were detected
by the island recorders in both spring and fall, and mist
net data contain only subtle differences. This suggests
that the lake is not an insurmountable barrier to any
of the nocturnal migratory species or species groups
that we studied. Previous radar studies demonstrate
that nocturnal migrants cross Lake Erie in significant
numbers in both spring and fall (Diehl e¢ al. 2003),
although they do not reveal which species are cross-
ing the lake. All species and species groups that we
could distinguish do indeed cross over Lake Erie in
both fall and spring migration. Further research on sea-
sonal variation in migration paths (especially through
the use of geolocators; e.g., Stutchbury et a/. 2009), as
well as multi-year studies that explore annual variation,
will provide important insight into the migration of
birds through the Great Lakes.

In the spring, the first birds of each species or species
group were detected earlier by the Pelee Island re-
corders than those on the mainland. This matched our
expectation and is consistent with the seasonal direction
of migration. However, contrary to our expectation, we
did not find the reverse pattern during fall migration,
when the dates of first detection were not statistically
different at the mainland and island sites. Further inves-
tigation 1s required to explore whether this anomaly is
due to Lake Erie serving as a temporary barrier to
spring migrants, but not to fall migrants. Further exam-
ination of the stopover behaviour of birds on Pelee
Island might determine whether they are spending a
longer time on the island in the spring than in the fall,
which might produce the significant delay in detection
on the mainland. This idea stands at odds with the gen-
eral consensus that the spring migration window is
shorter, with many species flying faster and stopping
for briefer periods (e.g., American Redstart, Setophaga
ruticilla, Morris and Glasgow 2001). Furthermore, our
banding data showed no significant differences between
mainland and island sites in first arrival date in spring
and fall.

Although Lake Erie may not be an insurmountable
barrier to migrants, poor weather conditions may ampli-
fy the risk of crossing open water. Wind strength and
direction, visibility, cloud cover, and temperature play
a role in determining the intensity of migration events
(Elphick 2007; Gagnon er al. 2011). Low cloud cover
may concentrate migrants at lower altitudes, resulting
in higher acoustic detection rates (Evans and Mellinger
1999). Furthermore, artificial light may cause disori-
entation and serve to reorient birds in flight toward the
island or cause individuals to increase their calling rate
(Evans et al. 2007; Poot et al. 2008). Artificial light
may have a more pronounced effect under poor or de-
teriorating weather conditions when stars are obscured
by cloud cover (Farnsworth and Russell 2005: Htippop

2014 SANDERS AND MENNILL: MONITORING OF MIGRATORY BIRDS OVER WESTERN LAKE ERIE

and Hilgerloh 2012). An investigation into the com-
position of nocturnal migrants detected over Pelee
Island on nights preceding and during strong winds or

precipitation could lead to a better understanding of

which species are most affected by poor weather while
navigating the Great Lakes.

In the western basin of Lake Erie, acoustic and band-
ing data suggest that Pelee Island is an important geo-
graphic feature for migratory birds. Many individuals
of all species passed over the island in spring and fall,
with little indication that they avoided crossing Lake
Erie, suggesting that islands in the middle of poten-
tial geographic barriers are important for migratory
birds. Given that many species of North American
migrants are in decline, studies that monitor migratory
bird behaviour at smaller geographic scales are impor-
tant for local management and development decisions
in regions where birds are known to concentrate sea-
sonally.

Acknowledgements

We thank M. Watson for assistance in collecting
recordings. We thank C. Berek, A. Demarse, K. Drea,
R. Gough, C. Lajoie, J. Mahal, A. Mamo, L. Mehdi, B.
Poisson, M. Watson, and A. Weinz for assistance with
annotating recordings. We thank the Holiday Beach
Migration Observatory for their ongoing volunteer-
based efforts to study avian migration in southwestern
Ontario. We appreciate the work of bird banders on
Pelee Island, and we thank the Canadian Wildlife Serv-
ice of Environment Canada for providing banding data
from Pelee Island. We thank the Essex Region Con-
servation Authority, Ontario Parks, and Parks Canada
for access to conservation areas and parks. We thank
C. Francis and an anonymous reviewer for comments
that improved this manuscript. Support for this study
was provided by the Natural Sciences and Engineering
Research Council of Canada, the Canada Foundation
for Innovation, the Government of Ontario, the Univer-
sity of Windsor, and Caldwell First Nation.

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Received 7 October 2013
Accepted 20 November 2013

Parental Care by Lone Male Ferruginous Hawks (Buteo regalis),
Rough-legged Hawks (Buteo lagopus), and Great Horned Owls
(Bubo virginianus) was Limited to Providing Food

ae = 1.5 oe N 3
JOSEF K. SCHMUTZ!:°, MARTIN A. GERARD?, GORDON S. Court?, and R. WAYNE NELSON?

Qe Ea erie efainaniy : ; >< (8

School of Environment and Sustainability, University of Saskatchewan, 15 Campus Drive, Saskatoon, Saskatchewan S7N 5C8
Canada

2! Caron Crescent, Grasswood, Saskatchewan S7T 1A8 Canada

‘Departunent of Environment and Sustainable Resource Development, 9920 108 Street, Edmonton, Alberta TSK 2M4 Canada

‘4218 63rd Street, Camrose, Alberta T4V 2W2 Canada

>Corresponding author: joe.schmutz@usask.ca

Schmutz, Josef K., Martin A. Gérard, Gordon S. Court, and R. Wayne Nelson. 2014. Parental care by lone male Ferruginous
Hawks (Buteo regalis), Rough-legged Hawks (Buteo lagopus), and Great Horned Owls (Bubo virginianus) was limited
to providing food. Canadian Field-Naturalist 128(2): 145-150.

In three long-term studies of Ferruginous Hawks (Buteo regalis), Rough-legged Hawks (Buteo lagopus), and Great Horned
Owls ( Bubo virginianus), we observed rare but regular occurrences of superabundant prey at nests where nestlings were hungry,
emaciated, or dead. In these cases, a male appeared to be the lone parent; the female parent was found dead, kept away by dis-
turbances, or simply absent. We conclude that the male parents, whose normal role is to provide food, were unable to expand
their care to include morseling, feeding, and brooding. Faced with the stress of incessant food begging by nestlings, the male
parents responded by bringing more and more food to the nest, to the point where food spoiled while the young starved amid
plenty. We provide and interpret detailed observations for 11 such cases and suggest several variables that would influence the
nestlings’ chances of survival.

Key Words: Ferruginous Hawk; Buteo regalis; Rough-legged Hawk; Buteo lagopus; Great Horned Owl; Bubo virginianus:

parental care; food provisioning; feeding; brooding; starvation; Rankin Inlet; Nunavut; Hanna; Alberta; Saskatoon;

Saskatchewan

Introduction

Studies of birds have revealed fascinating patterns
of mate choice and parental care, including elucidation
of the apparent ecological correlates that predictably
lead to such patterns (e.g., Cockburn 2006). Typical pat-
terns include brood parasitism; male only, female only,
and bi-parental care; and cooperative breeding (Cock-
burn 2006).

Care of young by monogamous pairs is the most
common pattern among raptors and birds in general;
81% of all species exhibit it. Among raptors, three over-
lapping phases occur: at first, the female assumes brood-
ing and shading duties almost continuously; later, the
female leaves the partly grown young for part of the
time to roost or even hunt nearby; and, finally, both
male and female may be away hunting when the young
approach fledging (Newton 1979, page 160).

A difference in body size influences the way in which
dominance relationships and parental roles are appor-
tioned within a raptor pair. Although a difference in size
between females and usually larger males is common
among animals (e.g., Blanckenhorn 2005), this is re-
versed among raptors with females about a third larger
than males (e.g., Olsen and Olsen 1987). The nest 1s
the female’s domain, and it is common among broad-
winged hawks, and possibly less so among falcons, for
the male to spend little time at the nest (Newton 1979,

page 159).

Using an inductive style of inquiry (e.g., Schmutz
1992), we report on observations at nests and empha-
size context and detail in our field observations and
related literature. We examine a male’s limits in parental
care and ask whether a male parent can adequately com-
pensate for the loss of a female and under what condi-
tions this may be possible.

Study Area and Methods

Ferruginous Hawks (Buteo regalis), studied near Han-
na, Alberta, were in moist, mixed-grass prairie where
rangeland grazing of cattle was the dominant land use.
Richardson’s Ground Squirrel (Urocitellus richara-
sonii) was the hawks’ main prey. Our observations were
part of a long-term population study varying in inten-
sity from banding and monitoring of breeding densi-
ties each year to detailed monitoring of prey, growth of
nestlings, and survival and dispersal of colour-marked
young and breeding adults (Schmutz ef al. 2008). The
number of nests and visits to nests varied accordingly,
from near daily during the nestling period to once a sea-
son. The use of an intensively studied core area and a
peripheral area also affected the number of visits.

Observations of Rough-legged Hawks (Buteo lago-
pus) were made in the vicinity of Rankin Inlet, Nunavut,
within an area defined for the long-term investigation of
the breeding biology of Arctic Peregrine Falcons (Falco
peregrinus;, Court et al. 1988).

145

146 THE CANADIAN FIELD-NATURALIST Vol. 128

Great Horned Owl (Bubo virginianus) studies were 2
conducted from a base in Saskatoon, Saskatchewan. a 5
Nests were reported by area residents or spotted from < = =
roads while trees were still devoid of foliage. Once & sl a
located, nesting areas were visited annually to deter- = si ie
mine occupancy. Owlets were banded and prey items g 3 ii Be eee
recorded (Gerard et al. 2009). ss) 2 6 i g o 3
For convenience, we refer to cases or nests by num- : = 5 Z a) as ee
ber, preceded by F, R, or G, for Ferruginous Hawks, as = = Na 5 ts eB a c=
Rough-legged Hawks, and Great Horned Owls, respec- @, 5 E s ae 2 2228
tively (Table 1). » 3] 88338 $s S2588
= 4 yee 8) Sa Se SS
on
Results 5
Observations of parental care deficits were rare in ES < 5
our studies (< 1% in over 5000 visits to all nests) and z z =
occurred at different stages in the nestling period and g : Bota tae Gain Sees = Be
' : oO = oS: © © fey 5) SaONDgy OO
in different years. S >) S555 55 B2SESS
Ferruginous Hawk nests os Bh Se ete eS ae
F-1: On our arrival at the study site on 9 June 1975, 5
two nestlings less than 1 week old were peeping as if a
hungry, although prey was in the nest and a dark morph 3 3
female, judging by size, was roosting on the nest rim. 2 ‘5 >
On I! July, both nestlings were dead, a light morph 2 I = |
Ferruginous Hawk was circling overhead, and the dark Se a0 =
morph bird was not seen. In the nest, we found 14 dead 805 Bb 5
Richardson’s Ground Squirrels. Our interpretation was S aS, a 8 is
that the nestlings vocalized out of hunger, to which the BS a a 3. 2
male parent responded by bringing more food but did ees é 3 WS a 5 Eps
not feed them. ces aes 3 Z he
F-2: On 30 June 1989, a mass of about 20 Richard- she 2 Nee aes 2925
son’s Ground Squirrels was found in a nest used during S n s| Psex = a es Bic
that year, but without young or adults visible at the Ss z a = q 2 = 23 = 4
time. Many of the squirrel bodies were whole. They had Bgf4| Boat 5H =-Z2-zZz
decayed and dried, suggesting that they had been cap- ia é
tured and deposited 2-3 weeks earlier. This would have 2e
placed the prey deliveries about mid-way through the & § =
normal nestling period for Ferruginous Hawks in that 6 & & 5 3 g
year. 5 °= |] 2 F% ,» 22 Baouane
F-3; On II June 2002, a nest with two nestlings and 38 = B 5 5 s 3 3 2 3 3 3 3
no prey was attended by a light morph Ferruginous 26 |Z be Sess a E ef L3 a Laas
Hawk circling overhead. On 4 July, one nestling was 3 =
dead and one alive. The live nestling gave monosyllab- ZO
ic, flutelike calls suggesting hunger (see Powers 2003, Sai 3S 3
page 98). Of 30 partial or whole ground squirrels in < 5 5s £ 8
the nest, 11 were gutted but otherwise uneaten, yet no . 3 = 3. =
parent was visible. The live nestling weighed 905 g at 5 5 z 2a 8 S
a primary length of 150 mm, the dead nestling 600 g at 3 ey = 5 S. pe
132 mm (see Schmutz 1977, Appendix 3 for age deter- es ae ag & &
mination). Assuming that the emaciated nestlings were as Ss te z s 2p ©
smaller-bodied males, they were still 180 and 470 g a= ie] 2 ¥ = D 2S Saha
below normal weight for their age, respectively. The S S =| 2 ies 3 a 3 5 & SS&66§
most plausible interpretation was that one nestling had roe c\$sss : ne a Saaaaa
starved to death and the second was near death. Ba wale lle a= 222222
F-4: On 8 June 1976, a single, distant adult called in ES = es 8
defence of a nest where a 410-g nestling emitted peep- 5 5 S S, g
ing sound (presumed hunger), although seven Richard- Se ile iS £ S
son’s Ground Squirrels were present. On 10 June, the = fs s = So 3
nestling was dead with the prey still in the nest. We Se lzit hie eee So ey 3 oe

2014 SCHMUTZ ET AL.: LIMITED PARENTAL CARE BY MALE RAPTORS 147

assumed that a nestling reared to 11 days must have
had a female parent initially, and, later, the remaining
adult was a male unable to feed the nestling.
Rough-legged Hawk nests

R-1: From 29 to 31 July 2000, a blind was installed
for photographic purposes approximately 15 m from
the nest of a pair of Rough-legged Hawks, about 10 km
north of Rankin Inlet, Nunavut. The pair had three

young, approximately 3 weeks of age. Each member of

the pair was easily identified to sex based on plumage.
The adult male was apparently not affected by the
installation of the blind and delivered several Siberian
Brown Lemmings (Lemmus sibiricus) to the nest each
day when photographers were present. At no time was
the male observed attempting to feed nestlings, and the
nestlings did not feed themselves. The adult female did
not attend the nest when photographers were in the
blind, and no feedings were witnessed during photo-
graphic sessions lasting up to 6 h. The young were
apparently fed by the female after the photographers’
departure; when the photographers arrived to film each
day, the previous day’s food was absent and the female
flushed on their arrival. We recorded several prey deliv-
eries by the adult male over the 3 days. Our interpre-
tation at the time was that male Rough-legged Hawks
do not feed young despite ample opportunity and appar-
ent need.

R-2: On 8 August 2005, a visit to a Rough-legged
Hawk nest on the Barrier Islands, Rankin Inlet, revealed
only an adult male present. The female parent was not
recorded at the territory during the visit, which lasted
more than 1.5 h, unusual for this species at this location
and time of year. The nest contained one live but ema-
ciated nestling and two dead nestlings that were ex-
tremely emaciated and had apparently starved to death
(Alastair Franke, Research Associate of the Circum-
polar Institute at the University of Alberta and the
Principal Investigator for Arctic Raptors, personal com-
munication, 8 August 2005). The living nestling was
standing on or beside the carcasses of at least five un-
eaten Arctic Ground Squirrels (Spermophilus parryi)
and one Siberian Brown Lemming. We concluded from
these observations that the female had died and that the
male was not feeding the starving young, despite the
availability of ample prey.

Great Horned Owl nests

G-1: On 29 April 2002, a telephone call alerted us
to three orphaned Great Horned Owl nestlings. On 26
April 2002, a female had been electrocuted on a trans-
former, which subsequently required repair. On MAG’s
arrival, a male flushed from a roost and came to within
5 m of the nest tree in a farm shelterbelt. Three young,
approximately 12, 16, and 20 days old, were in the
nest, along with a whole, fresh, male Western Mead-
owlark (Sturnella neglecta). As the young were mak-
ing hunger calls, MAG fed the meadowlark to the
two larger owlets. The youngest owlet was removed

and introduced into a foster nest with only one nestling
near Saskatoon. On | and 3 May 2002, the remaining
two young were reportedly doing well. We assume that
the two nestlings were large enough to feed themselves,
possibly by consuming small prey whole.

G-2: On 6 May 2002, a nest 6 m above ground in a
Trembling Aspen tree (Populus tremuloides) was snow
covered after a snowstorm the previous day. The nest
contained a freshly dead, 400-g Great Horned Ow]
nestling and a shivering 660-g nestling, plus the whole,
fresh carcass of a Black-billed Magpie (Pica hudsonia).
The living chick appeared to be cold and was “chit-
tering” continuously. It readily ate offered morsels of
the magpie. A shy adult male owl was observed nearby
— apparently the sole survivor of the nesting pair —
and was behaviourally unable to brood and tear prey for
the nestling. We believe we saved or prolonged the life
of the second nestling by removing snow and feeding it.

G-3: On 13 May 2005, a female owl that had been
banded at this nesting area in 2002 was found dead
below a nest containing one young. The female was
partly decomposed and estimated to have been dead
for 7-10 days. Prey, including a blackbird (Icterinae),
a rabbit head (Lagomorpha), and a Grey Partridge
(Perdix perdix), were on the ground below the nest,
and a shy Great Horned Owl, judged by size to be a
male, was nearby. At banding time on 2 June 2005, the
estimated 7-week-old fledgling owl flew poorly. The
adult male was still present and being mobbed by
American Crows (Corvus brachyrhynchos). It was pro-
tective of the young and aggressive toward our black
Labrador dog. By removing the dead female, we may
have encouraged the male to place prey in the nest sub-
sequently.

G-4: On 6 May 2008, we observed a nest containing
one dead and three live owlets, estimated to be 25, 28,
and 30 days old. An adult owl attended the nest, but
without venturing from nearby cover. The young were
vocalizing as though hungry. The nest contained three
young Snowshoe Hares (Lepus americanus), five
Northern Pocket Gophers (Thomomys talpoides), one
Northern Shoveler (Anas clypeata), one Long-eared
Owl (Asio otus), and the remains of a Northern Har-
rier (Circus cyaneus). The hungry young consumed a
whole hare and part of the shoveler when these were
offered in morsels. The smallest nestling was fed and
introduced into a foster nest attended by two adults.
We took signs of hunger despite superabundant food
to indicate that parental care was compromised. All re-
maining prey appeared to be too large for the young to
swallow whole or tear apart on their own.

G-5: On 5 May 2012, two 4-week-old owls were
banded. Their nest contained eight American Coots
(Fulica americana), one Gadwall (Anas strepera), one
rabbit, and one Virginia Rail (Ra/lus limicola). Some
of the coots contained maggots. A male adult owl was
in attendance. Several days later, the nest was revisited
and, again, only a male was in attendance. At 4 weeks

148

of age, we assumed that the young owls were able to
feed themselves, possibly swallowing small prey when
offered while leaving the large coots, ducks, and rabbit
uneaten.

Discussion
Is the males parental care role inflexible?

A common theme throughout these observations was
food in all 11 nests and so much excess food that it
spoiled in five of them (Table 1). Another common fea-
ture was the presence of a male parent only, except in
nests R-1 and, possibly, in nest F-1 with F-2 unknown.
Adult gender was judged based on raptor size, plumage,
in-hand inspection (G-1, G-3), in-flight/roosting obser-
vation (F-1, G-2), or presumed based on the fact that
the food-provision role (male) was satisfied while the
brooding and feeding role (female; Bechard and
Schmutz 1995; Houston ef al. 1998: Bechard and
Swem 2002) was not (F-3, F-4, R-2, G-4, G-5).

We know that two females had died (G-1, G-3). We
assume that other females had also died (F-1, F-2, R-2,
G-2, G-4, G-5) or that they were somehow kept away
from the nest (e.g., R-1). Nestlings were starving, hun-
gry, or cold, indicating that females were absent for at
least several crucial hours (e.g., during a snowstorm)
and for the days it took the males to amass excess prey,
some of which had begun to spoil. We concluded that
the female member of the pair was lost and that the
male was induced to keep hunting by the hungry young.
Males were unable to make the needed shift to feeding
and brooding.

We are cognizant of the additional possibility that
some nestlings may have died because of congenital
problems or poisons in the environment (e.g., Schmutz
et al. 1989) and that such nestlings voice physical pain,
which would induce the male to continue hunting. This
is unlikely in cases where nestlings survived or were
transplanted (F-3, G-1, G-3, G-5). In case of F-3, the
surviving nestling fledged from its foster nest, although,
later, its long-dead carcass was found. Its early near
starvation state may have compromised normal devel-
opment, preventing this nestling from maturing normal-
ly and becoming able to hunt and survive on its own.

Flexible parental roles in related species

Kenward (2006) summarizes observations and re-
ports that suggest some Northern Goshawk (Accipiter
gentilis) males are also unable to care fully for their
young when the female is lost. As in our cases, young
starve in the face of ample food. Also similar to some
of our owl cases, however, young Goshawks fledged
from five nests where females disappeared at nestling
age 12-20 days, when they begin to feed themselves.

When nestlings are very young and exposed to
weather, the ability of their parents to brood is as im-
portant to their survival as the availability of food. In
nest G-2, we believe that the failure of the male owl to
brood the young during a snowstorm led to the death
of one nestling; the other was likely saved by our clear-

THE CANADIAN FIELD-NATURALIST

Vol. 128

ing snow off the nest. For comparison, RWN has ob-
served a male Peregrine Falcon flying to a nest when
the female had left temporarily and begin to cover
and brood the young. Similarly, JKS observed a male
Swainson’s Hawk (Buteo swainsoni) cover the eggs in
a nest after the male had brought prey which the pre-
viously incubating female accepted from him and ate
atop a fence post.

In clear contrast to our observations of limited par-
ental care, a study of a captive male Red-tailed Hawk
(Buteo jamaicensis; Hamerstrom and Hamerstrom 1971)
suggests that males of this species can participate in
all facets of reproduction. The hawk studied had been
taken from its nest at the nestling stage and used for
falconry. Its gender was confirmed by ejaculation elicit-
ed by the falconer on whom the hawk was imprinted.
The male hawk wove sticks into a nest base, formed
a lined nest cup when straw was provided, covered a
chicken egg, and developed a 5-cm * 6-cm brood patch,
but without significant blood vascularization. The male
hawk accepted a total of three Red-tailed Hawk nest-
lings 1, 6 and 21 days old offered over 2 years. Without
prior experience, it responded positively to the vocal-
izations of the newly introduced | and 6-day-old nest-
lings and displayed great care closing its talons and slid-
ing mostly on his tarsus over the nestlings to brood. It
fed the nestlings pieces of flesh until they were 9 days
of age. The pieces were not regurgitated, but still moist
from saliva that seemed to flow more during feeding.
After 9 days, the male tended to lay small pieces of meat
on the nest rim for the nestlings to eat. By 24 days, the
nestlings began to tear and consume mice and day-old
chickens on their own. When the nestlings were 16 days
old, the male gradually stopped brooding them at night,
in a mid-May Wisconsin climate.

It may be surprising that hawks of the same genus
differ so greatly in terms of parental care. The Red-
tailed Hawk described above capably conducted all
facets of parental care, including the development of
a rudimentary brood patch. In contrast, we observed
Ferruginous and Rough-legged Hawks, and Great
Horned Owls, fail to tear prey and feed morsels to beg-
ging nestlings, a simple act that might have prevented
their starvation. Species-specific behaviour repertoires
may vary to that extent. Another possibility is that plas-
ticity in parental care exists initially, but is lost through
experience and discouraged by the dominant female.

Are female roles similarly inflexible?

At a different nest, a series of observations sug-
gested that a female Ferruginous Hawk was able to
assume the food provisioning role of males and raise
her brood alone. Here, a male was observed with the
female on two visits (6%) just before eggs hatched,
then not again for the remaining 32 visits during the
nestling period. This deviated from the typical pattern
of two adults defending a nest in 38% of 164 Visits in
IW (G—9A2 P—0.002) Prey was less often present
in the female-only nest, on only 21% of 29 nest checks

2014

compared with 54% of 420 visits (G = 10.77, P = 0.001)
to all other nests. Despite the lone attendance of the
female parent and scarce prey in the nest, the young
appeared as well fed as others. At least one young had
food in its crop on 57% of 7 crop records, compared
with 41% of 96 crop records (G = 0.72, P = 0.396) from
nests with both parents in attendance.

These results suggest that this female Ferruginous
Hawk was able to both hunt and care for her young,
raising two young from four eggs to-fledging. She like-
ly left her young exposed during favourable weather,
hunted when the young were hungry, and fed them im-
mediately on return. This pattern would account for the
reduced frequency at which food was found in the nest,
yet also account for well-fed young with an average
rate of food present in their crops. This male-loss or
abandonment occurred in a year of exceptional food
availability (Schmutz et a/. 2008).

Is the mating system of Ferruginous Hawks female-
centred?

In a female-centred mating system, the male is not
only dependent on the female for reproduction, but also
takes its cues from the female. Some observations sug-
gest that this is the case in Ferruginous Hawks.

Observations from a blind in 1990, by Janet Foster
(co-owner of John and Janet Foster Productions Ltd.,
personal communication, 26 June 1990) confirm the
nestling-care role of females (see also Powers 2003).
On several occasions, all four nestlings patiently wait-
ed their turn without struggle to be fed. Only once did
two or more nestlings pull on the same piece of food
offered by the female. When the offered piece of food
was too large and the nestling struggled to swallow, the
female took it back and ate the food herself. These
nestlings were 28-31 days old at the time.

Older nestlings are able to feed on their own. Observ-
ing the same nest as above from a blind in 1989, Dan
Wood (owned Wild Prairie Photographics Ltd., person-
al communication, 7 July 1989) observed a male, judg-
ing by size, leave a hill where it and the female had
been roosting. The male returned | h later and delivered
a ground squirrel directly to the nest, and then rejoined
the female on the hill. One of the three young took the
delivered ground squirrel from the nest rim and even-
tually all three nestlings fed on their own at ages 39
and 41 days.

A film sequence taken without audio in 1990 at the
nest mentioned above shows male and female Fer-
ruginous Hawks roosting side by side on the nest rim
(Foster and Foster 1992*). Twice, the female turned her
head toward the male with crown feathers erect and
beak opening, suggestive of a dominance cue, where-
upon the male left the nest to land on a branch in the
same tree only 2 m away. This behaviour combined
with the pattern of a male’s rare and brief presence at
nests with young, suggests that the female discourages

the male from tending young.

SCHMUTZ ET AL.: LIMITED PARENTAL CARE BY MALE RAPTORS

149

Other observations support the female’s primary role
where the nest and young are concerned. In dozens of
images obtained using time-lapse photography of nests
(JKS, unpublished), in only one case did both male and
female roost on the nest rim together; usually it is only
the female. Similarly, Powers (2003) describes adult
males delivering prey directly to the nest, as described
by Dan Wood above, but “he never tarried at the nest
but sat only briefly before departing within a few sec-
onds of his food delivery, presumably to hunt for more
food” (Powers 2003, page 67). Powers (2003, page 93)
concluded that a male remained only 5—15 s at a nest
after a prey delivery, and the longest observed stay was
5 minutes.

In the context of the female’s primary care at the nest
and, possibly, even enforced dominance over the male,
the F-3 male’s inability to channel its care away from
excessive hunting to feeding is plausible. Repeated ex-
clusion from the nest by a female, possibly over several
years, likely reinforced nest avoidance, and flexibility
in nestling care, if any, was lost through learning. This
role development with experience and learned irrever-
sibility could also apply to Great Horned Owls and
Rough-legged Hawks.

Female loss and prey size

Our observations suggest that three broods of Great
Horned Owls (G-1, G-3, G-5) survived in the absence
of a female. Large owls and especially Great Horned
Owls are known for their ability to swallow sizeable
prey whole. This could have been a significant factor
in the survival of some of the owl broods after female
loss. Furthermore, the prey used by the owls was varied
and included smaller items such as voles, mice, and
songbirds.

In contrast, the diet of Ferruginous Hawks and
Rough-legged Hawks (R-2) included primarily relative-
ly large, tough-skinned ground squirrels. Our observa-
tions of the Ferruginous Hawks showed that 28-31-
day-old nestlings still waited for the female to feed
them. At 39-41 days, nestlings tore ground squirrels
and fed on their own.

Nests F-3, F-4, R-2, G-2, and G-4 are noteworthy as
nestlings were hungry and even starving amid plenty
of food. The F-3 nestlings were an estimated 28 and 33
days old, and apparently could not tear and consume the
ground squirrels that were present. The older starving
nestling may have tried to eat on its own as it had dried
blood on its beak.

Even with females present, a parental miscue was
noted by Wayne and Alora Nelson (personal commu-
nication). A captive, 9-year-old, human-imprinted fe-
male Ferruginous Hawk incubated and tended to her
hatchling normally, but persisted in presenting food
morsels 5 cm above the nestling’s beak. The Nelsons
had to feed the young until it was 1 week old and able
to touch the female’s beak, which led to normal feed-
ing by the female soon after.

150

In conclusion, we suggest that starving nestlings
amid a superabundance of food in nests is a rare but
widespread occurrence and that, in these cases, the loss
of the female parent should be considered. We also pro-
vide evidence that male loss is less severe than female
loss. Furthermore, we suggest several variables that
might influence reproductive outcome after female loss:
age of nestlings, prey size related to the ability of nest-
lings to swallow it whole, and, possibly, age of the male
parent. We postulate that first-breeding males may ex-
hibit more behavioural flexibility in feeding young than
males whose role as a mere provider has been en-
trenched by the female over several breeding seasons.

Acknowledgements

We thank landowners for access to their land and
helpful information. We were assisted in the hawk study
by John and Janet Foster, Arnold Grandt, David Moody,
Skylar Rickabaugh, and Dan and Gwen Wood and in
the owl study by Jean and Keith Harris, Stuart and
Mary Houston, Sig Jordheim, Art Lepp, Bruce and
Hilda Noton, Glen Peterson, Kevin and Bergit Pul-
vermacher, Dennis Siegel, Marten Stoffel, Glen and
Mary Walker, and Dan Zazelenchuck. Alastair Franke
(arcticraptors.ca) provided access to the hawk nest at
Rankin Inlet in 2005; the Department of Sustainable
Development, Government of Nunavut, provided ac-
cess to the nest visited in 2000. Ricardo A. Figueroa,
André Gérard, Matthew D. Giovanni, Lynn W. Oliphant,
Katherine McKeever, and three anonymous reviewers
provided helpful comments on the manuscript.

Documents Cited (marked * in text)

Foster, J., and J. Foster. 1992. Wings over the prairie. John
and Janet Foster Productions Ltd. 25 minutes. Prepared for
TV Ontario as part of a nature series entitled The Global
Family. Aired 27 March 1992, 1930 EST.

Literature Cited

Bechard, M. J., and J. K. Schmutz. 1995. Ferruginous Hawk.
In The birds of North America. American Ornithologists
Union and Academy of Natural Sciences of Philadelphia,
Philadelphia, Pennsylvania, USA.

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Bechard, M. J., and T. R. Swem. 2002. Rough-legged Hawk.
In The Birds of North America. American Ornithologists
Union and Academy of Natural Sciences of Philadelphia,
Philadelphia, Pennsylvania, USA.

Blanckenhorn, W. U. 2005. Behavioral causes and conse-
quences of sexual size dimorphism. Ethology 111(11):
977-1016.

Cockburn, A. 2006. Prevalence of different modes of parental
care in birds. Proceedings of the Royal Society Series B
273, 1375-1383.

Court, G. S., C. C. Gates, and D. A. Boag, 1988. Natural his-
tory of the Peregrine Falcon in the Keewatin District of the
Northwest Territories. Arctic 41: 17-30.

Gérard, M, J. B. Clarke, K. Kozi, and D. Zazelenchuk.
2009. A sample of prey remains found in Great Horned Ow]
nests in Saskatchewan in 2008. Blue Jay 67(2): 71-76.

Hamerstrom, F., and F. N. Hamerstrom. 1971. Potential
eines mannlichen Greifvogels (Buteo jamaicensis) in Bezug
auf Nestbau, Britten und Jungenaufzucht. Die Vogelwarte
26: 192-197,

Houston, C. S., D. G. Smith, and C. Rohner. 1998. Great
Horned Owl. /n The Birds of North America. American
Ornithologists Union and Academy of Natural Sciences of
Philadelphia, Philadelphia, Pennsylvania, USA.

Kenward, R. 2006. The Goshawk. T & AD Poyser, London,
United Kingdom.

Newton, I. 1979. Population Ecology of Raptors. Buteo
Books, Vermillion, North Dakota, USA.

Olsen, P. D. and J. Olsen. 1987. Sexual size dimorphism in
raptors: intrasexual competition in the larger sex for a scarce
breeding resource, the smaller sex. Emu 87: 59-62.

Powers, L. R. 2003. A Hawk in the Sun. Dimi Press, Salem,
Oregon, USA.

Schmutz, J. K. 1977. Relationships between three species of
the genus Buteo (Aves) in the prairie-parkland ecotone of
southeastern Alberta. MSc thesis, University of Alberta,
Edmonton, Alberta, Canada.

Schmutz, J. K. 1992. Hamerstrom science from a “gabboon’s”
point of view. Journal of Raptor Research 26: 206-210.
Schmutz, J. K., D. T. T. Flockhart, C. S. Houston, and P. D.
McLoughlin. 2008. Demography of Ferruginous Hawks
breeding in western Canada. Journal of Wildlife Manage-

ment 72: 1352—1360.

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Received 25 October 2013
Accepted 11 January 2014

Apparent Widespread Decline of the Boreal Chorus Frog (Pseudacris
maculata) in Eastern Ottawa

Davip C. SEBURN!:4, KARI GUNSON?, and FREDERICK W. SCHUELER?

'Seburn Ecological Services, 2710 Clarenda Street, Ottawa, Ontario K2B 7S5 Canada
“Eco-Kare International, 644 Bethune Street, Peterborough, Ontario K9H 4A3 Canada
_ishope Mills Natural History Centre, RR#2, Bishops Mills, Ontario KOG 1T0 Canada
Corresponding author: davidseburn@sympatico.ca

Seburn, David C., Kari Gunson, and Frederick W. Schueler. 2014. Apparent widespread decline of the Boreal Chorus Frog
(Pseudacris maculata) in eastern Ottawa. Canadian Field-Naturalist 128(2): 151-157.

The Boreal Chorus Frog (Pseudacris maculata) was once common in the eastern Ottawa area. To assess its current status, we
conducted auditory Surveys at 184 wetlands in 2011 and 2012. Boreal Chorus Frogs were heard at only five (2.7%) of the
surveyed sites. These five sites were spatially aggregated, with only 0.5—7.5 km between any two sites. Sites occupied by
Boreal Chorus Frogs in eastern Ottawa were surrounded by significantly greater agricultural cover (at 1.0-, 1.5-, and 2.0-km
radii), less forest cover (1.0- and 2.0-km radii), and less wetland cover (1.5- and 2.0-km radii) than occupied sites in western
Ottawa. Sites in eastern Ottawa that were apparently unoccupied were surrounded by significantly greater agricultural cover
(only at the 2.0-km radius), similar forest cover (all radii), and less wetland cover (all radii) compared with occupied sites in
western Ottawa. Boreal Chorus Frog populations are commonly subject to extirpation resulting from stochastic events. The
reduced wetland cover in eastern Ottawa may be accompanied by reduced wetland connectivity, making recolonization of
wetlands difficult or impossible. Our data do not show whether wetland connectivity has been reduced, but future research
should address this important topic.

Key Words: Boreal Chorus Frog; Pseudacris maculata; amphibian decline; Ottawa; Ontario; amphibian habitat; wetland

connectivity

Introduction

Amphibian populations are known to be declining
around the world (Wake and Vrendenburg 2008; Collins
and Crump 2009; Collins 2010). A global assessment of
all known amphibian species revealed that a third were
threatened with extinction (Stuart ef a/. 2004). Although
multiple causes have been identified, land use change
resulting in habitat destruction is the leading cause of
amphibian decline and extinction (Collins and Crump
2009; Collins 2010). Amphibian populations have also
declined within protected areas where habitat destruc-
tion is not a threat. Such declines have been linked to
numerous factors including introduced fish species (e.g.,
Knapp and Matthews 2000), disease (e.g., Rachowicz
et al. 2006), pesticide drift (e.g., Davidson 2004), and
climate change leading to reduced hydroperiod in tem-
porary wetlands (e.g., McMenamin ef al. 2008). In
addition, sublethal effects of contaminants such as pes-
ticides may make amphibians more susceptible to dis-
eases (e.g., Davidson ef al. 2007).

In eastern Ontario and western Quebec the Boreal
Chorus Frog (Pseudacris maculata), formerly assigned
to the species Western Chorus Frog (P. triseriata), has
undergone substantial declines. It was once considered
common southeast of Montréal (Bleakney 1959), but
now appears completely absent from that area (Daigle
1997) and is extremely rare in Quebec (Desroches and
Rodrigue 2004; COSEWIC 2008*). It has also become
less widespread in areas of northern New York state
(Gibbs et al. 2005; Corser et al. 2012) and eastern

Ontario near Cornwall (Seburn ef a/. 2008) and appears
to have been extirpated from Vermont (Andrews 2013).
Largely as a result of declines in Quebec, the Great
Lakes/St. Lawrence—Canadian Shield population (under
the name Western Chorus Frog) was designated threat-
ened by the Committee on the Status of Endangered
Wildlife in Canada (COSEWIC 2008*).

The trilling chorus frogs form a clade of closely relat-
ed species within the genus Pseudacris (Moriarty and
Cannatella 2004). A range-wide analysis of the mito-
chondrial DNA of trilling chorus frogs substantially re-
vised our understanding of species limits in this group
and determined that populations in eastern Ontario and
western Quebec should be assigned to the Boreal Cho-
rus Frog species (Lemmon ef a/. 2007). These taxo-
nomic changes have been widely accepted (e.g., Corser
et al. 2012; Dodd 2013), which requires that the liter-
ature on trilling chorus frogs be examined carefully to
determine the current species allocation. Even recent
papers (e.g., Sacerdote and King, in press) do not al-
ways follow the current taxonomy.

Recent surveys of the Boreal Chorus Frog in eastern
Ontario have produced mixed results. Boreal Chorus
Frogs were detected at only 5% of historical locations
surveyed in the Cornwall area (Seburn ef a/. 2008), yet
were found at two-thirds of historical locations in west-
ern Ottawa (Seburn and Gunson 2011), approximately
100 km away. The cause (or causes) of the declines
remains unknown. Although wetland destruction is a
major threat in some areas of Quebec (Daigle 1997;

151

Picard and Desroches 2004*), populations have also
been lost in areas where wetlands remain (Gibbs ef al.
2005, Seburn et a/. 2008, Seburn and Gunson 2011).

The Boreal Chorus Frog is known to have been wide-
spread east of urban Ottawa during the late 1950s and
early 1960s, although precise locations are mainly lack-
ing. Knowledge of the distribution is based largely on
surveys conducted by herpetologist Sherman Bleakney
(1959) and Anne and Garnet Hanes (personal commu-
nication) (Figure 1). The agonistic or territorial call of
a Spring Peeper (Pseudacris crucifer), which is also a
trill (COSEWIC 2008*), can be confused with the call
of the Boreal Chorus Frog, but it is unlikely that the
experienced surveyors of the 1950s confused these two
species.

The first indication that Boreal Chorus Frogs were
rare east of urban Ottawa resulted from a survey con-
ducted by one of us (FWS) along with Aleta Karstad
(Bishops Mills Natural History Centre) and the late
Mike Rankin (Canadian Museum of Nature) from 27
to 30 April 1997. Although Spring Peepers were detect-
ed at 34 sites, Boreal Chorus Frogs were detected at
only two sites (FWS, unpublished data). To assess the
current distribution of the species across this area, we
undertook extensive auditory surveys of roadside wet-
lands and flooded ditches. We hypothesized that land
cover variables, particularly differences in wetland, for-
est, and cropland cover, might explain why the Boreal
Chorus Frog was rare 1n eastern Ottawa and beyond
compared with western Ottawa. As nitrate levels of
10 mg/L or more can cause significant mortality of

Ontario

THE CANADIAN FIELD-NATURALIST

“Russell

Vol. 128

Western Chorus Frog tadpoles (Hecnar 1995), we ob-
tained surface water-quality data to determine whether
nitrate contamination was contributing to local Boreal
Chorus Frog decline.

Study Area and Methods

All surveys were conducted in suburban and rural
eastern Ottawa and western Prescott and Russell coun-
ty (Figure 2), an area of just over 800 km?. Daytime
auditory surveys were conducted during the calling
season of the Boreal Chorus Frog in 2011 (11, 12, 19,
22, and 27 April) and 2012 (20, 21, 22 March and 12,
13, and 18 April). A known site with Boreal Chorus
Frogs in western Ottawa was visited at the start of each
survey to confirm calling was occurring that day. Al-
though Boreal Chorus Frogs in some populations may
not call consistently during the day (FWS, personal ob-
servation), diurnal surveys in western Ottawa suggest
that calling is widespread even among small popula-
tions (Seburn and Gunson 2011). Auditory surveys were
conducted at wetlands and flooded ditches visible from
roadsides, with the exception of a few locations in pub-
lic parks where wetlands away from the road could be
monitored. Surveys lasted 2—5 minutes per site depend-
ing on weather conditions and traffic noise.

We obtained records of 2012 surface water (0.5 m
depth) nitrate and nitrite levels from monitoring sta-
tions across Ottawa from the City of Ottawa. Because
nitrate can be reduced to nitrite in biological systems
(Hecnar 1995), we pooled the readings for both com-
pounds for our analyses. As the City of Ottawa moni-

u
75°20'W

Quebec

45°26'N -

United Counties of
Prescott-Russell

errr

N

A

10
[__} Kilometres

FIGURE 1. Historical records of the Boreal Chorus Frog (Pseudacris maculata) in eastern Ottawa and western Prescott and Russell
Localities were approximated from an unpublished map prepared by Anne and Garnet Hanes that included sites cis

their auditory surveys conducted in 1962 and records from Bleakney (1959) and the Canadian Museum of N

ature.

2014

SEBURN ET AL.: BOREAL CHORUS FROG DECLINE 153

!
75°20'W

Ontario
Quebec -
"A
oe 5) —— e
ie) 69) o re ie
Y OQ ome)
QY °
QO Q OOO
Aes SF?
45°26'N -
oO re re)
Se \\ Ov. ro @
1e)
he 0 5e fe)
((
u > United Counties of- ——
oe fetes Oo Prescotl-Russell
ie) Oo
Russell o

N

10
Kilometres A

FIGURE 2. Location of 184 sites, at which auditory surveys were conducted in spring 2011 or 2012 or both, in eastern Ottawa
and western Prescott and Russell. Open circles indicate sites where the Boreal Chorus Frog (Pseudacris maculata)
was not detected; filled circles indicate the five sites where it was detected.

tors water quality a number of times during the year at
each monitoring station, we took the highest combined
reading of nitrate and nitrite for each station.

The Ontario Land Cover Data Base, Ist edition
(Spectranalysis 1999*) was used to distinguish types
of agricultural cover, specifically to separate old field/
pasture (a habitat that can support Boreal Chorus Frogs)
from cropland (a habitat not suitable for Boreal Chorus
Frogs). This database uses spectral character obtained
from satellite images to classify land cover. Land use
information was also obtained from the Southern On-
tario Land Resource Information System (SOLRIS).
The layer represents the landscape in 15 m x 15 m
pixels and is derived from a combination of satellite
images, topographic maps, and aerial photographs from
2000-2003 (OMNR 2007*).

Three land use classes were used for analyses: forest
cover (> 60% tree cover, including plantations), wet-
lands (> 0.5 ha in area), and agriculture (a broad cat-
egory that includes intensive croplands as well as old
fields and forest opening). We calculated the percent-

age of land devoted to each land use class for areas of

radii 0.5, 1.0, 1.5 and 2.0 km around each survey site.
Land use variables on a larger scale may be correlated
with frog landscape ecology (¢.g.., Gibbs ef al. 2005),
but in this case, a larger radius would result in some
areas extending across the Ottawa River into adjacent
Quebec, which did not seem ecologically meaningful.
Land use data from sites in eastern Ottawa were com-
pared with known Boreal Chorus Frog sites In western

Ottawa (Seburn and Gunson 2011). AreMap 10.0 (Esri,
Redlands, Cal.) was used for all spatial analyses and
Minitab 8.3 (Minitab Inc., State College, Pennsylvania)
for all statistical analyses.

Results

Auditory surveys were conducted at 144 locations
in eastern Ottawa and western Prescott and Russell
in 2011. In 2012, auditory surveys were conducted at
48 of the 144 sites (33.3%) surveyed in 2011 as well as
40 additional sites in the area. In total, 184 sites were
surveyed at least once during the two years (Figure 2).
No Boreal Chorus Frogs were heard at any location in
eastern Ottawa and western Prescott and Russell in
2011. One of us (FWS) heard Boreal Chorus Frogs call-
ing near Russell during other amphibian surveys on
23 March, 2012 (Karstad et a/. 2012; Schueler and
Karstad 2012*). Subsequent surveys focused on that
area to attempt to locate additional sites with Boreal
Chorus Frogs. In total, Boreal Chorus Frogs were heard
at five of the 184 sites (2.7%) surveyed. Eight of the
48 sites (16.7%) surveyed in both 2011 and 2012 had
no standing water in 2012. One additional 2011 site was
in the process of being tile drained when surveyed in
2012. Only one of the 48 sites (2.1%) surveyed in both
years had Boreal Chorus Frogs calling in 2012 but not
ZOLL.

Maximum combined nitrate and nitrite level in sur-
face water was 6.6 mg/L in eastern Ottawa and 6.2 mg/L
in western Ottawa. There was no significant difference

154

between the combined nitrate and nitrite levels in east-
ern Ottawa (median = 1.6 mg/L, n = 34) and in western
Ottawa (median = 1.7 mg/L, n = 25; W= 816.0, P =
(sil sy).

Ground truthing of cropland cover identified by the
Ontario Land Cover Data Base indicated that this data-
base was not reliable. For example, one known Boreal
Chorus Frog site in an old field habitat was classified
as 59% cropland. Therefore, all subsequent analyses
used only the SOLRIS database.

The five eastern Ottawa sites with Boreal Chorus
Frogs were clustered in three patches in an area near
Russell (Figure 2). The distance between these occu-
pied sites ranged from 0.5 km to 7.5 km, and all sites
were separated by at least one road. Despite this spatial
aggregation, land use varied across the five sites. For
the surrounding area within a 1.0-km radius, median
agricultural cover was 64.8% (range: 52.6—75.0%),
median forest cover was 3.7% (range: 2.7—23.4%), and
median wetland cover was 11.2% (range: 3.6-28.2%).

THE CANADIAN FIELD-NATURALIST

Vol. 128

The occupied sites in eastern Ottawa had significantly
greater agricultural cover (within a 1.0-, 1.5-, and 2.0-
km radii), less forest cover (1.0- and 2.0-km radii),
and less wetland cover (1.5- and 2.0-km radii) than
occupied sites in western Ottawa (Table 1). Sites in
eastern Ottawa that were apparently unoccupied had
significantly greater agricultural cover (only for a 2.0-
km radius), no significant difference in forest cover (all
radii), and less wetland cover (all radii) compared with
occupied sites in western Ottawa (Table 2). Many of
the eastern Ottawa sites had low percentages of wet-
land cover. For example, although median wetland coy-
er values for a 2.0-km radius differed by only 2.1%,
30% of eastern Ottawa sites had < 10% wetland cover,
compared with only 14% of the western Ottawa sites.

Occupied sites in eastern Ottawa had greater agri-
cultural cover (2.0-km radius only) than apparently
unoccupied sites in eastern Ottawa, but there were no
significant differences in forest or wetland cover for
all radii (Table 3).

TABLE |. Land use surrounding sites with Boreal Chorus Frogs (Pseudacris maculata) in eastern Ottawa (n = 5) and western
Ottawa (n = 42) at 0.5-, 1.0-, 1.5-, and 2.0-km radii. Median percentages were compared using the non-parametric Mann-
Whitney test (W). Numbers in bold indicate a significant difference between eastern and western occupied sites (P < 0.05).

Land Radius Eastern Ottawa Western Ottawa

use class (km) sites (median %) sites (median %) W Ie

Agriculture 0.5 67.4 45.0 954 0.0649
1.0 64.8 46.5 948 0.0401
Is 65.4 47.2 932 0.0092
2.0 69.0 48.1 922 0.0032

Forest 0.5 9.9 17.0 1061 0.0701
1.0 3b 18.1 1066 0.0473
5 6.0 DRG 1064 0.0555
2.0 4.7 7743) 1073 0.0261

Wetland 0.5 9.4 ll 1061 0.0701
1.0 il 18.6 1049 0.1623
IL 9.7 19.6 1067 0.0435
2.0 9.3 18.1 1083 0.0102

TABLE 2. Land use surrounding apparently unoccupied sites in eastern Ottawa (m = 179) and sites with Boreal Chorus Frogs
(Pseudacris maculata) in western Ottawa (n = 42) at 0.5-, 1.0-, 1.5-, and 2.0-km radii. Median percentages were compared
using the non-parametric Mann-Whitney test (W). Numbers in bold indicate a significant difference between apparently
unoccupied eastern and occupied western sites (P < 0.05).

Unoccupied sites Occupied sites

Land Radius in eastern Ottawa in western Ottawa

use class (km) (median %) (median %) W (ie

Agriculture 0.5 529) 45.0 20469 0.1079
1.0 54.5 46.5 20346 0.2014
1.5 S25) 47.2 20488 0.0972
2.0 50.8 48.1 20606 0.0483

Forest 0.5 14.0 17.0 19413 0.2219
1.0 Wc) 18.1 19507 0.3324
le) 17.0 21.6 19316 0.1385
2.0 7 2251 19203 0.0743

Wetland 0.5 11.1 7 18951 0.0139
1.0 13.0 18.6 18975 0.0166
1.5 14.3 19.6 18812 0.0046
ZA) 16.0 18.1 18790 0.0038

2014

TABLE 3. Land use Surrounding sites in eastern Ottawa with (7
(Pseudacris maculata) at 0.5-, 1.0-, 1.5-, and 2.0-km radii. Median percent

SEBURN £7 AL.: BOREAL CHORUS FROG DECLINE 155

~ 5) and apparently without (” = 179) Boreal Chorus Frogs

ages were compared using the non-parametric Mann-

Whitney test (IY). Numbers in bold indicate a significant difference between occupied and unoccupied sites (P < 0.05),

Land use Radius Occupied sites
class (km) (median %)
Agriculture 0.5 67.4

1.0 64.8

lS 65.4

2.0 69.0
Forest 0.5 ; 9.9

1.0 3

1.5 6.0

2.0 4.7
Wetland 0.5 9.4

1.0 11.2

les OR

2.0 Wh!

Unoccupied sites

(median %) W P
52.9 16448 0.3534
54.5 16407 0.2016
5275 16371 0.1133
50.8 16320 0.0436
14.0 16698 0.2333
WS 16720 0.1679
17.0 16711 0.1927
el 16742 0.1172
11.1 16655 0.4089
13.0 16628 0.5512
14.3 16669 0.3447
16.0 16745 0.1114

aees—s—sosaooS70—{—[sas=asas_—_——X—X—X—X

Discussion

Our surveys indicate that the Boreal Chorus Frog is
rare in eastern Ottawa, and a decline seems to have
occurred since the 1950s. Although our surveys un-
doubtedly failed to detect some populations, our survey
technique has proven successful in the past. Using this
technique, we found 30 previously undocumented Bor-
eal Chorus Frog locations in western Ottawa (Seburn
and Gunson 2011). One limitation of our survey tech-
nique is that auditory surveys were largely limited to
wetlands visible from roads, and Boreal Chorus Frog
populations may persist away from roads. However,
only about 27% of the study area is more than 500 m
from a road, the approximate distance at which a chorus
can be heard.

Six broad classes of threats have been identified as
causes of amphibian declines: commercial use (1.e., col-
lecting), introduced/exotic species, land use change,
contaminants, climate change, and infectious diseases
(Collins and Crump 2009). Commercial use can be
ruled out because the Boreal Chorus Frog is too small
to be easily collected by humans. It is also unlikely that
exotic species have caused the decline. Although intro-
duced plants such as Purple Loosestrife (Lythrum sali-
caria) and European Common Reed (Phragmites aus-
tralis ssp. australis) have altered wetlands, they are
common across all of eastern Ontario including areas
where the Boreal Chorus Frog remains widespread. The
metabolite emodin from the invasive European Buck-

thorn (Rhamnus cathartica) can cause mortality of

Boreal Chorus Frog embryos (Sacerdote and King, in
press), but this shrub appears common in both east-
ern and western Ottawa (Soper and Heimburger 1985).
It also seems unlikely that climate change would cause
Boreal Chorus Frogs to decline in eastern Ottawa but
not nearby in western Ottawa. This leaves three broad
classes of threats: contaminants, infectious disease, or
land use change.

Contaminants can be lethal to the Boreal Chorus
Frog. Exposure to low levels (700 ppb) of the glypho-
sate herbicide Roundup WeatherMax (Monsanto Cana-

da Inc., Winnipeg, Manitoba) resulted in 80% mortality
of Boreal Chorus Frog tadpoles (Williams and Seml-
itsch 2010). The high mortality rate is likely a result of
the unique surfactant in the product, as Roundup Origi-
nal (Monsanto Canada Inc.) did not cause high levels
of mortality (Williams and Semlitsch 2010). However,
Roundup WeatherMax was not available in Canada
until the early 2000s, and the Boreal Chorus Frog was
declining before that (e.g., Daigle 1997), so it likely
does not explain the declines. Nitrate contamination
of surface waters from nitrogen-based fertilizers can
also cause significant mortality of Western Chorus Frog
tadpoles at levels of 10 mg/L or more (Hecnar 1995),
but surface water data from the City of Ottawa suggest
that nitrate levels are consistently below 10 mg/L, al-
though it is possible they were higher in the past.

Infectious disease is another possibility. The chytrid
fungus Batrachochytrium dendrobatidis has been de-
tected in over 50 countries and over 500 species of
amphibians (Olson ef a/. 2013). Infection can lead to
chytridiomycosis, a fatal disease implicated in the de-
cline of over 200 amphibian species (Fisher e¢ a/. 2009).
The fungus has been detected in Boreal Chorus Frogs
in southern Quebec, but no increased mortality or mor-
bidity of frogs was observed (Ouellet ef a/. 2005). Thus,
although chytridiomycosis cannot be excluded, again it
is unlikely that it would affect frogs in eastern Ottawa
but not western Ottawa.

Given that land use change resulting in habitat
destruction is the leading cause of amphibian decline
(Collins and Crump 2009; Collins 2010), it seems plau-
sible that habitat loss is a prime driver of the decline
in the Boreal Chorus Frog. The fact that unoccupied
sites in eastern Ottawa had significantly lower wetland
cover values at all spatial scales compared with occu-
pied sites in western Ottawa (Table 2) suggests that
landscape differences could explain the apparent de-
cline; however, we have no direct evidence that wet-
land cover in eastern Ottawa has been reduced since
the 1950s. One problem with this hypothesis is that oc-
cupied sites in eastern Ottawa did not differ in wet-

156

land cover from apparently unoccupied sites in eastern
Ottawa (Table 3). Local conditions, such as hydrology,
wetland configuration or connectivity, and the number
of wetlands too small to be detected by our analyses,
could explain the localized persistence of Boreal Cho-
rus Frogs at occupied sites in eastern Ottawa. Wetland
cover in all of Ottawa-Carleton declined from an esti-
mated 47.2% circa 1800 to an estimated 13.2% in 1967,
but no significant change in wetland cover occurred
from 1967 to 2002 (Ducks Unlimited 2010*). Intensive
farming practices in the 1950s and 1960s likely result-
ed in wetland loss, although we cannot document the
extent of any loss.

The Boreal Chorus Frog is a short-lived frog, with
more than 90% of adults breeding only once (Whit-
ing 2004). Individuals typically remain within 275 m
of breeding ponds (Desroches ef al. 2001*) and wet-
land occupancy is correlated with the number of neigh-
bouring occupied wetlands (Scherer ef a/. 2012). In the
closely related Western Chorus Frog, stochastic events,
such as premature drying of breeding ponds, can lead
to annual extinction rates exceeding 50% (Werner ef al.
2009). In addition, dry summers may limit successful
immigration of Boreal Chorus Frogs to unoccupied
wetlands (Schueler and Karstad 2013*). These factors
suggest that wetland connectivity is important in ensur-
ing that re-colonization can occur from neighbouring
populations. Such lack of connectivity appears to be a
factor in western Ottawa where isolated populations are
more apt to become extirpated (Seburn and Gunson
2011).

Boreal Chorus Frogs likely persist across the land-
scape as a metapopulation (e.g., Gill 1978). If reduced
wetland cover results in fewer wetlands, then it is plau-
sible that wetland connectivity is reduced in eastern
Ottawa. For example, a simulation of the loss of small
wetlands in Maine demonstrated that this would result
in a loss of total wetland area of 19%, but increase the
inter-wetland distance by 67% (Gibbs 1993). Whether
wetland connectivity has been reduced in eastern Ot-
tawa cannot be explicitly tested using SOLRIS data,
as wetlands smaller than 0.5 ha cannot be identified
(OMNR 2007*). A detailed examination of wetland dis-
tribution and abundance in eastern and western Ottawa
is required to determine whether wetland connectivity
could explain the apparent decline of the Boreal Cho-
rus Frog.

The most similar species that co-occurs with the
Boreal Chorus Frog 1s the congeneric Spring Peeper
(Pseudacris crucifer), which remains widespread in
eastern Ottawa (DS, personal observation). Why should
one species decline and the other remain widespread?
In general, the closely related Western Chorus Frog is
more common in ephemeral sites that tend to lack
aquatic predators, whereas the Spring Peeper is more
common in less ephemeral sites with more abundant
aquatic predators (Skelly 1996). This also appears to be
the case for the Boreal Chorus Frog in eastern Ontario
(DS, personal observation). Small, ephemeral wetlands

THE CANADIAN FIELD-NATURALIST

Vol. 128

are often first lost when cropland expands and the result
could be a landscape more suited to Spring Peepers
than Boreal Chorus Frogs.

There is compelling evidence that the Boreal Cho-
rus Frog has declined across much of eastern Ottawa
and western Prescott and Russell. The lack of wetland
cover in eastern Ottawa may have contributed to this
decline, although additional research on abundance and
distribution of small wetlands is required. Finding ex-
planations for the widespread persistence of western
Ottawa populations may also provide guidance for the
conservation of this species.

Acknowledgements

Mike Oldham of the Natural Heritage Information
Centre of the Ontario Ministry of Natural Resources
graciously provided data from the Ontario Herpeto-
faunal Atlas. We thank Garnet Hanes for the data and
maps from the surveys he conducted with the late
Anne Hanes. Funding was provided by the Canadian
Wildlife Federation. This paper benefited from review
comments from Carolyn Seburn and four anonymous
reviewers.

Documents Cited (marked * in text)

COSEWIC (Committee on the Status of Endangered
Wildlife in Canada). 2008. COSEWIC assessment and
update status report on the Western Chorus Frog Pseudacris
triseriata Carolinian population and Great Lakes/St. Law-
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Desroches, J. F., D. Pouliot, and S. Coté. 2001. Evaluation
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Ducks Unlimited. 2010. Southern Ontario wetland conver-
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Schueler, F. W., and A. Karstad. 2013. Preliminary results:
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Received | November 2013
Accepted 15 December 2013

Activity and Diet of Bats in Conventional versus Organic Apple
Orchards in Southern Michigan

BRENNA L. Lonc! AND ALLEN Kurta!+?

'‘Department of Biology, Eastern Michigan University, Ypsilanti, Michigan 48197 USA
Corresponding author: akurta@emich.edu

Long, Brenna L., and Allen Kurta. 2014. Activity and diet of bats in conventional versus organic apple orchards in southern
Michigan. Canadian Field-Naturalist 128(2): 158-164.

Organic farming practices have been suggested as a conservation strategy that can provide productive foraging sites for
insectivorous bats in agricultural areas. We tested the hypothesis that the number and diversity of insects captured and the
resulting activity and diet of bats would differ between organic and conventional apple orchards. During the summer of 2009
and 2010, we captured 131 Big Brown Bats (Eptesicus fuscus) and five Eastern Red Bats (Lasiurus borealis) by mist netting in
four organic and four conventional orchards in Michigan. Acoustic monitoring revealed that most calls (59%) were produced
by Big Brown Bats, followed by Hoary Bats (L. cinereus), Eastern Red Bats, and unidentified species of Myotis. Most insects
(96%) captured in light traps were Coleoptera, Diptera, Hemiptera, Lepidoptera, and Trichoptera; fecal analysis indicated
that Coleoptera dominated the diet of Big Brown Bats (79% by volume), followed by Diptera (14%). There were no Si gnificant
differences between organic and conventional orchards in number of insects captured with light traps, bats captured with mist
nets, acoustic recordings of bats, or proportion of Coleoptera and Diptera consumed by the bats. However, the taxonomic compo-
sition of captured insects and acoustic recordings was not homogenous among orchards in either group, suggesting that factors
other than farming practices were more important in determining which insects and bats were found in the small orchards
typical of southern Michigan.

Key Words: Big Brown Bat; Eptesicus fuscus; Eastern Red Bat; Lasiurus borealis; Hoary Bat; Lasiurus cinereus, bat activity:
bat diet; apple orchard; organic farming; pesticides; Michigan

Introduction

Use of synthetic pesticides in conventional agricul-
ture not only causes health concerns, but also incurs
social and environmental costs (Cleveland et a/. 2006).
Although pesticides are helpful in controlling injurious
species, such compounds can have a negative impact
on beneficial organisms (Morales 2002). In addition,
regular application of synthetic pesticides can result in
a community of insects more resistant to specific chem-
icals (Turgut ef a/. 2011) and a need to develop new
substances or apply more of the old, thus increasing
production costs (do Amarante ef a/. 2008). A growing
concern of both consumers and farmers over the use of
pesticides has led some growers to move from conven-
tional to organic practices (Meisterling ef al. 2009;
Turgut et a/. 2011). Organic management excludes the
use of synthetic pesticides and fertilizers, fungicidal
and insecticidal soaps, botanical insecticides, and sulfur
and copper compounds (Holb ef al. 2003; Peck et al.
2006).

Differences in farming practices between organic and
conventional farms should lead to differences in their
communities of insects in terms of number of individu-
als, taxonomic composition, or both (Letourneau and
Goldstein 2001; Bengtsson ef a/. 2005; Minarro et al.
2009; Padmavathy and Poyyamoli 2011; Mates ef al.,
2012). These presumed differences in insect communi-
ties should be reflected in the level of activity and diet
of bats, the primary nocturnal predators on insects
(Genghini et a/. 2006; Braun de Torres 2011). In the
United Kingdom, for example, Wickramasinghe ef al.

(2003) claimed that organic farms provided islands of
high-quality foraging habitat for insectivorous bats
and proposed organic practices as a potential tool for
increasing the numbers and kinds of bats in agricul-
tural areas.

In this study, we tested the hypothesis that organic
procedures affect the number and diversity of bats and
insects in orchards that grow apples, an economically
important crop in Canada and the United States (On-
tario Ministry of Agriculture and Food 2007*; USDA
2012*). We predicted that the abundance and diversi-
ty of both bats and insects would be greater at organic
sites. Based on the presumed greater abundance of in-
sects on organic farms, we also predicted that reproduc-
tively active adult female bats, which must bear the
high energetic cost of pregnancy and lactation (Kurta
et al. 1989, 1990), would be more common in organic
orchards, relative to adult males and non-reproductive
females, than at conventional sites. Finally, many spe-
cies of bats are selective opportunists, generally con-
centrating on certain types of prey but often increasing
consumption of locally abundant taxa (Fenton and Mor-
ris 1976; Murray and Kurta 2002). Consequently, we
predicted that the diet of bats would vary between
organic and conventional sites, reflecting the assumed
differences in composition of the insect community.

Study Area

Our study took place in four organic and four con-
ventional apple orchards in southern Michigan, within
200 km of Jackson (41°51' to 43°1'48"N, 83°40'12" to

158

2014

84°S1'36"W; Smith 2012). All orchards were in rural,
agricultural areas, with corn and soy as the most com-
mon crops. The apple trees in the eight orchards were
either small dwarf varieties (1—3 m in height) or semi-
dwarf trees (3—5 m in height). Orchards varied in size
from 5.7 to 24.3 ha, with a mean of 13.2 ha. Average
distance between an orchard and the closest other
orchard used in the study was 112 km + 10 (SE). All
fieldwork took place between 5 June and 13 August
2009 and between 18 May and 12 August 2010.

Methods

Field and laboratory techniques

To sample insects, we used a light trap (Model
2815A, BioQuip, Gardena, California), powered by a
12-volt battery. Light trapping occurred at only one
orchard on any given night and coincided with mist
netting for bats (see below). The trap was turned on | h
before sunset and deactivated about 5 h later, imme-
diately after the mist-netting system was dismantled.
Captured insects were preserved in 70% ethanol for
later counting and identification to order with the aid
of a dissecting microscope.

Bats were captured using mist nets made from 50-
denier, braided nylon (Avinet, Dryden, New York).
Netting systems were 6—9 m high and 6-12 m wide,
depending on the size of available flight corridors. Mist
netting was typically carried out 3-5 times a week,
depending on weather, but in only one orchard on any
given night. Netting began at sundown and continued
for 4 h. Ambient temperature was recorded at the begin-
ning and end of mist netting. Each captured bat was
identified to species, sexed, and classified as adult or
juvenile, depending on fusion of the epiphyseal plates
at the metacarpal-phalangeal joints (Anthony 1988).
Females were further identified as non-reproductive,
pregnant, lactating, or post-lactating based on body
mass, abdominal palpation, and ability to express milk
from the nipples (Racey 1988).

Fecal samples were collected from each captured
bat by placing the animal in a paper lunch bag until
defecation occurred. Fecal pellets were frozen for later
examination, when each pellet was soaked in 95%
ethanol and teased apart under a dissecting microscope.
The resulting insect fragments were identified to order,
and the percent volume of each order was estimated
visually for each pellet (Whitaker 1988). .

To complement mist netting, we used ultrasonic
detectors to assess activity of bats. Each monitoring
unit consisted of an ultrasonic detector (Anabat I, Tit-
ley Electronics, Ballina, Australia), a zero-crossings
analysis interface module (ZCAIM), and a small bat-
tery (12-volt, 5 A-h) housed in a protective case. A 5-cm
diameter opening was cut into the side of the case, and
a piece of polyvinylchloride pipe in the shape ofa flat
U was attached (Smith 2012). When 1n use, the micro-
phone of the detector rested inside the inner opening
of the pipe. The detector sensed ultrasound produced

LONG AND Kurta: BATS IN CONVENTIONAL VERSUS ORGANIC ORCHARDS

159

by each bat, whereas the ZCAIM transformed the
sound into a digital signal and recorded the frequency-
versus-time structure of the call as a file on a compact
flash card inside the ZCAIM. Detectors were calibrat-
ed against each other before use (Larson and Hayes
2000) with an ultrasonic distance-measuring device
(Model 10065, Sonin, Charlotte, North Carolina).

Eight monitoring units were established, one at each
orchard. Each unit was programmed to record from
about 30 minutes before local sunset to about 30 min-
utes after local sunrise. During recording, the cases were
positioned on a pole, about 6 m above the ground. After
downloading files each week, we moved the unit to a
new, randomly selected site in each orchard.

The number of files recorded per night was used as
a measure of bat activity. If a particular monitoring unit
did not function for the entire night, then all results
from that night were discarded. Recorded calls were
examined using Analook W software (version 3.3q,
2006, http://hoarybat.com) and assigned to one of four
groups: Myotis spp., Eastern Red Bat (Lasiurus bore-
alis), Hoary Bat (Lasiurus cinereus), or Big Brown
Bat (Eptesicus fuscus) (Flaquer et al. 2007; Brooks
2009; Morris et al. 2010). Although a few calls cate-
gorized as Big Brown Bat may have been made by
Silver-haired Bats (Lasionycteris noctivagans), the
latter species is rare, representing less than 0.1% of
bats that are mist netted in southern lower Michigan
in summer (Winhold and Kurta 2008; Kurta 2010). No
attempt was made to categorize sequences of calls with
fewer than three individual pulses or consisting of puls-
es of low quality (Johnson ef al. 2002; Francl 2008;
Johnson and Gates 2008).

Statistical analyses

We focused our analysis on Big Brown Bats because
they were the predominant species in the area. To
determine differences in diet between the two types
of orchards, we used two-sample independent /-tests.
Chi-squared tests were used to examine differences
in the composition of insects (by order) captured and
differences in calls of bats recorded at the two types of
orchards. In addition, chi-squared tests were used to
analyze differences between the two types of orchards
in terms of the proportion of reproductive versus non-
reproductive female Big Brown Bats and male versus
female adult Big Brown Bats. Both types of tests were
conducted using VassarStats (Lowry 2008*).

A nested analysis of variance was used to detect dif-
ferences between organic and conventional orchards
in the number of acoustic records, bats netted, and in-
sects captured. Only orders of insects that contributed
at least 5% of the total amount captured were includ-
ed in the comparison. Minimum ambient temperature
was used as a covariate in these three analyses, because
temperature can affect the number and diversity of in-
sects that are active (e.g., see Ciechanowski et al. 2007;
Le Lann et al. 2011). To obtain normally distributed
data, we added 1 to the raw counts obtained from

160 THE CANADIAN FIELD-NATURALIST Vol. 128

acoustic monitoring, mist netting, and light trapping
and applied a logarithmic transformation before per-
forming statistical analyses. Analyses were performed
primarily with Systat 10.2 (Systat Software, Inc., Rich-
mond, California) or Excel (Microsoft, Redmond,
Washington).

A nested analysis of variance was also used to com-
pare the percent volume of various orders of insects
found in the diet of bats from organic and convention-
al orchards. The percent volume represented by each
insect order in the fecal pellets produced by each bat
was converted to a proportion, and the square root of
each proportion was arcsine-transformed to attain nor-
mal distribution of the data (Zar 1999). Only orders
that contributed at least 5% of the total volume were
included in these comparisons.

Different practices in conventional and organic
orchards could affect not only the overall level of activ-
ity of bats and insects, but also the structure of the local
communities (Bengtsson ef al. 2005; Macfadyen et al.
2009; Power and Stout 2011). Consequently, we also
calculated taxonomic diversity, using the level of
species or genus for the bats and order for the insects
that were captured or identified in the feces. As a meas-
ure of diversity, we used Simpson’s index, which is
equal to 1 —[}' n(n, — 1)] / N (N— 1), where n, is the
number of individuals in each taxonomic group and NV
is the total number of captured species or orders (Brow-
er and Zar 1984). We calculated diversity for each
orchard and compared mean diversity of organic and
conventional orchards using a standard ¢-test. All data
are presented as the mean and standard error (SE), and
alpha was set to 0.05 for all statistical tests.

Results

Over the 2 years of our study, we trapped insects on
36 nights in organic sites and 38 nights in conventional
orchards for a total of 5233 insects (138 + 22 insects/
night) from the organic orchards and 4179 insects (116
+ 20 insects/night) from conventional orchards. There

was no significant difference between the two types of
orchards in number of insects trapped per night (/, ,
= ().03, P = 0.87). Five orders of insects — Coleoptera
(beetles), Diptera (flies), Hemiptera (true bugs), Lep-
idoptera (moths), and Trichoptera (caddisflies) —
accounted for 96% of the insects captured. There was
no significant difference between the two types of
orchards in numbers of insects by order captured per
night (all F, , < 0.54, P = 0.49; Table 1). The mean
diversity at the ordinal level was also not significantly
different (t,, = 1.95; P= 0.13): organic locations (0.71

+ 0.03) and conventional orchards (0.76 + 0.00).

Beetles were the most abundant order of insects in
both types of orchard, representing 40% and 31% of
captures in organic and conventional orchards, respec-
tively (Table 1). The next most common order at organ-
ic sites was moths (21%), but at conventional orchards
flies were the second most abundant order (23%). A
preliminary chi-squared test indicated significant het-
erogeneity in the composition of the five most com-
mon orders of insects among both the organic (7°, =
293.57; P < 0.001) and conventional (y*,, = 615.74;
P < 0.001) orchards, which precluded a statistically
valid comparison of the overall composition of insects
between types of orchards using chi squared.

Mist netting was carried out on 37 nights in organ-
ic and 38 nights in conventional apple orchards and
yielded 136 bats. Of these bats, 54 (39.7%) were net-
ted in organic orchards (1.5 + 0.4 captures/night), and
82 (60.3%) in conventional orchards (2.2 + 0.6 cap-
tures/night). The difference in number of captures per
night between organic and conventional orchards was
not significant (F, , = 0.26, P = 0.63).

We caught 131 Big Brown Bats and 5 Eastern Red
Bats — the two most common species in the region
(Winhold and Kurta 2008; Smith 2012) — and we
focus our analyses on the more-abundant Big Brown
Bat. They consisted of 44 adult females, 27 juvenile
females, 32 adult males, and 28 juvenile males. Of the
adult females, 14 reproductive (six pregnant, three lac-

TABLE |. Average number (%) of insects of the five most abundant orders captured per night at each orchard in southern Michigan.
Orders contributing less than 5% of the total number of insects trapped in both types of orchard were not included (Smith,

2012); thus, row percentages do not total 100%.

Orchard Lepidoptera Coleoptera Diptera Hemiptera Trichoptera Total
Orchard
AlMar E 3.6 (17.4) 8.9 (42.7) pron (2522) 0.5 2.4) eRe (lee) 20.8 (99.8)
AlMar SE 6.1 (18.7) 11.8 (36.0) 6.2 (19.0) 4.3 (13.2) 4.1 (12.7) 32.5 (99.6)
Apple Schram 11.4 (32.1) 15.2 (42.7) 3.8 (10.8) EOIGeS) 2.9 (8.3) 35.2 (99.4)
Country Mill 3.4 (12.8) 10.9 (40.6) SESH (ORG) 2.8 (10.5) 4.3 (15.9) 26.7 (99.4)
Total 24.6 (21.2) 46.8 (40.3) 20.6 (17.7) 9.6 (8.3) 13.9 (12.0) 115.5 (99.5)
Conventional
Erwin 5.1 (18.4) 9.8 (35.6) 4.4 (15.9) 1.3 (4.6) 6.8 (24.6) 27.4 (99.1)
Spicer 625,38) S823) 3.8 (20.7) 1.9 (10.6) 1.9 (10.5) 18.1 (99.4)
Swinderman N Seo.) 18.5 (27.9) 19.6 (29.5) Pera eell)) 20.2 (30.6) 64.3 (97.1)
Swinderman S PEAS) UES) 8.9 (34.8) 4.1 (16.1) 1.2 (4.7) 1.2 (4.7) 18.3 (71.6)
Total 15.9 (11.6) 43.1 (31.3) Blesn(2Selh) WN (2) 30.1 (21.8) 128.0 (93 0)
Overall average 20.1 (15.9) 44.9 (35.3) 26.4 (20.7) 8.3 (6.5) 22) 121.9 (95.9)

2014 LONG AND Kurta: BATS IN CONVENTIONAL VERSUS ORGANIC ORCHARDS 16]

tating, and five post-lactating) and 3 non-reproductive by Big Brown Bats (59%), followed by Hoary Bats
bats were netted in organic orchards and 21 reproduc- — (28%), Eastern Red Bats (13%), and Myotis spp. (< 1%;
tive (4 pregnant, 13 lactating, and 4 post-lactating) and — Table 2). The mean diversity of bats in organic orchards
6 non-reproductive bats were netted in conventional (0.53 + 0.02) was not statistically different (¢, = 0.78;
orchards. The difference in proportion of reproductive — P = 0.23) from that in conventional orchards (0.57 4
and non-reproductive female Big Brown Bats between 0.05). A preliminary chi-squared test indicated consid-
organic and conventional orchards was not signifi- erable heterogeneity in taxonomic composition of the
cant (7°, = 0.0; P= 1). The ratios of male to female calls of bats in both organic (y*, = 109; P < 0.001) and
adults (15:17 in organic orchards and 17:27 inconven- conventional (7. = 424: P < (0.001) orchards, which
tional orchards) were also not significantly different prevented a statistically meaningful comparison of

Qc, = 0.23; P = 0.63). overall composition between the two types of orchards
Acoustic monitoring records for the combined field — using chi squared.
seasons consisted of 25 945 files containing sounds We examined 131 fecal pellets from Big Brown Bats

made by bats during 569 detector-nights; a detector- and found that, overall, their diet included five orders
night was defined as a complete night of recording, of insects: Coleoptera (79.3% by volume), Diptera
from before sunset to after sunrise, by one detector. Of (13.2%), Lepidoptera (0.6%), Hemiptera (0.7%), and
these files, 10 217 were recorded on 279 detector-nights | Hymenoptera (0.7%; Table 3). Mean diversity of the
in the four organic orchards, whereas 15 728 were re- diet at the ordinal level was not significantly differ-
corded on 329 detector-nights in the four convention- — ent (¢, = 0.95; P = 0.19) between organic locations
al orchards. Although more files per night were record- (0.30 + 0.04) and conventional orchards (0.36 + 0.05).
ed in conventional orchards (51 + 14 files/night) than There were no significant differences between organic
in organic orchards (37 + 8 files/night), the difference and conventional orchards in percentage volume of the
was not significant (F, , = 0.20, P = 0.67). two most common orders: Coleoptera (F , = 0.21, P
We assigned 11 045 (43%) of the recordings to genus = 0.66) and Diptera (F, , = 0.01, P= 0.92).
or species. Most calls from all orchards were produced

TABLE 2. Numbers (and %) of bats identified from recordings by ultrasonic detectors in four organic and four conventional
apple orchards in southern Michigan.

eee SS
ooo OO <<<

Species Organic orchard Conventional orchard Total
Big Brown Bat 2943 (63.7) 3554 (55.3) 6497 (58.8)
Hoary Bat 1030 (22.3) 2030 (31.6) 3060 (27.7)
Eastern Red Bat 631 (13.7) 826 (12.9) 1457 (13.2)
Myotis spp. 15 (0.3) 16 (0.3) 31 (0.3)
Total 4619 6426 11 045

eS 00s ay —>woOO

Note: Big Brown Bat, Eptesicus fuscus; Hoary Bat, Lasiurus cinereus; Eastern Red Bat, L. borealis.

TABLE 3. Percent volume of insects, by order, consumed by Big Brown Bats, Eptesicus fuscus, in four organic and four con-
ventional apple orchards in southern Michigan based on fecal peliet analysis. Columns may not add to 100 because of

rounding errors.

Volume consumed (%)

Organic orchard Conventional orchard Total
Insect order (n = 52 bats) (n= 79 bats) (n= 131 bats)
Coleoptera 133 78.0 78.6
Diptera 132 13.9 13%
Lepidoptera 0.6 1.3 1.0
Hemiptera 0.7 lel 0.9
Hymenoptera 0.7 0.2 0.5
Unknown Sy) 4.5 5.0

SS SSeS —m—w[wyw]{Tr omwwee eee

Discussion because they were predicated on an assumed difference

We predicted that adult female bats would be more In the insect community between organic and conven-
common in organic orchards than conventional ones, tional orchards. However, light trapping revealed that
but the sex ratio of adult Big Brown Bats did not dif- __ total abundance and diversity of insects, as well as the
fer between types of orchard, nor did the ratio of repro- abundance of the five most common orders of insects,
ductive to non-reproductive adult females. Although did not differ between types of orchards (Table 1).

our predictions were not upheld, this is not surprising Availability of prey at the ordinal level, therefore, was

162

similar in organic and conventional apple orchards, sug-
gesting no reason for energetically stressed females to
favour one type of orchard over the other.

To date, only one quantitative study has been pub-
lished concerning activity of bats on organic and con-
ventional farms of any type (Wickramasinghe ef al.
2003). The authors compared bat activity on organic
and conventional farms in England and Wales, using
acoustic monitoring and captures via mist net, as we
did in Michigan. However, they reported that bat ac-
tivity was 61% greater on organic than conventional
farms. Also, in a companion study, Wickramasinghe
et al. (2004) detected a significantly greater number
of insects belonging to five families at organic loca-
tions, although pairwise statistical comparisons of 13
other families showed no differences between organic
and conventional sites.

However, the farms in the studies by Wickramas-
inghe ef al. (2003, 2004) differed greatly from ours.
Fifty-four percent of the farms in Wickramasinghe ef
al. (2004) were classified as livestock operations and
another 41% were mixed, involving both livestock and
unspecified crops. The farming practices and poten-
tial chemicals used in pastures and orchards differed.
For example, avermectin, an antihelminthic drug often
used with cattle and sheep on conventional farms, can
reduce the number of invertebrates using ungulate
feces, 1.e., dung beetles (Scarabaeidae and Geotrupi-
dae; Strong 1992), which are commonly eaten by
some British bats (Wickramasinghe ef al. 2004). Fur-
thermore, Wickramasinghe et a/. (2003, 2004) sampled
bats and insects in various habitats, and most differ-
ences they reported involved pastoral and aquatic habi-
tats; these habitats generally were not present in the
orchards we studied (Smith 2012). Finally, the average
size of their farms (about 51 ha; Wickramasinghe e¢ al.
2003) was more than double the area of our largest
orchard (24 ha).

Wickramasinghe ef al. (2003) proposed organic prac-
tices as a potential conservation strategy for increasing
the numbers and kinds of bats in agricultural regions.
However, we were unable to document differences in
the number, richness, and diversity of bats; in the
dietary composition of the bats; in overall activity and
diversity of insects at the ordinal level; or in the abun-
dance of particular orders of insects. The fact that tax-
onomic composition of the captured insects and the
acoustic recordings of bats were not homogenous a-
mong either conventional or organic orchards suggests
that factors other than farming practices, such as dif-
fering patterns of land use and habitats in the surround-
ing landscape, were more important in determining the
types of insects and bats found in apple orchards typ-
ical of southern Michigan.

Apple orchards in other parts of North America,
though, can be as large as 1200 ha (Washington Apple
Commission 2010*), and a suggested area of further
study is to examine the relation between size of orchard
and potential differences in their associated commu-

THE CANADIAN FIELD-NATURALIST

Vol. 128

nities of insects and bats. Large size might insulate the
interior of organic orchards from surrounding conven-
tional farms and potentially allow development of dis-
parate communities, as found by Wickramasinghe e/
al. (2003, 2004). In addition, recent advances in molec-
ular techniques, such as high-throughput sequencing
(Bohmann ef a/. 2011; Razgour et al. 2011), now allow
biologists to sequence the DNA of every item of prey
in the fecal material produced by a bat. Although many
sequences obtained in this manner cannot be identi-
fied to species or other taxonomic level below order,
use of molecular operational taxonomic units offers a
promising way of comparing diversity of prey in dif-
ferent types of orchards at a more detailed level than
visual analysis (Chesters et a/. 2013; Rolfe et al. in
press). Further research, concerning organic practices
and bats is warranted, because bats are experiencing
unprecedented mortality in North America from wind
turbines (Kunz et al. 2007) and white-nose syndrome
(Turner ef al. 2011), and determining factors that
promote the conservation of these flying mammals is
essential to their long-term survival.

Acknowledgements

We thank the farmers and their assistants for pro-
viding access to their orchards: J. Bush (Apple Schram
Orchard), B. Erwin (Erwin Orchard), J. Koan (AlMar
Orchard), A. Spicer (Spicer Orchard), J. Swinderman
(Apple Wood Orchard), and S. Tennes (Country Mill
Orchard). S. Francoeur and C. Rockey provided
advice on statistics and acoustic monitoring, respec-
tively. We thank Mckinley Ophelia for keeping BLL
company on many long nights in the orchards when
everyone else was asleep. A. Rolfe and J. Veilleux pro-
vided helpful comments on the manuscript. Financial
support was provided by a grant from the Organic
Farming Research Foundation to S. Tennes; a Faculty
Research Fellowship from Eastern Michigan Univer-
sity awarded to AK; and a Hellwig Research Award
given to BLL by the Department of Biology at Eastern
Michigan University.

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Received 8 November 2013
Accepted 28 January 2014

Decline in Breeding of the Great Black-backed Gull, Larus marinus,

ie = Herring Gull, LZ. argentatus, on Boot Island, Nova Scotia, 1986
to

COLIN M. MACKINNON!:? and ANDREW C. KENNEDY!

Environment Canada, Canadian Wildlife Service, Atlantic Region, P.O. Box 6227, Sackville, New Brunswick E4L 1G6
Canada

Van! . <a a

“Corresponding author: Colin.MacKinnon@ec.ge.ca

MacKinnon, Colin M., and Andrew C. Kennedy. 2014. Decline in breeding of the Great Black-backed Gull, Larus marinus,

and the Herring Gull, ZL. argentatus, on Boot Island, Nova Scotia, 1986 to 2010. Canadian Field-Naturalist 128(2):
165-172.

For over 50 years, Boot Island, Nova Scotia, has supported a significant mixed bird colony: Great Black-backed Gull (Larus
marinus), Herring Gull (L. argentatus), Great Blue Heron (Ardea herodias), and Double-crested Cormorant (Phalacrocorax
auritus). In 2002, the largest Great Black-backed Gull colony in Canada was located there. Over the last quarter century, the
Herring Gull colony has shown a dramatic and near-linear decrease from 727 nests in 1986 to 67 in 2000; in 2010, only two
nests remained. The number of Great Black-backed Gull nests has also declined by 44%, from 1467 nests in 1992 to 819 in
2010. These reductions may be partly attributed to factors external to the colony, such as changes in regional fisheries and
better landfill management. However, a more immediate problem may be nest predation and disturbance by American Mink
(Neovison vison), Raccoon (Procyon lotor), Coyote (Canis latrans), and Bald Eagle (Haliaeetus leucocephalus).

Key Words: Great Black-backed Gull; Larus marinus; Herring Gull; Larus argentatus; Boot Island; National Wildlife Area;

Nova Scotia; nesting; population decline

Introduction

Following the indiscriminate slaughter of many sea-
birds in the late 19th and early 20th century, often for
the millinery trade, Great Black-backed and Herring
Gulls recovered with a dramatic increase in numbers,
resulting in great overlap in their breeding ranges (Drury
1973). In the first half of the 20" century, Herring Gulls
were by far the more numerous (Drury 1973). How-
ever, after reaching a peak in the 1970s and 1980s, their
numbers along the east coast of North America start-
ed to decline, while Great Black-backed Gull numbers
continued to increase (Brown ef a/. 2001 in Rome and
Ellis 2004; Shoals Marine Laboratory 2010*). Cur-
rently, in eastern Canada, the Great Black-backed Gull
is deemed “not currently at risk” while Herring Gull
populations are of “moderate” concern as there is an
apparent population decline (Milko et al. 2003*).

These colonial birds require remote, disturbance-free
habitat in which to nest. The Great Black-backed Gull,
Larus marinus, and the Herring Gull, L. argentatus,
are ground-nesting birds that are highly susceptible to
mammalian predation and, thus, seek out isolated breed-
ing locations (Pierotti and Good 1994). Such colonies
are frequently situated on remote islands where moni-
toring by counting the number of nests is often difficult
because of poor and limited site access and a narrow
survey window. .

Surveys of Great Black-backed and Herring Gull
colonies are conducted regularly in eastern Canada.
However, when time and resources are limited, a trade-
off is made between monitoring a few colonies inten-
sively and conducting infrequent surveys of many col-

onies over a wide geographic area. Broad-scale surveys,
frequently conducted from the air, predominate as the
desired method of monitoring populations at a region-
al scale. However, aerial counts are not as accurate as
ground surveys (Drury 1973; Boyne and Hudson
2002*) and are often of limited value in tracking pop-
ulation trends of individual colonies.

Gull colonies in Boot Island National Wildlife Area
(Kings County, Nova Scotia) present a unique monitor-
ing opportunity for eastern Canada because biennial
surveys of colonial nesting birds have been undertaken
as part of the National Wildlife Area’s management
regime since 1984. In this article, we assess changes in
the size of both Great Black-backed and Herring Gull
colonies from 1986 to 2010 and discuss possible fac-
tors affecting the multiyear decline in both species.

Study Area

Boot Island National Wildlife Area is located in the
southern bight of Minas Basin at the mouth of the
Gaspereau River, 8 km northeast of Wolfville, Kings
County, Nova Scotia (45°08'N, 64°16'W). The 107-ha
wildlife area consists of approximately 91 ha of salt
marsh and 16 ha of uplands (Figure 1). The uplands
comprise the 15.5-ha main island (2.0 ha forested and
13.5 ha of open habitat) as well as a 0.5-ha treeless and
much smaller “Cyril’s Island” (Little Boot Island). The
upland portion of Boot Island slopes gently upward
from the salt marsh in a northeast direction, such that
the seaward face consists of an actively eroding 6-m
high mud and sandstone cliff. The small forested area is
dominated by a dense stand of dead and dying White

165

166
N
Uplands
ot Boot Island
Saltmars National Wildlife
—, Area

Long Island

Minas
Basin

FIGURE |. Location of Boot Island National Wildlife Area,
Kings County, Nova Scotia.

Spruce (Picea glauca) with scattered Red Maple (Acer
rubrum), Trembling Aspen (Populus tremuloides), and
Balsam Fir (Abies balsamea). Decay of the forest has
been exacerbated by the deleterious effects of tree-
nesting Great Blue Herons (Ardea herodias) (see
MacKinnon and Kennedy 2006*).

The area of open field habitat, where most Great
Black-backed and Herring Gull nests are located, is
dominated by two species of wild mustard: Wild Radish
(Raphanus raphanistrum) and Hairypod Hedgemus-
tard (Sisymbrium officinale). The remaining area is
composed of open field and rough cover consisting of
a nearly impenetrable thicket of Virginia Rose (Rosa
virginiana), American Red Raspberry (Rubus idaeus)
and Northern Bayberry (Myrica pensylvanica) (Newell
et al. 2006*).

Historically, Boot Island supported large colonies
of Great Black-backed and Herring Gulls. The island
also supports a large colony of Great Blue Herons and
Double-crested Cormorants (Phalacrocorax auritus),
and once supported a significant winter roost of Amer-
ican Crows (Corvus brachyrhynchos) (Milton 1983),
However, around 1991 the crows relocated 19.5 km to
the southwest to the town of Kentville on the mainland.
The inter-tidal mudflats in the vicinity of the island are
recognized as a RAMSAR site (a wetland of interna-
tional importance) and support a diversity of migrant
shorebirds (Elliot 1977; Hicklin 1981, 1984: Mawhin-
ney 1991*; Gilliland 1992). These flats also sustain a
commercial bait (Bloodworm, Glycera dibranchiata)
industry (Klawe and Dickie 1957*).

Until the late 1800s, Boot Island was connected to
the mainland and it has a long history of human use
(Mitcham 1986). The island was farmed intensively,
and an extensive system of dykes and ditches draining
the adjacent salt marshes existed until agricultural
activities ceased in approximately the 1930s.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Over many decades, biologists and naturalists have
visited Boot Island and reported wildlife observations.
The late Cyril Coldwell, curator of Acadia University’s
ornithology collection, provided most colonial bird
observations throughout the 1960s. The first detailed
survey of colonial birds on Boot Island was carried out
by Barkhouse (1976*), who also summarized past ob-
servations, reporting that gulls started nesting on the
island around 1948 followed shortly after by breed-
ing Great Blue Herons. Double-crested Cormorants
were first reported nesting in 1967.

Methods

Between 1976 and 2010, 14 surveys of gull nests
were completed, recording both occupied and unoc-
cupied nests. All surveys were centred at the peak or
the end of the Great Black-backed Gull’s egg-laying
period in mid-May (survey dates ranged from 8 to 19
May, average 13 May). Although Herring Gulls lay
slightly later than Great Black-backed Gulls, full (three-
egg) clutches of the former were frequently encoun-
tered, and thus we believe that this single survey win-
dow adequately captured the number of nests occupied
by both species. Slight variations in survey date were
due to weather and travel conditions, as the tidal waters
between Boot Island and the mainland are extremely
hazardous. No surveys were conducted during cold or
wet conditions (see Diamond 2001*).

Following the census method described by Bark-
house (1976*), surveys were carried out by walking a
series of parallel transects across the narrowest width
of the colony. Observers walked abreast and within
close proximity to each other. The outermost observers
temporarily marked the flanks of each transect with red
flagging tape. On completion of a transect, the group
wheeled around in unison so that on the return tran-
sect, the person who had been on the outermost flank
followed his or her previously marked line. One person
in the centre, usually the crew leader, was the recorder.
The crew leader set the transect width, based on observ-
er visibility and nesting density and recorded and con-
firmed observations relayed from the rest of the team.
Observations reported orally by crew members were
repeated by the recorder for confirmation. The crew
leader also kept track of crew observations visually to
prevent duplication or missed nests. All movement
within the colony was kept to a minimum. To reduce
disturbance, hand signals were often used to relay in-
structions, such as change direction, stop, proceed, etc.

Contents of Great Black-backed and Herring Gull
nests (empty, number of eggs and/or number of chicks)
were recorded. Any structure more elaborate than a
simple scrape with some built-up edge was deemed a
nest (Diamond 2001*). To distinguish between Great
Black-backed and Herring Gull eggs, the following cri-
teria were used: a

Locations of grouped Herring Gulls within the larger
colony of Great Black-backed Gulls were noted.

2014 MACKINNON AND KENNEDY: GULL BREEDING DECLINE ON Boot ISLAND, NOVA SCOTIA 167

The approximate position of individual Herring
Gulls on nests in relation to adjacent features (bush,
rock, etc.) was visually tracked.

The sizes of all eggs, in each nest, were visually at-
tributed to species; Herring Gull eggs are about 19%
smaller than Great Black-backed Gull eggs (Bent
1963*), although there is some variation in size be-
tween regions (see Pierotti and Good 1994; Good
1998). The collective size of all eggs in a specific nest,
along with the above supporting criteria, were used to
identify it as either Great Black-backed or Herring Gull.

A survey typically required 4-6 hours to complete.
To minimize further disturbance within the gull colony
and adjacent Great Blue Heron and Double-crested
Cormorant colonies, nests were not marked to deter-
mine a correction factor (e.g., Lincoln index).

To test whether the population decline of gulls was
significant, we used a linear regression analysis. The
regression and regression equations were calculated
using the statistical package within Microsoft Excel
while t and P values were calculated as per Schefler
(1980); applying the two-tailed test.

Results and Discussion

Herring Gull nests were first recorded in significant
numbers in 1976 (712 nests; Figure 2). The number of
nests peaked in 1986 (727 nests), but by 2010, only 2

Great Black-backed
Gull

NUMBER OF NESTS

a

1975 1980 1985 1990

FiGuRE 2. Number of nests (occupied

nests were recorded in the survey. The near-linear
decline in the number of Herring Gull nests on Boot
Island from 1986 to 2000 suggests a constant pressure
on these birds (R* = 0.99, y = —46.394x + 92 857,n =7
surveys, t= 25.7, P< 0.01) (Figure 2).

The number of Great Black-backed Gull nests were
also first recorded in significant numbers in 1976 (N
1005). The number of nests peaked in 1992 (N=1467),
and by 2010, the number of nests had decreased by
44% to 819. The decline of Great Black-backed Gull
nests was significant but a more variable decline from
1992 to 2010 (R? = 0.82, y = -31.434x + 64 010, n=9
surveys, t= 5.74, P < 0.01) (Figure 2).

Although the survey window remained the same
between 1976 and 2010, the ratio of occupied to un-
occupied Great Black-backed Gull nests fell from a
high of 22.3:1 in 1986 to just 2.3:1 in 2010 (Figure 3).
While the Great Black-backed Gull colony was declin-
ing, there was a noted increase in the proportion of
empty nests. This could be a result of young and inex-
perienced breeders nesting later, although we believe
that the increase in empty nests reflects an increase in
predation on eggs and/or chicks. For Herring Gulls, the
ratio of occupied to empty nests ranged from 3.7:1 in
1986 to an average of 4.1:1 from 1996 to 2000 (Figure
4). The observed differences in the ratios may reflect
both the timing of heavier predation and an artefact

i
"Ores ow
i TO Oe

1995 2005 2010

YEAR

2000

and unoccupied) of Great Black-backed Gulls (Larus marinus) and Herring Gulls (L.

argentatus) found at Boot Island National Wildlife Area, Nova Scotia, 1976 to 2010.

168 THE CANADIAN FIELD-NATURALIST

of the survey window, as Great Black-backed Gulls
begin egg laying earlier than Herring Gulls. It 1s note-
worthy that there was an increase in the ratio of occu-
pied to empty Great Black-backed Gull nests from
2006 to 2010 when the Herring Gull colony had been
reduced to fewer than 25 nests (Figures 3 and 4). Dur-
ing this same period, Coyote began to visit the island
and predation by this species became a more obvious
and regular event (MacKinnon ef a/. 2007).

It is well documented that Great Black-backed Gulls
are more aggressive and may out-compete Herring
Gulls as well as preying on young birds (Threlfall 1968;
Rome and Ellis 2004; Cotton 2009). This may at least
partly explain the earlier onset of the Herring Gull de-
cline. However, by the early 1990s, both species were
in decline and other factors must be considered.

During this period of decline in both gull species
and the increasing proportion of empty Great Black-
backed Gull nests, the breeding numbers in the adja-
cent Great Blue Heron and Double-crested Cormorant
colonies remained relatively constant. From 1984 to
2010, Great Blue Herons and Double-crested Cor-
morants averaged 53 nests (range 38-75, n = 13 sur-
veys) and 232 nests (range 147-351, n = 13 surveys),
respectively (MacKinnon et al. 2010*). No obvious
overall change or trend in colony size was observed.
The factors influencing the gull decline are apparently

1600
1400
1200
1000

80

oO

60

NUMBER OF NESTS
Oo

40

oO

20

oO

=)

1976 1980 1984 1988

Vol. 128

not affecting the health of the adjacent tree-nesting
colonial species.

It is worth noting that, based on 2002 aerial survey
data (753 pairs), the Great Black-backed Gull colony
on Boot Island, Nova Scotia, was the largest in Canada
(Cotter et al. 2012*). Furthermore, if Herring Gulls
were deserting Boot Island for other locations, current-
ly the closest large Herring Gull colony is situated in an
inland bog 195 km to the southwest on Briar Island,
Nova Scotia. (Environment Canada 2008*; Cotter e¢
Ge 202):

Food Availability
Human refuse and landfills

Changes in the handling of human refuse, such as
the closure or reduction of landfills, have been cited as
one reason for decreases in gull populations (Drury
1973; Shoals Marine Laboratory 2010*). In Nova Sco-
tia, a solid waste management strategy was released in
1995 (Friesen 2000), although organic garbage was
not banned from the province’s landfill sites until 30
November 1998 (Speed 2000*). By this time, the Her-
ring Gull colony on Boot Island had already collapsed
and Great Black-backed Gull numbers were in decline.
Although competition for limited food resources may
have been partly responsible for the changes in the
colony, we do not have any direct evidence that the
changes in management of human refuse in the vicinity

HB 2

1996 2000 2004 2008

YEAR

FIGURI 3. Comparison of occupied nests (solid bars) with empty nests (open bars) of Great Black-backed Gulls (Larus mar-
inus) at Boot Island National Wildlife Area, 1976 to 2010.

20 AC , =
2014 MACKINNON AND KENNEDY: GULL BREEDING DECLINE ON Boot ISLAND, NOVA ScoTiIA 169

NUMBER OF NESTS

1986 1990 1994

lli =

1938
YEAR

2002 2006 2010

FIGURE 4. Comparison of occupied nests (solid bars) and empty nests (open bars) of Herring Gulls (Larus argentatus) at
Boot Island National Wildlife Area, Nova Scotia, 1986 to 2010.

of the colony played a significant role in the observed
reductions in gull numbers.

The cod fishery

In 1992 the North Atlantic Cod (Gadus morhua)
population collapsed and the Canadian government
placed a moratorium on the fishery. Chapdelaine and
Rail (1997) were able to link this decline in the Atlantic
cod fishery on the north shore of the Gulf of St.
Lawrence with similar changes in nesting Herring Gull
numbers on nearby islands. We do not know where the
Boot Island Herring Gulls spend their time outside

the breeding season, although some birds may winter

along the eastern seaboard of the United States (see
Drury and Nisbet 1973*; Gaston er al. 2008*). It may,
however, not be coincidental that the rapid decline in

Herring Gull numbers on the north shore of the Gulf

of St. Lawrence from 1988 to 1993 (Chapdelaine and
Rail 1997: Cotter and Rail 2007) roughly coincides
with the Boot Island population crash from 1987 to
2002. Furthermore, changes observed at the Boot Island
colony generally reflect regional trends for Great Black-
backed and Herring Gulls in Nova Scotia from 1971
to 2002 (Cotter et al. 2012*).

Predation and disturbance
Coyote

The decline in both gull species coincide with the
expansion of Coyote into Nova Scotia (Parker 1995).
The range expansion and population increase of Coy-
otes in the province was rapid: provincial fur harvesters
removed 777 animals in 1990-1991, with an increase
to 1.887 animals in 1994-1995 (O’Brien and Boudreau

1998*). Coyotes are opportunistic predators (Berg and
Chesness 1978; Parker 1995) and, in recent years,
tracks and scat have been found on a number of off-
shore islands. Coyote sign (scat) was first recorded on
Boot Island in 1998 with additional evidence (tracks
and scat containing small bones) in 2000 and 2002. In
2004, one animal was observed feeding on Great Black-
backed Gull eggs (MacKinnon ef al. 2007). This ani-
mal readily swam across the narrow channel dividing
Boot Island from the mainland and, on arrival, prompt-
ly started consuming gull eggs. The presence of Coy-
otes on the island during nesting is now likely a year-
ly occurrence, as one animal was observed in the gull
colony in 2010 and again on 12 May 2014 (MacKin-
non et al. 2010*).

American Mink and Raccoon

We also recorded the presence of American Mink
on the island in 2004, 2008 and 2010 and, more recent-
ly, Raccoon in 2010. Both are notorious consumers of
eggs and the young of ground-nesting birds and Rac-
coons have a significant impact on gull colonies (Kad-
lec 1971: Ellis ef al. 2007). American Mink have also
been predators in off-shore Nova Scotia tern colonies,
such as those on Country Island and The Brothers
(D’Eon 2005*).

Bald Eagle

Bald Eagles are frequently observed over Boot Is-
land and, in 2008, a breeding pair took up residence
in a large White Spruce (Picea glauca) adjacent to the
gull colony. Eagles are known predators of gulls (Bue-
hler 2000) and, although we did not observe a preda-

170

tion attempt during our surveys or search for prey re-
mains under the nest, there was a conspicuous absence
of gull nests in the immediate vicinity of the eagle nest.
As approximately 50% of the gull colony is within
300 m of the eagle nest, the close proximity of eagles
may influence the reproductive success of breeding
gulls through disturbance of normal activities during
the breeding season (White e7 al. 2006).

Egg collection by humans

Hébert (1989) reported that, among other factors,
human predation on gull eggs for food (egging) was
partly responsible for the observed decline in Herring
Gulls on Kent Island, New Brunswick. No comparable
activity has been reported for Boot Island, and we
attribute this absence to the hazardous waters surround-
ing the island and the lack of an “egging tradition” in
the largely agriculture-based adjacent community. Thus,
egging is not considered to be a factor in the Boot Is-
land gull decline.

Conclusion

Populations of nesting Great Black-backed and Her-
ring Gulls have been in decline on Boot Island, Nova
Scotia, for about 25 yrs. The earlier decrease in Her-
ring Gull numbers may be attributed to aggression by
the larger and competing Great Black-backed Gulls;
however, declines in both species suggest that other,
likely multiple, factors were involved. The loss of the
Atlantic cod fishery in 1992 has been linked to gull
declines in a number of colonies (Chapdelaine and Rail
1997) and this regional reduction of food availability
may also be a contributing factor in the loss of Herring
Gulls on Boot Island.

Coyote access to near-shore islands, following their
expansion into Nova Scotia in the early 1980s, has re-
sulted in nest loss from predation on Boot Island. Addi-
tional predation of eggs and chicks by American Mink,
Raccoons, and Bald Eagles may have further con-
tributed to an increase in nest failure in recent years.
These stressors have become cumulative to the point
that the Herring Gull colony on Boot Island has been
essentially extirpated. In 2010, the Herring Gulls and
nests were almost completely absent, while the Great
Black-backed Gull colony had declined to 56% of its
peak in 1990. It is important to note that this study is
based on short duration field observations conducted
over a number of years and that any definitive cause
and effect is beyond the scope of this paper. Further
work, such as determining the foraging and wintering
areas of the Boot Island gulls is required.

A combination of many factors appears to be respon-
sible for the observed declines. A broad reduction in
food resources caused by changes in fisheries and bet-
ter landfill management; the arrival of the Coyote along
with other nest predators, such as American Mink and
Raccoon; and the recent establishment of an active
Bald Eagle nest within the colony may all have con-
tributed to the observed decline in gulls on Boot Island.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Acknowledgements

The survey data for Boot Island National Wildlife
Area have been collected with the assistance of a num-
ber of participants over many years. We thank the fol-
lowing crew leaders, staff, and volunteers for their
efforts: Jeff Atkinson, Peter Barkhouse, Bill Barrow,
Barb Boiduk, Susan Bowes, Adam Campbell, Cyril
Coldwell, Donald Colpitts, Graham Daborn, Tony
Erskine, Keith Galbraith, Jocelyne Gauvin, Gay Han-
son, L. Hanson, Peter Hicklin, Randy Hicks, Matthew
Horsman, Ron Hounsell, Etta Hudgins, Jason Hudson,
Larry MacDonnell, Andrew Macfarlane, Nancy (Lutz)
MacKinnon, Adam MacPherson, Randy Milton, Reg
Newell, Darren Ogden, Cindy Pauley, Mark Pulsifer,
Mathew Robinson, Sylvie Roussel, Vishalla Singh,
Peter C. Smith, and Jim Wolford. Our apologies to
those we may have missed. We also thank Paul Cham-
berland, Kevin Davidson, Tony Gaston, Peter Hicklin,
and two anonymous reviewers for commenting on
earlier drafts of this paper.

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Received 6 December 2013
Accepted 5 February 2014

Distribution and Abundance of Baling Twine in the Landscape

Near Osprey (Pandion haliaetus) Nests: Implications for Nestling
Entanglement

RENEE SEACOR!', KAYHAN Ostovar!, and MARCO RESTANI2:3

Environmental Sciences Program, Rocky Mountain College, 1511 Poly Drive, Billings, Montana 59102 USA
“Department of Biological Sciences, St. Cloud State University, St. Cloud, Minnesota 56301 USA
*Corresponding author: restani@stcloudstate.edu

Seacor, Renee, Kayhan Ostovar, and Marco Restani. 2014. Distribution and abundance of baling twine in the landscape near
Osprey (Pandion haliaetus) nests: implications for nestling entanglement. Canadian Field-Naturalist 128(2): 173-178.

Polypropylene baling twine used by Ospreys (Pandion haliaetus) during nest construction creates a risk of entanglement for
nestlings and adults on the Yellowstone River, Montana. In 2013, we evaluated the abundance of twine in 2-km-radius buffer
zones centred on 38 nests for three categories of road density. We found more twine per kilometre along roads in low (7 = 19)
and moderate (n = 13) road density nest buffer zones than in high road density nest buffer zones (n = 6). The estimated total
amount of twine found along roads in nest buffer zones ranged from 0 to 2602 m and did not differ among road density strata.
The percentage of Osprey nests containing twine was highest in low (63.2%) and moderate (61.5%) road density nest buffer
zones and lowest (33.3%) in high road density buffer zones, which reflected a gradient from rural and suburban to urban land-
scapes. The estimated total amount of twine within a nest buffer zone did not predict whether a nest contained twine. The
amount of twine found in seven nests destroyed by wind or power company personnel ranged from 0 to 206 m and was not
correlated with the amount of twine found in their buffer zones. During the 2012 and 2013 breeding seasons, four of 120 nestlings
(3.3%) became entangled in twine: two were cut free and fledged normally, one died, and one was euthanized. The abundance
of twine in the environment surrounding nests and its slow rate of biodegradation mean that vigilance by citizen scientist nest
monitors and assistance from power companies are the only short-term solution to reducing mortality resulting from entanglement.

Key Words: baling twine; entanglement; Montana; mortality; Osprey; Pandion haliaetus; Yellowstone River

Introduction

Many bird species incorporate atypical materials into
their nests (Hansell 2000). A variety of species use
anthropogenic nesting materials, such as plastic trash,
twine, and rope, and the prevalence of plastics in nests
is thought to be increasing (Montevecchi 1991; Antczak
et al. 2010; Votier et a/. 2011). Because plastics degrade
slowly, using the percentage of nests containing plas-
tics as an index of their abundance in the environment
over time requires both a constant search effort and
knowledge of bird population trends (Ryan ef al. 2009).
Moreover, the presence of plastics in large, long-lasting
nests suggests that such an index is best considered
over decades rather than years (Bond et al. 2012).
Although some individuals within a species benefit by
signaling conspecifics through the use of artificial nest-
ing materials (Hansell 2000), plastic rope and twine
present a potential risk of nestling and adult entangle-
ment. In fact, mortality associated with the use of these
nesting materials is well documented in White Storks
(Ciconia ciconia; Kwiecinski et al. 2006), Northern
Gannets (Morus bassanus; Votier et al. 2011), and Great
Grey Shrikes (Lanius excubitor; Antezak et al. 2010).

The stick nests of Ospreys (Pandion haliaetus) often
include both natural items (e.g., wings, bones, dung,
and sod) and artificial objects (e.g., clothing, shoes,
toys, and rope) (Allen 1892). The propensity of nest-
ing Ospreys for using discarded polypropylene baling
twine, which is used by farmers and ranchers to store

hay and straw, has recently elicited some conservation
concern. For example, all nests near agricultural fields
at two study areas in Montana contained twine, which
had entangled young in nearly 5% of nests (Blem ef al.
2002). In west-central Saskatchewan, approximately
12% of nestlings had become entangled (Houston and
Scott 2006). Both groups of researchers concluded that
entangled nestlings would have died without human
intervention. Baling twine hanging from Osprey nests
on power poles has come into contact with electrical
lines and is thought to have ignited fires, which further
endangered nestlings (personal communication with
power company linemen).

During visits to Osprey nests in 2012, we noticed
that adults had often incorporated baling twine into
nests. Various lengths of twine were found in the nest
bowl, wrapped around large sticks, or hanging from the
nest rim. Three out of 50 (6%) nestlings had become
so badly entangled in twine that they had to be rescued,
had died, or had to be euthanized (Figure 1). While
driving between nests, we often observed remnants of
baling twine along the shoulders of secondary roads.
The prevalence of plastic string, rope, and twine in the
environment surrounding nests has been cited as a rea-
son for their occurrence in nests (Blem ef al. 2002;
Antezak et al. 2010; Bond et a/. 2012), and our obser-
vations prompted us to investigate the abundance of
discarded baling twine in the landscape surrounding
Osprey nests. Our objectives were to estimate and com-

We)

174

THE CANADIAN FIELD-NATURALIST

FiGure |. An Osprey (Pandion haliaetus) nestling badly entangled in baling twine and later euthanized, Yellowstone River,
Montana, 2012. Compare the normal right foot and tarsus with the grossly swollen and disfigured left foot and tarsus.

Photo: Cameron Sapp.

pare the amount of twine along roads near Osprey nests
across a gradient from rural to urban landscapes and to
assess the relation between the presence of twine in
nests and the amount of twine found along roads near
nests. Our results have allowed us and our partner, the
Yellowstone Valley Audubon Society (Y VAS), to tar-
get specific areas for cleanup and specific landowners
for conservation education.

Study Area

The study area extended approximately 945 km
along the Yellowstone River floodplain from the Wy-
oming—Montana border (44°59'29"N, 110°30'S8"W)
to the Montana—North Dakota border (47°45'28"N,
104°02'42"W). As the river flows downstream, it
changes from a high-gradient, clear, cold mountain sys-
tem to a low-gradient, turbid, warm-water system. The
geomorphology of this dynamic, unregulated river
comprises wide multi-channel reaches, forested islands,
gravel bars, and straight channels with cliffs. Vegeta-
tion along the riparian corridor reflects the decrease
in elevation, from higher-elevation (1750 m) forests
dominated by conifers (e.g., Juniperus, Pinus, Picea,
and Pseudotsuga spp.) to river bottoms (615 m) com-
posed of a mixture of shrubs (e.g., Salix spp.) and
deciduous trees (e.g., Populus spp.). Anthropogenic
land uses along the river include small grain farming,
livestock grazing, recreation, and urban-centred indus-
tries, such as oil refining and coal-fired power gener-
ation. Billings (population 106 000) is the largest city
along the river and is situated near the middle of the
study area. Climate is semi-arid.

Methods
Along with 15-20 citizen scientists from YVAS, we
conducted fieldwork during the 2012 and 2013 breed-

ing seasons. Beginning in April, we used binoculars
and spotting scopes to survey the study area for nesting
Ospreys. We used a Global Positioning System (GPS;
Garmin GPSMAP 62) to record the location of nests
and uploaded these coordinates into a geographic infor-
mation system (GIS). Nests were observed from the
ground at approximately 1-week intervals throughout
the breeding season to determine occupancy (Steenhof
1987) and to estimate dates of egg laying, hatching, and
fledging; brood size; and reproductive success, i.e.,
number of young fledged (Harmata ef a/. 2007).

We used ArcGIS (v. 10.1, Environmental Systems
Research Institute, Redlands, California, USA) to cre-
ate a 2-km-radius buffer zone around each of the 71
Osprey nests located in 2012 to identify roads for twine
sampling. The size of the buffer zones was large enough
to contain an adequate number and length of roads to
survey based on our sampling protocol (see below) and
small enough to allow us to survey the entire Osprey
study population in one season. We obtained a state
roads GIS from the Montana Geographic Information
Clearinghouse (National Resource Information System
2013*), which categorized roads into seven types:
United States Forest Service, primary, secondary, Na-
tional Highway Administration non-interstate, inter-
State, city-county, and urban. The roads layer was
added to the georeferenced nest buffer zones, and roads
were then highlighted and clipped to each buffer zone.
Because of safety considerations, we did not survey
Interstates 90 or 94 and, therefore, for each buffer zone
containing those roads, we subtracted their lengths from
the total length of roads. We also omitted six nest buffer
zones from further consideration because they con-
tained less than 10 km of roads, the lower limit of our
sampling protocol (see below). We partitioned the re-
maining 65 nest buffer zones into three road density

2014 SEACOR ET AL.: BALING TWINE ON ROADS NEAR OsprEY NESTS 175

strata based on the total length of roads they contained:
low, 10 to < 20 km; moderate, 20 to < 40 km: and high,
> 40 km). These categories spanned a gradient from
agricultural land (rural) to cities (urban) across the het-
erogeneous study area.

We did not have any a priori information (e.g., vari-
ation in metres of twine per buffer zone) to guide sam-
pling effort or allocate kilometres of road per stratum
to survey for baling twine, which are required for Ney-
man-like optimization (Cochran 1977). Therefore, we
used square-root of stratum size (i.e., total kilometres
of road within each stratum) as a reasonable compro-
mise between equal and proportional allocation of sam-
pling effort (Bankier 1988). The allocated kilometres
of road to survey for twine in each nest buffer zone (i.e.,
sampling effort) increased with road density stratum:
low (10-12 km), moderate (14-16 km), and high (>
18 km). When the buffer zones of different nests over-
lapped by = 20%, we randomly selected one zone to
sample for baling twine, and then, in the field, we did
not survey any road segments in the overlapping area
to maintain statistical independence in analyses.

In the field during the 2013 nesting season, we used
an iPad (Apple, A1430) and the ArcMap application
to select and number sequentially each 2-km road seg-
ment within a nest buffer zone. A coin flip determined
whether a particular segment was surveyed for baling
twine, a process repeated up to the allocated number
of kilometres to be surveyed for each buffer zone
depending on its road density (i.e., low: 10 or 12 km,
moderate: 14 or 16 km, and high: > 18 km). For exam-
ple, we surveyed either five or six 2-km road segments
in each nest buffer zone in the low road density stratum
for a total search effort of either 10 or 12 km. Ina few
instances, we were unable to survey a selected 2-km
road segment because access was restricted. Alterna-
tive roads were randomly chosen where possible, but
when these were unavailable, we reduced the number
of kilometres surveyed in that zone. Travelling by bicy-
cle allowed us to look for twine along both sides of
selected roads. We collected twine from the road and
pavement edge to fencing along the shoulder (about
4 m in width). Twine was bagged and labelled by nest
buffer zone, and its length (to the nearest metre) was
estimated with a tape measure in the laboratory.

Data transformations were unsuccessful in achiev-
ing normal distributions; thus, we used the Kruskal-
Wallis independent samples test to determine whether
the number of metres of baling twine collected per
kilometre of road differed among the three road den-
sity strata (SPSS v. 19.0, IBM, Armonk, New York,
USA). We conducted the same test to determine wheth-
er the total amount of twine in a buffer zone (1.¢., metres
of twine collected per kilometer of road multiplied by
total kilometres of roads within a buffer zone) differed
among road density strata. We used Dunn’s test to iden-
tify differences in rank among the three road density
strata following a significant Kruskal-Wallis test. Al-

though Kruskal-Wallis is a ranks (not a means or medi-
an) test, we have provided box plots of data distribu-
tion to aid interpretation.

In 2013, we noted which Osprey nests contained
baling twine during the banding of nestlings or by ob-
servation from the ground. We used logistic regression
to examine the relationship between estimated total
amount of twine in a buffer zone and the presence or
absence of twine in the nest. Goodness of fit was eval-
uated using the Hosmer—Lemeshow test. We also meas-
ured the amount of twine (metres) in nests that had
either blown down during storms or had been removed
by power company personnel. We used linear and
non-linear regression models to test for a relationship
between the amount of twine in those nests and the
amount of twine found along roads in the correspon-
ding buffer zone (per kilometer of road and total).
The significance of all statistical tests was accepted
atP= 0,05.

Results

Together with YVAS volunteers, we surveyed
approximately 925 km of the Yellowstone River for
Osprey nests in 2012 and 2013. All nests considered
in the baling twine analyses (n = 65) were built on
artificial structures (i.e., power poles, nest platforms,
or bridge spans). Of the buffer zones around these
nests, 25 were located in areas of low road density
(10 to < 20 km), 23 in areas of moderate road density
(20 to < 40 km), and 17 in areas of high road density
(> 40 km). The number of nests available for statistical
analysis decreased after we omitted buffer zones that
overlapped by > 20%, leaving us with 19, 13, and 6
nests in the low, moderate, and high road density strata,
respectively. The mean number of kilometres of roads
surveyed for twine in each nest buffer zone varied
across road density strata: low, 10.9 km (SD 0.7, range
10-12); moderate, 13.5 km (SD 1.2, range 14-16); and
high, 21.3 km (SD 2.9, range 18-28).

The ranks (amount) of baling twine collected per
kilometre of road surveyed within nest buffer zones
differed among low, moderate, and high road density
strata (H = 6.10, df = 2, P = 0.047); low road density
buffer zones had significantly higher ranks than high
road density buffer zones (Figure 2A). However, the
ranks of estimated total length of twine in buffer zones
did not vary with road density (1 = 3.12, df= 2, P=
0.210) (Figure 2B). The percentage of Osprey nests
containing twine was nearly twice as high in the low
(63.2%, 12 of 19) and moderate (61.5%, 8 of 13) road
density strata as in the high road density stratum (33.3%,
2 of 6).

The total amount of baling twine estimated within
a nest buffer zone did not predict whether a nest con-
tained twine (y7= 0.36, df= 1, P = 0.547). Although the
data fit the logistic regression model, only 23 of 38
nests (60.5%) were classified correctly; all but one con-
tained twine. We observed 93.7% more nests without

176

Amount of twine (m/km)

3000

2500

2000

1500

1000

Amount (m) of twine in buffer

500

Low (10 to <20 km)

THE CANADIAN FIELD-NATURALIST

Moderate (20 to <40 km)

Vol. 128

High (=40 km)

Road density within nest buffer

FIGURE 2. Amount of baling twine in 2-km-radius buffer zones around Osprey (Pandion haliaetus) nests along the Yellowstone
River, Montana, 2013. A: Metres of twine per kilometre of road; B: Estimated amount of twine (m) in buffer zones.
Note: lines mdicate median, boxes show the 25th and 75th percentiles, whiskers indicate the 10th and 90th percentiles,

and dots show outliers.

twine (15 of 16 nests) than predicted by the model. We
collected twine from seven nests that had either blown
down in storms or were removed by power company
personnel before or after the nesting season in 2013
(Table 1). No relationship existed between the amount
of twine found in nests and either the amount of twine
found per kilometre of road (linear: F = 1.01, df= Ihe
P = 0.362; non-linear: F = 1.03, df= 1, P=0.357) or
the total amount estimated in nest buffer zones (linear:
F= 1.05, df= 1, P= 0.353: non-linear: F = 1.09, df= 1,
P=0.354).

Ospreys occupied 28 nests in 2012 and 50 nests in
2013; the large increase in the number of occupied
nests located in 2013 reflects the greater field effort
on the part of the citizen scientists. Ospreys fledged
an average of 1.8 nestlings per occupied nest in 2012
and 1.4 nestlings per occupied nest in 2013. During
2012, two of 21 (9.5%) successful nests contained three
nestlings entangled in twine: one was found dead dur-
ing the banding visit, one had to be euthanized. and one
was cut free and appeared to fledge normally (three
of 50 [6.0%] nestlings entangled). During 2013, one

2014

TABLE 1. Amount of baling twine found in Osprey (Pandion
haliaetus) nests blown down during storms or removed by
power company personnel and the estimated amount of twine
in a 2-km-radius buffer zone around each nest, Yellowstone
River, Montana, 2013.

Length of twine, m

Per km of road

Nest ID no. In nest in buffer zone —_ In buffer zone
137 0 0.2 5

295 l 3.4 63

318 l Re > 22

350 206 2.2 43

354 0 2.3 52

374 131 26.3 447

598 64 4.3 74

of 35 (2.9%) successful nests contained an entangled
nestling; it was cut free and appeared to fledge nor-
mally (one of 70 [1.4%] entangled nestlings).

Discussion

We found significantly more baling twine per kilo-
metre of road in Osprey nest buffer zones in low and
moderate road density areas than in high road density
areas, which reflected the gradient from rural and sub-
urban to urban landscapes. Although we did not find a
direct relationship between the estimated total amount
of twine in a buffer zone and its presence in nests, the
percentage of nests containing twine was highest in
rural and suburban areas and lowest in urban areas. The
amount of twine found along roads surrounding nests
varied considerably: rural buffer zones, 0-778 m; sub-
urban buffer zones, 0-2602 m; and urban buffer zones,
0-506 m.

From our observations in 2012, we knew that bal-
ing twine accumulated along roads, and this prompt-
ed us to initiate this study, the first to estimate twine
abundance near Osprey nests. Surveying roads was
convenient and efficient. We believe that measuring the
amount of twine found along roads near nests under-
estimated its abundance in the landscape, because adja-
cent pastures and feedlots often contained discarded
twine. An untested assumption of our study was that
a positive relationship exists between the amount of
twine along roads and the amount of twine in the 2-
km-radius buffer zones around nests.

Most farmers and ranchers on our study area store
hay and straw in large, round bales, each of which is
a potential source of twine for nest-building Ospreys.
A single bale contains approximately 115 m of twine
(Houston and Scott 2006). One hay bale can feed one
horse or steer per month during winter; therefore, five
horses fed for five months require approximately 25
bales (and 2875 m of twine). It remains unclear how
twine is transferred from pastures and feedlots to roads,
but we suspect that most had blown off the flat beds
of ranch pickup trucks, where we often observed it
loosely coiled or piled. In fact, we observed one such

SEACOR ET AL.: BALING TWINE ON ROADS NEAR OsprEY NESTS

iy

instance while conducting twine surveys. Regardless
of the mechanism, Ospreys in southern Montana en-
counter an abundant and apparently available resource
for use as nesting material.

Without human intervention, 3.3% of Osprey nest-
lings on the Yellowstone River would have died from
twine entanglement, which is lower than the 11.7%
nestling mortality reported by Houston and Scott (2006)
in Saskatchewan. In our study, the percentage of suc-
cessful nests in which young were entangled (5.4%)
was higher than that reported from nearby central
Montana (4.6%; Blem et al. 2002). Current estimates
of mortality resulting from entanglement on the Yel-
lowstone River does not appear to be of conservation
concern regarding the potential for additive mortality
because the Osprey population had reproductive rates
in 2012 and 2013 greater than those needed to support
a stable population (Henny and Wight 1969; Postupal-
sky 1989). However, twine entanglements did raise
animal welfare concerns because nestlings and adults
suffered slow deaths from starvation, constriction, and
infection. The only practical, short-term solution strat-
egy to prevent deaths from entanglements is to have
citizen citizen-scientist volunteers regularly visit nests
and contact us if they observed entangled nestlings. In
each such instance, power company cooperators have
responded quickly to our requests for assistance.

The amount of baling twine we collected from sev-
en nests that had been destroyed by wind or removed
by power company personnel varied by two orders of
magnitude (0-206 m per nest). Similar variation was
reported in the mass of plastics found in five nests
constructed by Northern Gannets (Votier ef a/. 2011).
Several proximate factors have contributed to the ex-
tent that artificial materials are used in nest construc-
tion or as nest adornments: abundance in the environ-
ment (Blem et al. 2002; Bond et al. 2012), age of the
nest (Votier ef al. 2011; Bond et al. 2012), and age of
the nest builder (Sergio et a/. 2011). On the Yellow-
stone River, no relationship existed between the amount
of twine found in the landscape and the amount found
in Osprey nests. We lacked information on the age of
nests, but it appeared that larger nests, which were
presumably older, contained more twine; however, the
limited sample of seven nests made this inference ten-
tative. Although we found a variety of anthropogenic
materials in nests, the largest component was baling
twine, and such selectivity has been noted in a diver-
sity of bird species from seabirds to raptors to passer-
ines (e.g., Antezak er al. 2010; Sergio et al. 2011; Voti-
er et al. 2011; Bond et al. 2012). Whereas the cost of
using baling twine in Osprey nests was clear, any poten-
tial benefits have yet to be determined (e.g., conspe-
cific signaling; Sergio ef al. 2011).

Educating the public about the risks that baling twine
presents to nesting Ospreys is a component of our re-
search project. Together with our partner, Y VAS, we
mail informational brochures to landowners along the

178

Yellowstone River, with the hope of promoting respon-
sible handling and disposal of twine. We are also iden-
tifying what we believe are significant sources of the
twine that ends up along roads (e.g., feedlots). Poly-
propylene baling twine degrades slowly and clean-up
efforts along roads and in pastures and feedlots will
have to be coordinated for decades to reduce dangers to
Ospreys (see also Ryan ef al. 2009; Bond et al. 2012).
It appears that farmers and ranchers are slowly switch-
ing from using twine to bale hay and straw to using
plastic netting or wraps. We rarely find the latter mate-
rials in nests and view such a transition as a potential
long-term solution benefitting Ospreys.

Acknowledgements

This project was a cooperative effort between the
Yellowstone River Research Center (Rocky Mountain
College) and the Yellowstone Valley Audubon Society.
We thank C. Sapp and the dozens of citizen scientists
who dedicated countless hours in the field to monitor-
ing nests in 2012 and 2013. W. Newton, statistician at
the Northern Prairie Wildlife Research Center (United
States Geological Survey), kindly volunteered his time
and expertise to design the twine sampling protocol.
He, A. Harmata (Montana State University) and two
anonymous reviewers provided valuable comments on
the draft manuscript. Generous in-kind support was
provided by Beartooth Electric Cooperative, Montana—
Dakota Utilities, NorthWestern Energy, Park Electric
Cooperative, Yellowstone Valley Electric Cooperative,
and Yellowstone Valley Tree Surgeons. Many land-
owners, the United States Bureau of Land Manage-
ment, the city of Billings, and the Audubon Education
and Conservation Center graciously granted permis-
sion to monitor and access nests on their properties.
This research was approved by the Institutional Animal
Care and Use Committee at Montana Fish, Wildlife
and Parks (IACUC FWP07-2012) and through a Mon-
tana Scientific Collector’s Permit (2013-031) to K.
Ostovar and a Federal Bird Banding Permit (22518)
to M. Restani. Funding was provided by the Yellow-
stone River Research Center, Toyota TogetherGreen,
The Cinnabar Foundation, The Peregrine Fund, Yel-
lowstone Valley Audubon Society, Montana Audubon,
Rocky Mountain, College, St. Cloud State University,
and Montana Fish, Wildlife and Parks.

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Library, Bozeman, Montana, USA. Accessed 15 October
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Received 7 December 2013
Accepted 31 January 2014

Cues used by Predators to Detect Freshwater Turtle Nests may Persist
Late into Incubation

JuLia L. RiLey! and JACQUELINE D. LitzGus!?

Department of Biology, Laurentian University, 935 Ramsey Lake Road, Sudbury, Ontario P3E 2C6 Canada
“Corresponding author: jlitzgus@laurentian.ca

Riley, Julia Le and Jacqueline D. Litzgus. 2014. Cues used by predators to detect freshwater turtle nests may persist late into
incubation. Canadian Field-Naturalist 128(2): 179-188.

Previous studies have found that turtle nest depredation is concentrated immediately post-oviposition, likely because cues
alerting predators to nest presence are most obvious during this time. In Algonquin Provincial Park, Ontario, we examined
the frequency of nest depredation during the incubation period for Snapping Turtles (Chelydra serpentina [Linnaeus, 1758])
and Midland Painted Turtles (Chrysemys picta marginata [Agassiz, 1857]). Contrary to most past findings, nest depredation
occurred throughout the incubation period for both species. In fact, 83% and 86% of depredation interactions with Snapping
and Painted Turtle nests, respectively, occurred more than a week after oviposition at our study site. Peaks in nest depredation
(weeks with >10% nest depredation) occurred late in incubation and may have coincided with hatching. Trail cameras deployed
at four nesting sites revealed six predator species interacting with nests. The presence of predators at nest sites increased late in
the incubation period indicating a persistence or renewal (from hatching) of cues; additional research is necessary to determine
the nature of these cues. These findings have implications for both research and turtle conservation. Further research should
examine the relationship between temporal changes in predator species’ density and patterns of nest depredation. Additionally,
in areas where protective nest caging is used as a species recovery action, it may be important to ensure that cages remain in
place throughout the incubation period until emergence of hatchlings.

Key Words: Snapping Turtles; Chelydra serpentina; Midland Painted Turtles; Chrysemys picta marginata; Turtles; Algonquin

Provincial Park; Ontario; ecology; nest depredation; predator detection; predators; species recovery

Introduction

Freshwater and marine turtles are among the most at-
risk groups of vertebrate animals, and one of the many
threats to turtle populations is unnaturally high levels of
nest predation (Gibbons et a/. 2000; Spotila 2011). A
broad range of animals from multiple taxa depredate
turtle nests, including mammals, birds, reptiles, and
invertebrates (Ernst and Lovich 2009). High levels of
turtle nest depredation commonly occur in human-
disturbed landscapes because food resources associated
with human presence (e.g., trash, agricultural crops)
increase the abundance of subsidized predators of tur-
tle eggs (Christiansen and Gallaway 1984; Garber and
Burger 1995; Mitchell and Klemens 2000; Seburn
2007: Fordham et al. 2008; Kurz et al. 2011). In addi-
tion, the removal of top predators in urban areas has
caused an increase in the number of mesopredators,
leading to a concomitant increase in depredation of tur-
tle eggs (Barton 2003; Prugh er al. 2009; Ritchie and
Johnson 2009). In some turtle populations, subsidized
mesopredators annually depredate 100% of nests (e.g.,
Geller 2012), resulting in chronic prevention of recruit-
ment (Spinks et a/. 2003; Seburn 2007; Fordham et al.
2008). Substantial repeated reductions in recruitment
(approximately a 50% or more decrease in egg and/or
hatchling survival) perpetuate population declines
(Crouse et al. 1987; Crowder et al. 1994; Tomillo et al.
2008). World-wide and in Canada, turtle populations
are in decline (Gibbons et a/. 2000); thus, it is important
to understand all possible threats, including the ecology
and behaviour of nest predators, to increase the effec-
tiveness of management and recovery actions.

Many studies of turtle nest depredation report that
most depredation events occur within the first week
post-oviposition (Tinkle et a/. 1981; Christens and Bider
1987; Congdon er al. 1983, 1987; Robinson and Bider
1988; Burke et al. 2005; Strickland er a/. 2010; Geller
2012; Wirsing et al. 2012; Holcomb and Carr 2013).
For example, Congdon et al. (1983) found that 87% of
Blanding’s Turtle (Emydoidea blandingii) nest depre-
dation occurred within the first 5 days post-oviposition.
Congdon et al. (1987) found that of depredated Snap-
ping Turtle (Chelydra serpentina) nests, 59% were
depredated within the first 24 h, 73% of these nests were
depredated within the first 6 days, and 100% by the
324 day post-oviposition. Robinson and Bider (1988)
found that 57% of depredation events occurred within
72 h of nest construction, and 87% occurred within 5
days post-oviposition. Similarly, Desroches and Picard
(2007) found that 80% of nests were depredated in the
first week post-oviposition. Holcomb and Carr (2013)
found that 86% of Alligator Snapping Turtle (Mac-
rochelys temminckii) nests were depredated within the
first 24 h, and, on average, the time until depredation
was 19h.

The high peak of depredation immediately post-
oviposition is thought to occur because cues alerting
predators to the presence of a nest are most prominent
at this time. These cues might include olfactory stimult,
such as the scent of oviposition fluid (Legler 1954;
Congdon er al. 1983; Spencer 2002), and visual cues,
such as the presence of a female turtle (Congdon er al.
1987; Eckrich and Owens 1995) and soil disturbance

ivy

180

(Strickland ef al. 2010; Spencer 2002). Acceptance
of this evidence has led some researchers examining
cues for nest depredation (e.g., Marchand and Livaitis
2004; Strickland e¢ al. 2010) to base their methodology
on the premise that depredation of turtle nests is
restricted to the first week post-oviposition. For only
2 weeks, Marchand and Livaitis (2004) monitored fake
nests constructed to investigate depredation, and Strick-
land et al. (2010) monitored nests for 2 days post-
oviposition. Yet, is an early peak the only temporal pat-
tern of turtle nest depredation reported in the literature?

Although most studies of nest depredation report
that it occurs within a short time post-oviposition, a
few, largely neglected, studies document substantial
levels of nest depredation throughout or late in the
incubation period. Snow (1982) found that 55% of
depredated Painted Turtle (Chrysemys picta) nests were
older than 3 days; however, all of the nests were still
depredated within the first month of incubation. Brooks
et al. (1992) found that all Wood Turtle (G/yptemys
insculpta) nest depredation occurred 9 weeks after the
last nest was laid. Burger (1977) found that the risk of
depredation of Diamond-backed Terrapin (Malaclemys
terrapin) nests did not decrease over the course of
incubation; instead, nest depredation was significantly
higher 60—90 days post-oviposition (75% of remain-
ing nests depredated) than 1-30 days and 30-60 days
post-oviposition (27% and 20% of nests within those
periods, respectively). Gillingwater (2002) observed
nest predation throughout the incubation period and,
in some cases, Northern Map Turtle (Graptemys geo-
graphica) nests were depredated the following spring
after hatchlings had overwintered in the nest. Some re-
ports document bimodal predator activity, with peaks
around nesting and hatching. For example, Congdon
et al. (1983) found that a few nests (6%) were depre-
dated at the end of the incubation period during hatch-
ling emergence and, similarly, Desroches and Picard
(2007) found that 5% of freshwater turtle nests were
depredated during hatchling emergence. In these stud-
ies, because of a large peak in nest depredation imme-
diately post-oviposition, there may not have been many
nests left to depredate in the fall. Also, nest debris may
make nest-searching by predators more challenging.

Nevertheless, ‘predators often appear to cue in on
nests later in incubation. During fieldwork for a 41-
year study of the ecology of Snapping Turtles and a
35-year study of Midland Painted Turtles (Chrysemys
picta marginata) based out of the Wildlife Research
Station (WRS, 45°35'N, 78°30'W) in Algonquin Provin-
cial Park, Ontario (R. J. Brooks, University of Guelph
and J. D. Litzgus, Laurentian University), field tech-
nicians observed that nest depredation was occurring
throughout incubation. These observations and our sub-
sequent review of the literature led to our research ques-
tion: do predators detect nest cues that persist long
after oviposition and lead to later occurrences of nest
depredation?

THE CANADIAN FIELD-NATURALIST

Vol. 128

In the long-term Algonquin Park study, eggs are re-
moved from the nest cavity less than 4 h post-oviposi-
tion, measured, and reburied within 24 h (Samson ef
al. 2007). This study method is common in turtle re-
search, management, and conservation programs glob-
ally; thus, depredation patterns documented in our
study may be representative of what is occurring under
these circumstances. This method also provides the
unique opportunity to examine the temporal pattern
of nest depredation that occurs when fresh cues left
by the mother are reduced. The aim of our project was
two-fold: first, to quantify the frequency of nest depre-
dation throughout incubation for Snapping and Mid-
land Painted Turtles when nest cues are reduced imme-
diately post-oviposition and, second, to compare this
temporal pattern to patterns reported previously to
determine if nest cues and depredation peaks are pres-
ent later in incubation.

Study Area

The study took place along the Highway 60 corridor
on the west side of Algonquin Provincial Park, Ontario,
Canada. The study area is within the Algonquin—Lake
Nipissing ecoregion, which is a rugged landscape
underlain by Precambrian Shield outcrops (Ontario
Ministry of Natural Resources 1998). Forest cover
dominates, with predominantly upland mixed forests
of sugar maple (Acer saccharum), yellow birch (Betula
alleghaniensis), and eastern hemlock (Tsuga canaden-
sis) and lowland sites with spruce (Picea spp.) and
tamarack (Larix laricina) (Hughes 2003). Field sam-
pling was concentrated at two sites: the WRS and
Arowhon. The WRS is within the North Madawaska
watershed and nesting sites in this area vary from nat-
ural sand dunes beside lakes to gravel embankments
along access roads and Highway 60. At the Arowhon
site, nesting occurs on a decommissioned rail-line that
is used as a public hiking trail (Mizzy Lake Trail:
Schwarzkopf and Brooks 1985). Elevations on the west
side of Algonquin Park (370 — 570 m above sea level)
are higher than the surrounding landscape and expe-
rience a cool and wet climate (Ontario Ministry of
Natural Resources 1998); this climate is reflective of
the northern range limits of both turtle species.

Methods

From 30 May to 4 July 2011, Snapping and Mid-
land Painted Turtle nesting was monitored daily. Nest
sites were monitored visually by researchers on foot
and using binoculars. Monitoring occurred from dawn
(about 0500) to about 1000, and from just before to
dusk (about 1700) until after dark, as long as nesting
activity was occurring, to capture the time frames in
which Snapping and Painted Turtles experience peak
nesting (Ernst and Lovich 2009). The nesting activity
of each female was recorded as it occurred. Nest loca-
tions were marked with metal stakes and flagging tape.
Nests were carefully excavated, maintaining the eggs

2014

in the same order and orientation in which they were
found (Samson et al. 2007). After the eggs were re-
moved, the nest cavities were filled with soil excavated
during egg removal. The eggs were taken to a field
laboratory to record clutch data for the long-term study
(egg data were not used in the current study). Eggs
were reburied in their original nest chambers within
24 h post-oviposition, before the vitelline membrane
adhered to the inner shell surface and formed a “white
spot” on the top of the egg (Yntema 1964; Rafferty and
Reina 2012), thus ensuring no trauma to developing
embryos (Samson ef al. 2007). We cannot discount the

possibility that removal, processing, and reburial of

eggs by researchers in the first 24 h affected depreda-
tion rates, especially at the start of incubation. Although
we present data for the first 72 h post-oviposition, we
are aware they may not be indicative of depredation
rates on nests from which eggs were not removed (Con-
gdon et al. 1983; Tinkle et a/. 1981; Christens and Bider
1987; Burke et a/. 2005; Strickland er a/. 2010; Geller
2012; Wirsing ef al. 2012).

RILEY AND LITZGUS: TURTLE NEST PREDATION ls]

Each nest was randomly assigned to a caging treat-
ment (above ground, below ground, or wooden-sided
cage, or uncaged control) and cages were deployed
when the eggs were reburied (69 Midland Painted Tur-
tle and 52 Snapping Turtle nests). Various nest cage
types were used to test their effects on hatchling fit-
ness for another study (Riley and Litzgus 2013); here
we report only the details of predator interactions with
these nests. During the nesting season (June) and the
hatching season (September), the nests were monitored
daily for predation attempts and successful depreda-
tion events. During the remainder of the incubation
period (July and August), the nests were monitored
weekly (Burke ef a/. 1998; Kolbe and Janzen 2002)
because of logistic constraints. A depredation attempt
was recorded when substrate was cleared away from
around the nest or nest cage and/or the nest cage was
dug up but the predator did not reach the nest cham-
ber (Figure 1A). A successful depredation event was
recorded when eggs were dug up and/or eaten (Fig-
ure 1B). If predator tracks and scat were discernible

i set caoe with substrate cleared away from the sides indicating a depredation attempt. B. Suc-
FIGURE 1. A. An above-ground nest cage Ww ith substrate cleat \ '

cessful depred

ation of a nest protected by a wooden-sided cage indicated by the excavated nest chamber and eggshell
c 2 : - te

6 nts. C. A trail camera photograph of a Red Fox (Vulpes vulpes) interacting w ith a nest outfitted with an above-
ragments. C. all Cé ‘ 5 : : Fe Dosa R Dare ae Rees p

ee nd cage. All photographs were taken during summer 2011 in Algonquin Prov incial Park, Ontario, Canada.

grou age. ee 9

182

around a nest, the predator species was identified and
recorded. To gauge the predator assemblage patrolling
the nesting sites and to capture interactions between
predators and nest cages, four trail cameras (119456C,
Bushnell Corporation, Overland Park, Kansas, USA)
were set-up from | July to | October 2011 at four nest
sites.

The numbers of depredation attempts and success-
ful depredation events were combined for analysis to
create a total number of depredation interactions. This
total did not differ among cage types or from the un-
caged control nests over the incubation period (Riley
and Litzgus 2013). It was assumed that if the nest
cages were not protecting the clutches, a depredation
attempt would have been successful. If a nest was sub-
ject to multiple depredation attempts (i.e., a predator
targeted the same nest multiple times), only the first
attempt was included in the dataset.

We quantified predator interactions with nests in two
ways. First, we used a non-parametric product-limit
survival analysis (Kaplan and Meir 1958; Engeman
et al. 2006). The survival time equaled the time (days)
from oviposition to the first predator interaction. When
eggs hatched, those nests were considered ‘“‘censored”
after the time of that event, and thus removed from the
analysis. A Mann—Whitney—Wilcoxon test was used to
compare the survival curves between species. Second,
we undertook a more detailed analysis of the specific
timing of the depredation interactions during incuba-
tion. This part of the analysis included only nests that
were subject to depredation interactions during incu-
bation (Snapping Turtle, 29/52 nests; Midland Paint-
ed Turtle, 28/69 nests). For each species, the number
of depredation interactions was totaled for each week
of incubation (29 May to | October 2011). This num-
ber was divided by the total number of depredation
interactions over the incubation period and multiplied
by 100 to calculate the percentage (i.e., frequency) of
depredation interactions that occurred weekly. For each
species, this observed distribution of depredation fre-

THE CANADIAN FIELD-NATURALIST

Vol. 128

quency was compared with an expected distribution
based on previous quantified reports of depredation in
the literature, using a Kolmogorov-Smirnov goodness-
of-fit test. From all literature sources that contained
usable data, we averaged the reported data to generate
an expected distribution of depredation interactions by
week post-oviposition (Table 1). Also, using a Kol-
mogorov—Smirnoy goodness-of-fit test and our data,
we compared the distribution of weekly depredation
interactions post-oviposition for Snapping Turtles with
that for Midland Painted Turtles to determine whether
there were differences in pattern. Depredation peaks
were identified as weeks with > 10% nest depredation.
A significance level of a = 0.05 was used for all sta-
tistical tests. Statistical analyses were performed using
R (R Foundation for Statistical Computing, Vienna,
Austria).

For predator interactions where tracks or scat were
identified, the number of interactions observed for each
predator species was summarized monthly. Similarly,
the number of photographs of each predator species
taken by the trail cameras was summarized monthly
to elucidate temporal patterns of predator presence at
the nest sites over the course of incubation.

Results

For both turtle species, the probability of nest sur-
vival decreased steadily with number of days post-
oviposition (Figure 2). The survival curves did not dif-
fer between species (W, = 365, P= 0.11). In 2011, the
first Snapping Turtle nest was found on 7 June, and
the last nest was found on 24 June. The observed fre-
quency of weekly predator interactions with Snapping
Turtle nests over the incubation period was different
than the expected distribution of depredation interac-
tions post-oviposition (D = 0.63, P< 0.01; Figure 3A).
For Midland Painted Turtles, the first nest was found
on 2 June and the last was found on 2 July. The ob-
served frequency of weekly predator interactions with
Midland Painted Turtle nests over the incubation period

TABLE 1. Rates of turtle nest predation over the course of incubation reported in previously-published studies. The means of
these rates were used to create an “expected distribution” of depredation frequencies for comparison with our observed fre-
quencies for Snapping Turtles and Midland Painted Turtles in Algonquin Provincial Park using a Kolmogorov—Smirnov
goodness-of-fit test. For the purposes of the test, the mean depredation frequency estimated for weeks 4+ (3.7%) was split

over weeks 4—16 (0.3% each week).

To

Depredation frequency (%)

Study Species Week | Week 2 Week 3 Week 4+
Congdon et al. 1983 Blanding’s Turtle 87 5) 7 4
Christens and Bider 1987 Painted Turtle 86 0 0 14
Burke et al. 2005 Diamond-backed Terrapin 100 0 0 0
Wirsing ef al. 2012 Snapping Turtle 98 | 0 l
Painted Turtle 98 | 0 l
Geller 2012 Map Turtle spp. 90 7 0 3
Snow 1982 Painted Turtle 64 21 12 3

Mean expected distribution used
in Kolmogorov-Smirnov test

2014 RILEY AND LITZGUS: TURTLE NEST PREDATION 183

8
=
2
|
2)
i)
2
5
©
2
° H
Se ‘
eo '
_| A. Snapping Turtle
0 20 40 60 80 100 120
Days after oviposition
G
2
2
3
7)
ro
2
a + |
P (o)
g
o
N
ae
2 _| B. Midland Painted Turtle
[———sn: aa ot
0 20 40 60 80 100 120

Days after oviposition

FIGURE 2. Survival curves (solid lines) for (A) Snapping Turtle
(Chelydra serpentina) and (B) Midland Painted Turtle
(Chrysemys picta marginata) nests over the incuba-
tion period in Algonquin Provincial Park. Vertical tick
marks indicate when eggs hatched and their nest was
“censored” from the analysis. The grey area represents
the period when hatchlings were emerging from the
nests. Dashed lines indicate standard errors.

was also different from the expected distribution (D
= 0,63, P< 0.01; Figure 3B).

For Snapping Turtle nests, 17% of predator inter-
actions occurred in the first week post-oviposition.
Another peak of predator interactions (weekly interac-
tions ranging from 10% to 17% of the total depreda-
tion interactions) occurred from weeks 10 to 14 (64 to

A. Snapping Turtle
( Expected

@ Observed

Predator interactions per
week (%)

_. os..,1) ee

D2 8S 4 20 / 88) 10 B12 is 14 5 AG

Weeks after oviposition

B. Midland Painted Turtle

@ Expected
@ Observed

Predator interactions per
week (%)
>
oO

t- ia a i ee |
120394 5G 7 BOLO ep S415. 16
Weeks after oviposition

FiGureE 3. For (A) Snapping Turtle (Chelydra serpentina)
and (B) Midland Painted Turtle (Chrysemys picta
marginata) nests in Algonquin Provincial Park, ob-
served weekly frequencies of predator interactions
over the incubation period (29 May to 17 September
2011) were significantly different from those expect-
ed based on previous reports (see Table 1| for list of
studies). The grey area represents the period when
hatchlings were emerging from the nests.

91 days) post-oviposition. Predator interactions oc-
curred throughout the incubation period up to 105 days
post-oviposition. For Midland Painted Turtle nests,
14% of predator interactions occurred in the first week
post-oviposition. Another spike in predator interactions
occurred 3 and 4 weeks post-oviposition, when 18%
of total depredation interactions occurred each week.
Another peak in depredation occurred at 11 weeks (71—
77 days) post-oviposition, when 14% of total depre-
dation interactions occurred. We also observed elevat-
ed levels of predation in week 12 (7% of total depre-
dation interactions). Similar to Snapping Turtle nests,
predator interactions with Midland Painted Turtle nests
occurred throughout incubation until 109 days post-
oviposition. The number of predator interactions per
week over the incubation period did not differ between
species (D = 0.31, P =0,A2).

Six predator species were identified interacting with
nests and were present at the study sites throughout

184

incubation: Red Fox (Vulpes vulpes), Eastern Wolf
(Canis lycaon), Raccoon (Procyon lotor), American
Crow (Corvus brachyrhynchos), Common Raven
(Corvus corax), and Wild Turkey (Meleagris gallopa-
vo) (Figure 4). The number of photographs of preda-
tors at nest sites captured by the trail cameras was high-

at A. Visually monitored nests
2
@ Wild Turkey
» 10 CO) Raven & Crow
re)
= E3 Raccoon
i ©]
2 6 @ Red Fox
Q.
ao)
9 6
£
2
4
2
0
June July
90 .
B. Trail camera data
80
@ Wild Turkey
70 O Raven & Crow
g 60 M Raccoon
= & Eastern Wolf
a= (50
a @ Red Fox
ie?)
= 40
=|
2
30
20
10
vialaen
May June July

THE CANADIAN FIELD-NATURALIST

Vol. 128

est in August and September; specifically, the number
of photographs of canid (Red Fox and Eastern Wolf)
predators increased during this time. Of the predator
interactions for which species were identified, Red Fox-
es were the most common predators of nests from July
until the end of incubation.

August September

August

September October

FIGURI 4. Assessment of the predator assemblage at Snapping Turtle (Chelydra serpentina) and Midland Painted Turtle
(Chrysemys picta marginata) nesting sites in Algonquin Provincial Park by (A) number of predators identified from
tracks and scat interacting with nests and (B) number of photographs taken by trail cameras of each predator species

by month over the incubation period.

2014

Discussion

We observed nest predation throughout incubation,
and there were peaks in depredation (one for Snap-
ping Turtles and two for Midland Painted Turtles) more
than a week after oviposition. Our observed patterns
of nest predation were atypical compared to those gen-
erally reported in the literature. Only a handful of oth-
er published studies have reported nest predation
throughout or late in incubation (Snow 1982; Burger
1977). Also, the presence of predators at nest sites in
our study, particularly canid species, increased later in
incubation (August and September). These later peaks
in depredation may be a result of a larger predator pop-
ulation at this time of year because canid pups begin
to hunt independently in the fall (C. Callaghan pers.
comm.), and because predators are somehow cueing to
nests at hatching. Wild dogs have been documented to
target Leatherback Sea Turtle (Dermochelys coriacea)
and Green Sea Turtle (Chelonia mydas) nests during
hatching and to dig up nests and prey on hatchlings
before their emergence from the nest (Spotila 2011).
Burger (1977) found that crows and gulls depredated
terrapin nests only during oviposition, whereas mam-
mals (raccoons and foxes) depredated nests at a low rate
during oviposition and at a high rate during hatching.
Also, mammalian predators depredated Wood Turtle
(Glyptemys insculpta) nests 9 weeks post-oviposition
(Brooks et a/. 1992). In our study, removing eggs from
nests in the first 24 h post-oviposition (which may
have reduced early-incubation nest depredation) pro-
vided an opportunity to investigate depredation in the
later stages of incubation.

It appears that cues to nests exist and can be detected
by predators throughout egg incubation, but what are
these cues? The most obvious answer may be that the
predator-exclusion cages around several of the nests
monitored in our study may have served as visual cues
to the nests’ presence. However, our study also includ-
ed nests with no cages and nests protected by cages
that did not extend above ground (Riley and Litzgus
2013) and, therefore, did not provide a visual cue. In-
deed, predation interactions did not differ among the
nest caging treatments, including controls with no cages
(Riley and Litzgus 2013); thus, it does not appear that
predators were using nest cages as visual cues to a food
source. Other researchers have also found that mark-
ing nests did not increase depredation rates (Burke ef
al. 2005; Strickland et a/. 2010). In contrast, raccoons,
coyotes, and Corvus sp. have been found to use nest
cages and nest markers, respectively, as visual cues
for nests (Mroziak et a/. 2000; Rollinson and Brooks
2007; S. D. Gillingwater, pers. comm. ). This is most
likely a learned response (Rollinson and Brooks 2007),
and it appears that, at our study site, predators have not
yet learned to associate nest cages, which were present
for only 2 summers, with the presence of turtle eggs.

A few other studies that examined unprotected turtle
nests also found that nest predation occurred late in

RILEY AND LITZGUS: TURTLE NEST PREDATION

incubation (Snow 1982; Burger 1977; Brooks et al.
1992); thus, a natural cue must be attracting predators
to nests at that time. For Midland Painted Turtles, the
depredation peak at 3-4 weeks post-oviposition could
be associated with predators returning to the nesting
sites when these turtles lay their second clutches (ap-
proximately 2 weeks after the first clutch is deposited
at our study sites; Brooks e/ a/., unpublished data). The
return of nesting female turtles may present a visual cue
for predators. The presence of hatchlings at a nesting
site may also act as a visual cue for predators later on
in incubation and trigger further searching and preda-
tion of nests. At Rondeau Provincial Park and Long
Point, Ontario, raccoons and coyotes have been ob-
served following the tracks of early-emerging turtles
back to their natal nest and consuming the remaining
eggs and young (S. D. Gillingwater, pers. comm.).

How else would predators know where unmarked
turtle nests are located? Embryonic fluids released dur-
ing hatching could serve as olfactory cues to attract
predators to nests in August and September. In addi-
tion, rotting undeveloped or unfertilized eggs may pro-
duce olfactory cues. At our study sites, canid presence
increased in August and September, and this group of
animals is well-known for its outstanding olfactory
abilities (Spotila 2011). Another possibility for nest
detection may be auditory cues. Some turtle hatchlings
have recently been reported to vocalize within the nest
cavity after hatching (Ferrara ef a/. 2013).

In our study, the later peak in predator interactions
with nests was found at 64-94 and 71-77 days post-
oviposition for Snapping and Painted Turtles, respec-
tively. These peaks of depredation precede the begin-
ning of hatchling emergence for Snapping Turtles (77
days post-emergence; Riley ef a/. unpublished data)
by 13 days, and for Midland Painted Turtles (74 days
post-emergence; Riley e¢ al. unpublished data) by 3
days. Thus, these peaks could correspond to the release
of embryonic fluids (olfactory cues) and potential
vocalization by hatchlings (auditory cues). The study
by Brooks et al. (1992) lends support to this idea, as
mammalian predators began depredating nests at their
study site 9 weeks after the last nest was completed,
which corresponds to the hatching period and hatch-
ling emergence for Wood Turtles (Ernst and Lovich
2009). Although this is a convincing association, it is
impossible to correlate definitively the later peak in
predator interactions with the hatching period from
our study alone, as it is unknown when the eggs were
hatching. Hatchlings often remain in the nest chamber
after hatching, sometimes emerging 1—9 days later
(Burger 1976; Christens 1990) or, in the case of Painted
Turtles, in the spring after overwintering in the nest
(Storey ef al. 1988; Costanzo and Lee 1995: Costanzo
et al. 2000: Packard and Packard 2003, 2004). Future
work should examine olfactory, auditory, and visual
cues at nests throughout incubation to determine wheth-
er the presence or types of cues change with time.

186

In our study, given that we potentially changed the
nest cues during the first 24 h post-oviposition, we
were able to measure peaks in nest depredation later
in incubation for two freshwater turtle species. For
Snapping Turtles, there was a later peak in nest depre-
dation 10-14 weeks post-oviposition, and, for Midland
Painted Turtles, there were peaks in nest depredation
during weeks 3, 4, and 11 post-oviposition. There was
also an increase in the number of canid predators at
the study sites during these later peaks in incubation,
which corroborates the findings of other studies in
which mammal predator presence at nest sites increased
before hatchling emergence from nests (Spotila 2011:
Burger 1977). We report that there were comparative-
ly low peaks in nest predation during the first week
post-oviposition: only 17% and 14% of depredation
interactions with Snapping and Midland Painted Tur-
tle nests, respectively, occurred at this time. Some re-
searchers have observed similar trends at other study
sites, with nest predation occurring both during nesting
and hatchling emergence (Burger 1977; Spotila 2011;
Congdon ef al. 1983; Desroches and Picard 2007). At
various sites, differences in predator species densities
and interspecific differences in the cues used by pred-
ators to find nests (i.e., avian species using visual cues,
and mammal species using visual and olfactory cues)
may be responsible for geographic variation in depre-
dation frequency over incubation. Depredation frequen-
cy may also depend on individual predator behaviours
and how they vary among sites. More research is need-
ed to examine the effects of predator species and pred-
ator behaviour on temporal patterns of nest predation.

Management of at-risk turtles throughout North
America commonly includes the use of cages to pro-
tect nests from subsidized mesopredators (Seburn
2007; Riley and Litzgus 2013). Some strategies sug-
gest removal of nest cages halfway through incuba-
tion, as it is thought that depredation will not occur dur-
ing the last half of incubation. In a study by Rahman
and Burke (2010), nest protectors were removed from
nests after 21-25 days of incubation, and, in one study
area (out of three), depredation significantly increased
in the 11 nights after nest protector removal. Engeman
et al. (2006) also found that after removing nest cages
from sea turtle nests, depredation rates increased sub-
stantially. Soil disturbance during nest cage removal
may provide a cue to attract predators (Rahman and
Burke 2010). An alternative to removing nest cages
partway through incubation is including an opportu-
nity for turtles to escape from the cage (e.g., escape
hatches or holes). Our study, as well as others that doc-
ument nest depredation before hatchling emergence,
indicates that turtle nests can be at risk of depredation
throughout the full incubation period. Thus, it is impor-
tant to understand and study depredation timing at each
location, as it may be site or predator specific, to gauge
the effectiveness of a particular nest protection method
for that site. In some areas, species recovery may re-

THE CANADIAN FIELD-NATURALIST

Vol. 128

quire that nest protection measures remain in place
throughout incubation and extend to hatchling emer-
gence.

Acknowledgements

All work was carried out in compliance with the
Canadian Council for Animal Care guidelines and
under approved Laurentian University Animal Care
Committee protocol 2008-12-02. All work was author-
ized by permits from the Ontario Ministry of Natural
Resources (OMNR). Financial support for this work
was provided by the Natural Sciences and Engineer-
ing Research Council, the Canadian Wildlife Federa-
tion, the OMNR’s Species at Risk Stewardship Fund,
the Toronto Zoo, and Laurentian University. In-kind
contributions were provided by the Wildlife Research
Station (WRS) in Algonquin Provincial Park and the
University of Guelph. We thank R. J. Brooks for allow-
ing access to his long-term study sites. Thanks also to
the following people who assisted with fieldwork: K.
Hall, M. Keevil, H. McCurdy-Adams, P. Moldowan,
L. Monck-Whipp and staff and volunteers from the
WRS. Thanks to J. Baxter-Gilbert, C. Davy, P. Moldo-
wan, S. Gillingwater, and an anonymous reviewer for
their helpful comments on the manuscript.

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Notes

A Gray Wolf (Canis lupus) Delivers Live Prey to a Pup

L. DAvID MECH

U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711—37th Street S.E.. Jamestown, North Dakota 58401-

TaT ULS.A.

Mailing address: 1920 Fitch Avenue, University of Minnesota, St. Paul, Minnesota 55108 USA; email: mechx002@umn.edu

Mech, L. David. 2014. A Gray Wolf (Canis lupus) delivers live prey to a pup. Canadian Field-Naturalist 128(2): 189-190.

A two-year-old sibling Gray Wolf (Canis lupus) carefully captured an Arctic Hare (Lepus arcticus) leveret alive on Ellesmere
Island, Nunavut, Canada, and delivered it alive to a pup 28-33 days old. This appears to be the first observation of a Gray Wolf

delivering live prey to a pup.

Key Words: Canis lupus; Gray Wolf; Arctic Hare; Lepus arcticus; learning; feeding; Ellesmere Island; Nunavut

The capacity to hunt and kill is innate in canids, as
anyone who has raised a domestic dog (Canis lupus
familiaris) can attest (Fox 1969). However, because
of the difficulty of observing young free-ranging car-
nivores hunting and killing, information about the role
of learning in helping fine-tune these behaviors is still
developing (Caro and Hauser 1992; Thorton and Rai-
hani 2008), and I know of no such information on Gray
Wolves (Canis lupus).

Several felid species bring live prey to kittens (Kitch-
ener 1999). Meerkats (Suricata suricatta) feed disabled
scorpions (of the genera Parabuthus and Opistoph-
thalamus) to pups (Thornton and McAuliffe 2006). Bat-
eared Foxes (Otocyon megalotis) bring live inverte-
brates to their young (Nel 1999; Pauw 2000), and Red
Foxes (Vulpes vulpes) allow cubs to capture live earth-
worms (MacDonald 1980). The phenomenon is dis-
cussed and the literature is summarized in Caro and
Hauser (1992) and Thornton and Raihani (2008).

In Gray Wolves, the canine teeth are usually fully
emerged by about 7 months of age (Van Ballenberghe
and Mech 1975), and we know that by winter, when
young-of-the year are 7-10 months old, they accom-
pany adults and participate in the hunting and killing
of prey (MacNulty et a/. 2009). Even without parents
present, 8- or 9-month-old Gray Wolves can kill White-
tailed Deer (Odocoileus virginianus) (M. Jimenez, per-
sonal communication).

Young Gray Wolves accompanying adults probably
learn to perfect their killing technique by observing the
adults during the actual attacking and killing, and prob-
ably by imitating them. Yearling Gray Wolves seen up
close while helping their parents kill a Muskox (Ovi-
hos moschatus) calf bit the calf as close as possible to
where their parents were biting it on the head (Mech
1988, page 78), even though no Gray Wolf was biting
any other part of the body and the head holds were not
lethal.

Although the basic killing tendency is innate in car-
nivores, gaining experience at an early age would be
adaptive because prey is often so difficult to find, catch,
and kill that any advantage at any step in the process
would promote survival. Thus the following observa-
tion of a two-year-old sibling Gray Wolf providing a
pup 28—33 days old (the pup’s ears had only begun to
stand upright 1—2 days before) with an opportunity to
handle a live Arctic Hare (Lepus arcticus) is informa-
tive. I made the observation on 6 July 1994 on Elles-
mere Island, Northwest Territories (now part of Nuna-
vut), Canada, where I studied human-habituated Gray
Wolves around dens close-up each summer from 1986
through 2010 (Mech 1988, 1995, 2005).

The primary prey of those Gray Wolves was Musk-
oxen and Arctic Hares, and the composition of the pack
attending the den under observation during the present
study was a breeding male and female (named Whitey),
a two-year-old female (Explorer), a two-year-old male,
and a single pup (Mech 1995). It was common for the
adults to catch and quickly kill young Arctic Hares,
and I had watched Explorer do so several times both as
a yearling and as a two-year-old (Mech ef a/., in press).

Following are my observations as edited from my
field notes:

6 July 1994, 1534— left the north side of the den
ridge and started toward a hill to the east. Explorer fol-
lowed me and, when 150-250 m from the den, sur-
prised an Arctic Hare leveret about 20 cm long which
jumped up and fled. Explorer chased it for 4 minutes
back and forth along a creek bed and valley, catching it
several times with her paws, grasping it with her mouth,
shaking it, and dropping it. She could have killed it
several times but did not. Each time that she dropped
the hare it ran again, and she had to re-catch it. Finally
she secured it alive.

189

190

Just as Explorer caught the leveret, Whitey came
running down from the den (around 300 m away)
and chased Explorer trying to snatch the Arctic Hare.
Whitey was unable to catch Explorer, and Explorer
took the Arctic Hare to the den. My assistant observing
from another vantage point saw Explorer carry the live
leveret by the nape of the neck toward the den. She
dropped it on the way and re-caught it, and then deliv-
ered it alive to the pup. The pup was very interested
in the leveret and tried to eat it. Explorer eventually
grabbed the leveret from the pup, and she and the pup
chewed it. At some point during this process the leveret
died.

The obvious question about this observation is why
Explorer spent so much time and effort keeping the
leveret alive. Explorer found the leveret close to the
den, and the leveret was small. Both circumstances
were favorable for the live delivery. Delivering a small
live prey animal to a pup would serve to introduce the
pup to live prey when the prey is helpless, and that
would allow the pup to learn that the active animal rep-
resents food. An alternative explanation suggested by
a reviewer is that “the juvenile rabbit triggered sever-
al incomplete responses, with behavioral elements of
(a) prey killing, (b) food delivery, and (c) pup carry-
ing.” It was not clear whether the pup or Explorer actu-
ally killed the leveret, but the killing took place some
time soon after the pup encountered the animal alive.
Simple learning would have allowed the pup to asso-
ciate pawing the live leveret with feeding.

To my knowledge this is the first observation of a
Gray Wolf delivering live prey to a pup. Such behav-
iors might not be common in Gray Wolves, because
this incident is the only time I have seen it during 13
summers observing Gray Wolves and dens in the area
for 2-6 weeks each year. On the other hand, even
though I have seen Gray Wolves kill Arctic Hares about
25 times, I have never seen a Gray Wolf capture a lev-
eret so close to a den before.

This observation fits the definition of opportunistic
teaching (Caro and Hausner 1992), in that the two-year-
old Gray Wolf modified her behavior in the presence of
a pup with no killing experience without an immediate
benefit to herself and provided the pup with experience,
thus helping the pup improve its basic predatory skill.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Acknowledgements

This study was funded by the U.S. Department of
the Interior. I also thank Environment Canada and the
Polar Continental Shelf Project for logistical support;
Craig Johnson for his assistance with observations; and
Dan Stahler and two anonymous reviewers for offer-
ing helpful suggestions to improve the manuscript.

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Thornton, A., and K. McAuliffe. 2006. Teaching in wild
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Received 8 August 2013
Accepted 8 November 2013

2014

NOTES 19]

Assessing Capture Success of Small Mammals Due to Trap Orientation

in Field—Forest Edge Habitat

/ 2 . 2 3 aS
DANIEL M. WoLcotrt,!':? MADISON R. ACKERMAN,! and MICHAEL L. KENNEDY!

Ecological Research Center, The University of Memphis, Memphis, Tennessee 38152 USA

“Corresponding author: dmwS5(@txstate.edu

due to trap orientation in field—forest edge habitat. Canadian Field-Naturalist 128(2): 191-194,

The prediction that trap orientation would not affect the likelihood of capturing small-nonvolant mammals in field—forest
edge habitat was tested during late May and early June 2010 at 3 locations in western Tennessee. Traps were placed in pairs
along transects in edge habitats with the orientation of one trap facing outward, toward the field, and the other oriented
inward, toward the forest. Results reflected no differential capture success due to trap orientation among ages, sexes, species,
or locations. This finding should facilitate the inventorying and monitoring of small mammals in an abundant and potentially

species-rich habitat type found in many terrestrial regions.

Key Words: capture success; edge habitat; Sherman live trap; small mammal monitoring; trap success; trap susceptibility;

trap orientation

Introduction

Adjoining field and forest habitats (field—forest edge)
are common throughout many ecoregions. Generally,
variety and abundance of life are thought to be great-
est in and about edges (Smith and Smith 2012). How-
ever, the intensity, or even presence, of edge effects has
been questioned by many recent studies (reviewed in
Murcia 1995). Although field—forest edges are sites with
the potential for high biodiversity, the increase in such
sites due to habitat fragmentation raises the risk of dele-
terious effects on resident communities (Groom et al.
2006).

Field—forest edges are interesting sites for studying
aspects of biodiversity and sustainability of small mam-
mal populations. Murcia (1995) and Fagan ef al. (1999)
noted the need for investigations in edge habitats. Many
studies of natural history traits require live capture and
examination of individual animals. Choosing an effec-
tive, unbiased trapping method is important for collect-
ing reliable information on populations of small mam-
mals (Risch and Brady (1996). Wilson ef al. (1996)
described procedures for measuring and monitoring
biological diversity; however, they did not discuss ori-
entation of live traps in detail. At this time, the signif-
icance of trap orientation to capture success in field-
forest edge habitats is unknown.

The transition zone between field and forest habitats
often contains thick vegetation, which may be easier to
access from the field side. When sampling for small
mammals with live traps in many field—forest edges,
investigators must decide whether to place the traps
with opening toward the field or the forest. Given that
visibility and accessibility are greater from the field
side, traps are usually approached from this direction to
reduce the time and effort needed to check and rebait
them. The extra effort required to access a trap opening
that faces the forest is worthwhile if capture success
is enhanced but not if it is decreased or insignificant.
Therefore, the purpose of our investigation was to test

the prediction that trap orientation (opening of the trap
facing the field or the forest) has no effect on the like-
lihood of capturing small, non-volant mammals (here-
after, small mammals) in field—forest edge habitat.

Study Area

The study was conducted at three locations in west-
ern Tennessee: Ames Plantation, Meeman Biological
Station, and Shelby Farms Park (hereafter, Ames,
Meeman, and Shelby Farms, respectively). Ames was
located in Fayette and Hardeman counties (35°6.9'N,
89°12.7'W); Meeman (35°21.7'N, 90°1.1"W) and Shel-
by Farms (35°8.4'N, 89°50.2'W) were in Shelby County.
All locations represented sites with numerous anthro-
pogenic field—forest edge habitats. At Ames, fields were
either currently used for agriculture or were early suc-
cessional habitat. Forests consisted mainly of mature
bottomland hardwoods with some upland hardwood
forests (Baldwin et al. 2005). At Meeman, fields con-
sisted of maintained areas with some early succession-
al habitat, and forests comprised mature upland and bot-
tomland hardwood forests (Carver ef al. 2011). Shelby
Farms was the study area with the greatest anthro-
pogenic influence. Fields there were created by current
agricultural practices or were maintained areas or road
edges, and forests comprised upland and bottomland
hardwood forests (Wolcott et a/. 2012). Habitats exam-
ined in our investigation represented a mosaic of field—
forest edge types in both rural and urban locations.

Methods
Trapping

Our trapping protocol consisted of transect-sam-
pling using Sherman live traps (7.5 cm x 9.0 cm *
23.0 cm: H. B. Sherman Traps, Inc., Tallahassee, Flori-
da) following the method of Jones er al. (1996). Dur-
ing late May and early June 2010, traps were placed
along 54 transects in pairs with one trap facing the
forest and the other oriented toward the field. Each

transect included 30—40 total traps with pairs of traps
spaced approximately 5 m apart. Trap placement was
fully randomized throughout the study with an even
number of transects having the field-facing trap on the
left-hand side and other transects having the field-
facing trap on the right side. All traps were placed with
the door mechanism free from obstructions to ensure
efficient operation. Traps were baited with rolled oats
and checked daily.

We conducted sampling for |—3 nights along each
transect. Number of trap-nights (1.e., one trap set for
one night) was used as a metric to maintain equal com-
parisons between the two trap orientations. Species,
age, and sex of captured animals were determined. Ani-
mals were temporarily marked on the ventral surface
(lightly) with a black or blue sharpie marker (Sanford
LP, Oak Brook, Illinois) and released at the site of
capture. We followed the guidelines for the use of wild
mammals in research suggested by the American Soci-
ety of Mammalogists (Sikes et al. 2011) and the Insti-
tutional Animal Care and Use Committee of the Uni-
versity of Memphis approved our methods (IACUC
Protocol 0673).

Analysis

Logistic regression was used to assess capture suc-
cess based on trap orientation. The dependent variable
was categorized as binary: capture success of a field-
facing trap was coded as 0, and capture success of a
forest-facing trap was coded as 1. The predictor vari-
ables for sex, age, location, and species were coded as
dummy variables (see Sokal and Rohlf 2012). Ages
were categorized as juvenile, sub-adult, or adult (see
Martin et al. 2002). Reference categories were adult
for age, female for sex, Ames for location, and House
Mouse (Mus musculus) for species. Logistic regres-
sions considered possible demographic factors (sex,
age, and species) as well as locale, which could account
for differences in capture success due to trap orienta-
tion. The goodness-of-fit for each logistic regression
was assessed using a likelihood ratio test (Sokal and
Rohlf 2012). All statistical tests were performed in R
version 3.0.2 (R Foundation for Statistical Computing,
Vienna, Austria, 2012).

Results

Sampling effort (orientation of traps equally divid-
ed) was 10216 trap-nights: Ames 1596, Meeman 2280,
Shelby Farms 6340. In total, 408 individual small mam-
mals (191 males, 217 females) were captured a total of
480 times (230 males, 250 females): captures at each
location were Ames 147 (65 males, 82 females); Mee-
man 75 (38 males, 37 females); Shelby Farms 258 (127
males, 131 females). A total of six species were cap-
tured. The White-footed Mouse (Peromyscus leuco-
pus) was captured most often (n = 324: 157 males,
167 females) followed by the Hispid Cotton Rat (Sig-
modon hispidus; n = 80; 33 males, 47 females), the
Woodland Vole (Microtus pinetorum: n = 40: 19 males,

THE CANADIAN FIELD-NATURALIST

Vol. 128

21 females), the North American Deermouse (Per-
omyscus maniculatus; n = 23; 13 males, 10 females),
the House Mouse (n = 9; 6 males, 3 females), and the
Marsh Oryzomys (Oryzomys palustris; n = 4; 2 males,
2 females).

Likelihood ratio tests conducted for each model
showed that neither demographic factors nor location
significantly increased the likelihood of capture suc-
cess in a particular orientation: sex L = 0.441, df= 1,
P= 0.507; age L = 1.016, df = 1, P = 0.602; species
L=3.602, df= 1, P= 0.608; location L = 0.362, df= 1,
P=0,835).

Discussion

Our results support the prediction that trap orienta-
tion does not affect the capture success of small mam-
mals in relation to age, sex, species, or location in field—
forest edge habitat. Rana (1986) and Norton (1987)
showed that small mammals do not use areas random-
ly and that well-placed traps will enhance capture suc-
cess. Capture success for this group of mammals is also
influenced by several factors other than placement (see
Wilson et al. 1996). For example, sampling may vary
among habitat (e.g., Feldhamer et al. 1993; Schnell e¢
al. 2008). Capture rates are also known to be different
for males and females (Davis and Emlen 1956). Males
often have larger home ranges than females and, thus,
greater exposure to traps. Because of this, males are
often captured more frequently than females resulting
in sex biases in trapped samples (Buskirk and Lindstedt
1989; Poindexter et a/. 2013). Our findings suggest that
a sex bias is not present in the successful capture of
individuals at a particular orientation.

Differential capture among species may be related to
population density, with the most abundant species gen-
erally being captured in greatest numbers (see Nichols
1986; Hopkins and Kennedy 2004). As in our inves-
tigation, LaMountain (2007) reported that the White-
footed Mouse was abundant in edge and forest habitats
and the Hispid Cotton Rat was abundant in field habi-
tat in western Tennessee. Other studies have recorded
high densities of White-footed Mouse in edge (e.g.,
Adler and Wilson 1987; Manson et al. 1999) and forest
sites (e.g., Yahner 1992; Wolf and Batzli 2002). Foster
and Gaines (1991) and Brady and Slade (2001) noted
an abundance of Hispid Cotton Rat in field habitats. In
our study, we were able to capture species that are abun-
dant in each of these habitat types. Our findings demon-
Strate that there was no significant difference in captures
between the two trap orientations due to species,

Other factors are known to influence the capture of
small mammals. For example, trap type (Sealander and
James 1958; Hansson and Hoffmeyer 1973), type of
bait (Churchfield 1990), moonlight (Price ef a/. 1984).
chemical odours (Chabreck ef al. 1986, Heske 1987),
fire (Christian 1977; Gates and Tanner 1988), and
weather (Gentry ef al. 1966: Doucet and Bider 1974).
In addition, Barnett and Dutton (1995) noted the im-

2014

portance of trap position, spacing, quantity, and dura-
tion in the capture of small mammals. Because orien-
tation of the trap could be a consideration in several

of these untested factors, additional investigations of

trap orientation in relation to trapping methods could
provide new insight into procedures for studying small
mammals in various habitats.

Overall, we found no difference in capture success
due to trap orientation in field—forest edges in relation
to age, sex, species, or location. It ‘appears that the ori-
entation of a trap in field—forest edge habitat makes lit-
tle difference in the capture of small mammals. Given
this finding, we note that orienting trap openings toward
the field will likely minimize the trapping effort asso-
ciated with checking and rebaiting traps. This informa-
tion should be useful in future studies relating to the
challenging tasks of surveying and monitoring popu-
lations of small mammals.

Acknowledgements

We thank the personnel of the Ames Plantation,
Meeman Biological Station, and Shelby Farms Park
for permission to work on lands under their control.
We also thank A. Christopher Grow, as well as students
from the University of Memphis field techniques
course, who assisted in various aspects of the fieldwork
associated with our investigation.

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Received 7 November 2013
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2014 NOTES 195

Effect of Food Patch Discovery on the Number of American Crows
(Corvus brachyrhynchos) Using a Flight Lane

WILLIAM LANGLEY

Butler Community College, El Dorado, Kansas 67042 USA; email: blangley@butleree.edu

Langley, William. 2014. Effect of food patch discovery on the number of American Crows (Corvus brachyrhynchos) using a
flight lane. Canadian Field-Naturalist 128(2): 195—199,

In winter, American Crows (Corvus brachyrhynchos) move back and forth between night roosts and foraging sites along
flight lanes. If communal roosts act as information centres, we would expect more birds to use a particular flight lane after
discovery of a new food patch on that route. In this study, I investigated how the number of crows using different flight lanes
was affected by the establishment of artificial food patches, as well as how crows responded to multiple days of provisioning
and to the location of the food patch relative to the flight lane. A fter discovery of a food patch, the number of crows using the
flight lane closest to it increased, while numbers using adjacent flight lanes remained the same or decreased, particularly
when the patch was in the path of the flight lane and when food provisioning occurred for 2 consecutive days. These results

support the idea that crows using winter roosts may make use of information on food availability obtained at the roost.

Key Words: American Crow; Corvus brachyrhynchos; flight lane; winter roost; food patch; communication

Introduction

During the winter, American Crows (Corvus brachy-
rhynchos) gather in night roosting groups consisting
of thousands of birds (Kalmbach 1916; Haase 1963).
They move to and from these night roosts in flight lanes:
in the morning they radiate out to foraging areas and
in the evening congregate at pre-roosting aggregation
areas before flying to the night roost (Aldous 1944;
Moore and Switzer 1998). In some cases, crows use the
same flight lane regularly; in other cases, they change
flight lanes (Aldous 1944; Stouffer and Caccamise
1991; Caccamise ef a/. 1997). Large increases in the
number of crows using a flight lane have occurred, after
a food patch such as grain spilled from a railcar or an
un-harvested sunflower field, was discovered (Langley
1992*).

An important potential function of a communal roost
is that it can act as an information centre to facilitate
finding and exploiting new food sources (Ward and
Zahavi 1973: Weatherhead 1983). Common Ravens (C.
corax) and Hooded Crows (C. corone) that have dis-
covered a food patch recruit others at the night roost

to follow them to the food patch the next day (Marzluff

et al. 1996; Sonerud et al. 2001). Crows can also learn
by “local enhancement”, i.e., by watching other indi-
viduals and responding if a food patch is discovered
(Richner and Marclay 1991). The fact that tagged crows
at a food patch do not stay together at the pre-aggrega-
tion site or roost raises the possibility that more fluid
kinds of interactions may also occur among individuals
at a large roost (Moore and Switzer 1998). ;
The information centre hypothesis predicts that, after
discovery of a food patch on a flight lane, more birds
would use that lane subsequently. In this study, | inves-
tigated how the number of crows using a flight lane is
affected by the location of a provisioned food patch
relative to the lane and the number of days over which
it is provisioned. The approach was to present a food
patch just after a snow storm when wheat plants (777-

ticum aestivum) and un-harvested sorghum seed (Sorgh-
um bicolor), the two main food items for crows during
winter (Platt 1956), became less available.

Study Area

The study area was located in the northeast quadrant
of the area surrounding a winter roost at Wichita, Kansas,
where as many as 25000 birds a night have roosted
over the past 50 years (Langley 1999). The area forms
a rectangle of 969 km? including northeastern Sedgwick
County and the western edge of central Butler County
(Figure 1). Land use consisted of agriculture (mainly
wheat and sorghum fields), pasture, small farms, resi-
dences, and several small towns (Langley 1992*). Ob-
servations occurred from December to March, 1990—
1998.

Methods

Food patches were set out before dawn on the morn-
ing after a snowfall of at least 8 cm. A food patch con-
sisted of 65 kg of corn kernels or bakery items. Food
was spread over cleared frozen ground or on crusted
snow and directly in the path ofa flight lane and at least
10 km from the roosting site.

Three sometimes four flight lanes occurred in the
study area. I selected three with the targeted lane having
an adjacent lane on either side of it and identified the
trajectory of each during preliminary observations. For
one lane, I counted the number of birds by positioning
myself so that birds flew overhead. To ensure that all
the birds were counted, I took up a position that was
located at the eastern edge of Wichita before dawn and
counted until no birds passed overhead for 10 minutes.
A different strategy was used to count the number of
birds in the other flight lanes. First, | drove outward in
a zig-zag pattern along the trajectory of the flight lane
until no crows could be seen in flight or foraging in the
fields for two (2.56 kim?) sections (the roads border the
edges of a square-mile section). Then, I retraced the

196

THE CANADIAN FIELD-NATURALIST

ee

!

Sieg
= = cet: :
<i roe

Vol. 128

ee Se
ae mite ie
ies ¢

| Ni

FicureE 1. Northeast portion of Wichita, Kansas, showing night roosting site of American Crows (Corvus brachyrhynchos).
Flight lanes (lines) and food patches (oval) are depicted for the first experiment. Thinnest lines show the targeted flight
lane that was used in the study for only | year; a thicker line shows a lane used for 2 different years; and the thickest
line shows a lane that was used for 4 different years. Dashed lines beyond the food patches show where some crows

continued to fly and forage.

trajectory of that path counting all crows in the air or
foraging. Counting early in the morning and following
the trajectory out and back from the night roost reduced
the possibility of missing any dispersing crows. Again,
all crows were counted until none was observed for
two sections. A similar process was repeated for a third
flight lane. This counting procedure was conducted for
three consecutive mornings before a forecasted snow
event. Counting the number using a flight lane as they
flew directly overhead had to be done first and was

done this way for each flight lane on a different day so
that counts were done in similar manner for all 3 flight
lanes. The mean counts of those 3 days served as the
baseline number of crows using those flight lanes.
On the morning after establishing a food patch, |
counted the number of crows passing overhead from
a position directly in line with the food patch and
between it and the roost until no crows had arrived
from the direction of the roost for 20 minutes. After
that, I counted the number of crows using the adjacent

2014

flight lanes by tracing the trajectory from the roost and
back as was carried out for counting before establishing
a food patch. A survey of the crows at the food patch
and beyond was conducted after all 3 flight lane counts
were completed. These trials were conducted 14 times.

In a second experiment involving 8 trials, food
patches were provisioned for 2 consecutive days rather
than | day as in the first experiment. The change in the
number of crows using that flight lane was measured on
the third day. In a third experiment, a similar approach
and counting procedures were used, but the food patch-
es were offset from the flight lanes. They were placed
at a distance of 1.0-1.5 km away from the targeted
flight lane. These trials occurred 25 times.

Several non-parametric tests were used to compare
differences (Siegel 1956): a Wilcoxon T test was used
to compare the number of crows using a flight lane
before and after food provisioning; a Mann-Whitney
U test was used to compare the counts after | or 2 days
of provisioning; and a y* test was used to compare
the number of discovered food patches placed in or
to the side of a flight lane.

3000 5

2500 ~

2000

1500 ~

Mean number of crows

1000 ~

500 +

——_|——_ ———;

Se
After one-day
provisioning

Baseline 1

FIGURE

NOTES

Adjacent
before

197

Results

More crows used a flight lane after a food patch was
set out for | day: pre-food, 152 (SD 220) and post-food,
588 (SD 597) (Wilcoxon test, 7 = 8, n = 14, P< 0.02)
(Figure 2). Although in two trials the number of crows
using a flight lane decreased after the food patch was
set out compared with number of crows using the flight
lane beforehand, adverse weather (snow and fog) may
have contributed to this decrease. The change in the
number of crows using a flight lane ranged from —179
to 1992. In three of these trials, crows were observed
veering from an adjacent flight lane to the targeted
flight lane. In these cases, the flight lanes merged into
one that flew overhead at the counting location.

The number of crows using adjacent flight lanes de-
clined from 1396 (SD 1134) to 1165 (SD 894) after
the food patch was discovered (Figure 2), but the dif-
ference was not statistically significant (Wilcoxon test,
T = 42, n= 14, P= 0.66). The response of crows in
adjacent flight lanes veering and joining the targeted
flight lane passing over the food patch contributed to
the lower mean after the food patch was discovered.

Adjacentafter Baseline2 Aftertwo days

provisioning

2. Mean number (and standard deviation) of American Crows (Corvus brachyrhynchos) in target lane before (base-
li 1) and after | day of food provisioning, observed in adjacent lanes before and after food provisioning, and in the
ine d ; ?

target lane before (baseline 2) and after 2 days of food provisioning.

198

When food patches were provisioned for 2 days, the
increase in number of crows using the target lane more
than doubled from 528 (SD 625) to 1238 (SD 1140)
(Wilcoxon test, T= 0, n = 8, P< 0.01) (Figure 2). The
number of crows using a flight lane after 2-day pro-
visioning 1238 (SD 1140) was greater than the number
after 1-day provisioning 588 (SD 597) (Mann-Whitney
Untest0 S20, m= Semi =TAyde = 0502),

When food patches were offset from the flight lane,
different groups of crows discovered some of them.
On three occasions, crows from the Wichita roost dis-
covered the food patch as evidenced by the direction
from which the birds arrived (trajectory from roost), by
their early arrival (O800), and by the increase in the
number of birds in these three flight lanes from pre-
food 178 (SD 158) to post-food 291 (SD 131). These
counts were carried out as in prior experiments. Seven
other food patches were found by crows not from the
Wichita roost. These crows arrived from a different
direction than those from the Wichita roost, at later time
(1000) and in fewer numbers (1—10 birds). No other
crows were observed in the surrounding area. Of the
25 food patches set to the side of the path of flight lane,
15 were not discovered by any crows the day after
provisioning. The proportion of food patches discov-
ered by crows from the Wichita roost when they were
placed on the flight lane (12/14) was significantly great-
er than when they were offset from the flight lane (3/25)
(y° = 17.6, P< 0.001).

Discussion

The increase in the number of crows using flight
lanes where supplementary food had been placed the
previous day suggests that information obtained at the
roost enabled crows to adjust their foraging behaviour
to take advantage of the new food supply. This was
especially true when the new food patches occurred
in the path of the flight lane and were provisioned for
more than | day. These data support the hypothesis that
night roosts can act as information centres. In this study,
the roost was large (>25000 crows) and the number of
additional individuals using a flight lane exceeded 1000
on five occasions.

However, the response to discovery of food patch
was highly variable, suggesting that additional factors
may have influenced the number of crows using a par-
ticular flight lane. When a large number of crows re-
sponded to the discovery of a food patch, there were
too many to access or use the food. Furthermore, most
of these crows flew past the food patch, sometimes
more than 10 km, suggesting that some were foraging
in areas other than the food patch. Although the dis-
covery of a food patch proved to be an attraction, not
all arriving individuals necessarily made use of it.

Presumably, the use of flight lanes can be beneficial
to crows in more ways than enhancing the discovery
of food sources. Vigilance is important while foraging,
and flocks can provide an optimal tradeoff between

THE CANADIAN FIELD-NATURALIST

Vol. 128

feeding and watching for danger (Ward and Low 1997).
When competing for a carcass with a larger raptor, ag-
gregation in flocks ensures success more often for the
crows. For example, a flock of American Crows was
more successful in displacing a Red-tailed Hawk (Buteo

jamaicensis) from an animal carcass than a few indi-

viduals (Langley 2001).

During this study, crows occurred in three kinds of
associations. The most abundant were those in flocks
from the Wichita roost. Less common were groups of
2-6 birds that presumably represented territorial family
groups not associated with the roost (Knopf and Knopf
1983). A third type observed rarely and intermittently,
ranged in size from 10—100 individuals, possibly rep-
resenting non-breeding crows in the area. Groups of
non-breeding crows have been observed near urban
areas in California (Caffrey 1992).

Acknowledgements

I wish to thank Alan Maccarone for reviewing the
manuscript and the Chickadee Check-off of the Kansas
Department of Wildlife for financial support at the start
of the project.

Documents Cited (marked * in text)

Langley, W. M. 1992. Foraging behavior in winter roosting
crows in Wichita area. Non-game report. Kansas Depart-
ment of Wildlife and Parks, Pratt, Kansas. 55 pages.

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Langley, W. M. 1999. Changes in crow populations in Kansas.
Kansas Ornithological Society Bulletin 50: 35-38.

Langley, W. M. 2001. Crow and red-tailed hawk interaction
at animal carcasses. Transactions Academy of Science
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Marzluff, J. M., B. Heinrich, and C. S. Marzluff. 1996.
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Moore, J. E., and P. V. Switzer. 1998. Preroosting aggrega-
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Platt, D. 1956. Food of the crow, Corvus brachyrhynchos
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Richner, H., and C. Marclay. 1991. Evolution of avian roost-
ing behavior: a test of the information centre hypothesis

2014

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438.
Siegel, S. 1956. Non-parametric Statistics: for the Behavioral
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Sonerud, G. A., C. A. Smedshaug, and O. Brathen. 2001.
Ignorant hooded crows follow knowledgeable roost-mates
to food: support for the information centre hypothesis.
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Stouffer, P. C., and D. F. Caccamise. 1991, Roosting and
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NOTES

199

Ward, C., and B. S. Low. 1997. Predictors of vigilance for
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Ward, P., and A. Zahavi. 1973. The importance of certain
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Received 15 November 2013
Accepted 28 January 2014

200 THE CANADIAN FIELD-NATURALIST Vol. 128

Muskrat (Ondatra zibethicus) Interference with Aquatic Invertebrate
Traps

*
MICHAEL C. CAVALLARO!:3, ANSON R. MAIN!, and Curisty A. MorRISSEY!: 2

'School of Environment and Sustainability, Room 323, Kirk Hall, University of Saskatchewan, 117 Science Place, Saskatoon,
Saskatchewan S7N 5C8 Canada

"Department of Biology, W. P. Thompson Building, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan
S7N SE2 Canada

3Corresponding author: CavallaroMC 15@gmail.com

Cavallaro, Michael C., Anson R. Main, and Christy A. Morrissey. 2014. Muskrat (Ondatra zibethicus) interference with aquatic
invertebrate traps. Canadian Field-Naturalist 128(2): 200-203.

In field biology, interactions between wildlife and in situ equipment occur often. These interactions have the potential to induce
a variety of behaviours in local fauna. Here, we note the destructive behaviour exhibited by the Muskrat (Ondatra zibethicus)
following deployment of aquatic invertebrate traps for research purposes at 12 wetlands located in central Saskatchewan. Of
24 aquatic insect emergence traps used on seven wetlands in our study, 14 (58%) required recurring repairs. In addition, on
several occasions, leaf litter bags and their anchoring stakes were torn or chewed. The recurring damage took place in wetlands
with Muskrat lodges. We recommend structural modifications to aquatic invertebrate traps in wetland complexes densely inhabited
by Muskrats and other semi-aquatic rodents.

Key Words: Muskrat; Ondatra zibethicus; aquatic insect emergence traps; destructive behaviour; prairie wetlands; leaf litter bags

When conducting field studies, biologists occa-
sionally encounter unexpected obstacles arising from
natural causes, such as adverse weather, and wildlife
interference. Overcoming these obstacles can require a
substantial amount of time, energy, and resources and,
in some Cases, may result in gaps in data or a complete
loss of data. These experiences may prove to be valu-
able, and lessons learned can be applied to future exper-
imental design.

As part of a 2013 study monitoring aquatic insect
emergence and benthic macroinvertebrate communi-
ties in agricultural wetlands located near Alvena, Sas-
katchewan (52°31'0.12"N, 106°1'0.12"W), we deployed
standard aquatic insect emergence traps and leaf litter
bags (Merritt et a/. 1996; Dangles and Malmqvist 2004).
Our floating traps covered a surface area of | m? above
the water column. They consisted of a wooden frame
supporting mesh-netting sides, with a collection funnel
and flask at the top (Figure 1A). Unfortunately, the low-
er ledge of the frame at the water edge consisted of an
8-cm-wide ledge allowing Muskrats easy access for
perching or sitting.

Accounts of Muskrat biology and life history in Sas-
katchewan are well documented (e.g., Messier ef al.
1990; Virgl and Messier 1992). Muskrats’ diet consists
primarily of the roots, shoots, and rhizomes of emer-
gent hydrophytes (Virgl and Messier 1992); occasion-
ally, they will migrate to upland habitat to feed on row
crops, especially in agricultural wetlands (Bucci 2009).
Other feeding habits include the construction of feed-
ing huts or eating platforms. Built from mud and com-
pacted vegetation, these resemble Muskrat lodges, stand-
ing just above the surface of the water (Link 2005*).

In multiple instances, an individual Muskrat or a pair
moved building materials, such as mud, dead vegeta-

tion, and twigs, onto our traps (Figure 1A); this design
allowed the animals to move freely on and off the ledge.
No damage to the frame of the traps occurred, but the
netting that funneled imago insects was slightly torn.
To our knowledge, the only other account of Muskrat
interference with aquatic traps is Marcstr6m’s (1964)
description of Muskrats damaging fish traps in northern
Sweden.

We collected emergence trap samples every 3-4 days
and repaired damaged nets during these collections.
At some locations, Muskrats were directly observed
damaging a trap on multiple occasions, and this prompt-
ed modification of the traps. Subsequently, chicken
wire, 2.5-cm mesh (Cable Ben-Mor, Model #94002:
Rona, Boucherville, Quebec), was fastened around the
outside of the traps and across the bottom to protect the
netting.

We acknowledge that the addition of chicken wire
mesh structure below the floating trap could have an
influence on emerging insects, as several aquatic insect
taxa, such as Odonata, require substrate (e.g., emergent
vegetation) to achieve the final stage of metamorphosis
(Merritt et a/. 1996). Our placement of emergence traps
focused on both open water and emergent vegetation
habitats and the method for anchoring each trap was
identical. Samples collected from emergence traps —
with or without chicken wire — in open water habitats
did not contain an overabundance of taxa that require
Substrate. Also, frequent collection of samples allowed
for direct observation of potential bias by searching for
insect exuviae attached to the emergence traps. A bias
in sampling could be inferred by an overabundance of
insect exuviae from substrate emerging insects.

Several times, we observed Muskrats seeking refuge
under the emergence traps even after modification,

2014

However, the addition of chicken wire under the trap
prevented them from surfacing inside the trap, effec-
tively preventing damage (Figure 1B). Adult Muskrats
were unable to enter the trap, although a single juve-
nile Muskrat was observed moving freely over the
side of the chicken wire. This same juvenile collected
senesced wetland vegetation to create a resting area
(Figure IC). In addition to nesting materials, fresh
vegetation was also found on the traps. With a variety
of potential predators in the area (e.g., Coyotes, Canis
latrans; Red-tailed Hawks, Buteo jamaicensis: and
humans), this suggests that the floating traps provided
a safe vantage point to feed.

FIGURE |. (A) Aquatic insect emergence trap (1 m *
ledge.
trap after chicken wire was installed. In this photo,
and off the trap. (D)

NOTES 201

We assembled leaf litter bags to monitor shifts in
benthic macroinvertebrate communities throughout the
growing season. Roughly 10 g of dried leaf litter from
senesced native wetland vegetation (e.g., Broadleat
Cattail, 7ypha latifolia) were added to a mesh bag. The
bags were anchored in the sediment with |-m spruce
stakes. On multiple occasions, while removing leaf lit-
ter bags during the course of our experiment, we ob-
served teeth marks on the spruce stakes (Figure 1D).
On four instances during early spring, we found evi-
dence of Muskrats chewing open the leaf litter bags.
We are uncertain why they did this, as there was no
evidence of them feeding on the contents.

1 m) with Muskrat (Ondatra zibethicus) building material on its lower

Juvenile Muskrat seeking refuge under an aquatic insect emergence trap. (C) Aquatic insect emergence
en d the chicken wire is bent from the juvenile Muskrat moving on
Spruce stakes used to anchor the leaf litter bags showing teeth marks and evidence of chewing.

Muskrats display strong territorial behaviour during
breeding season; their aggressive behaviour reaches
a peak from late April to late May and diminishes in
early June (Beer and Meyer 1951). The frequency of
our trap repairs was highest from late May into early
June, when we suspect invasion of the territory by con-
specifics prompted destructive territorial behaviour.
Some Muskrats were noticeably agitated by our pres-
ence; individual animals were observed sitting on their
lodges and chattering their teeth or emitting a sharp,
whining noise. Mizelle (1935) recounts observations
of swimming muskrats “clapping” their fanned tails
on the surface of the water when startled. Other semi-
aquatic rodents are described as displaying defensive
behaviour that involves hissing, whining, or gnashing
their teeth toward other individuals (Leighton 1933).

In the absence of aquatic or semiaquatic plants,
overwintering Muskrats are known to consume woody
material such as tree bark (Lewis et a/. 2000). The late
spring—summer transition that occurred in 2013, our
study year, did not support wetland vegetation growth
until late May. During winter and early spring, Musk-
rats do not store food reserves and are confined to for-
aging beneath the ice (Virgl and Messier 1992). With
the lack of available food resources within the wet-
lands and upland habitat, spruce stakes placed in the
wetland flowing ice melt could serve as an easy source
of nutrients. However, the leaf litter bags were con-
structed with nylon and fibreglass mesh (0.25—1 mm),
with no apparent nutritional value. Muskrats exhibit
feeding plasticity, consuming diverse food items and
allowing them to occupy a variety of aquatic ecosys-
tems (Bucci 2009). Like most rodents, they must chew
regularly to wear their teeth adequately (Lewis et al.
2000). Dental wear in Muskrats is primarily driven by
diet and life history strategies — burrowing versus
lodging (Lewis et al. 2002). Muskrats with varying life
history strategies may require greater dental wearing
from other activities (e.g., chewing or gnawing woody
materials). The dominant plant species at each wetland
in our study area, a non-woody monocot, Broadleaf
Cattail, might not have been sufficient to achieve den-
tal wearing, which might explain why Muskrats gnawed
our stakes and leaf litter bags (Figure 1D).

The spring snow melt in 2013 caused severe flood-
ing throughout Saskatchewan. Reports indicated that,
at the height of the flooding, the South Saskatchewan
River rose 1.5 m (Water Security Agency 2013*). As
a direct effect of this flooding, Muskrats were able to
occupy new habitats and move freely throughout their
known range. Our field sites in Alvena, Saskatchewan,
were roughly 5.5 km from the South Saskatchewan
River; thus, flooding may have contributed to a high
influx of Muskrats in our study area (F. Messier, Uni-
versity of Saskatchewan Biology Department, August
2013, personal communication),

THE CANADIAN FIELD-NATURALIST

Vol. 128

Based on our experience over the season, we rec-
ommend modification of aquatic emergence traps to
include a narrower, or nonexistent, perching ledge at
the bottom. Where funds are insufficient to construct
new traps, shielding the netting and bottom of the traps
with chicken wire is effective (Figure 2). As a measure
of caution, and to be certain Muskrats cannot gain ac-
cess to the trap, chicken wire should be firmly attached
to all sides and bottom. To prevent damage to leaf lit-
ter bags, encasing them in a rigid mesh structure with
galvanized wire would fix them to a desirable location,
provide protection from Muskrats, and act as a suitable
alternative to anchoring with stakes; this strategy could
be used in both lotic and lentic freshwater systems.
Galvanized wire mesh has been used to protect the soil—
surface water interface bordering dams, dikes, canals,
and shoreline property from Muskrat burrowing and
many state agencies in the United States document
these methods in external reports (e.g., Link 2005*).
As a final level of precaution, surveys should be con-
ducted to determine whether Muskrats are present be-
fore invertebrate traps are installed in wetlands so that
modification may be made to protect the traps appro-
priately. However, because Muskrat lodges are not
always apparent, as they are typically hidden within
dense emergent vegetation, trap modifications may be
made as a preventive measure.

XX
PRR

ae QO

FiGuRE 2. Emergence trap with chicken wire fixed to sides
and bottom to prevent Muskrat (Ondatra zibethicus)
entry and damage.

In conclusion, the high Muskrat density in our study
area resulted in substantial additional work and time
spent during the field season, especially during peak
Muskrat breeding season (late April to late May). We
hope the advice in this note helps field biologists, ento-
mologists, and wetland scientists planning and exe-
cuting aquatic invertebrate experiments in habitats oc-
cupied by Muskrats or other semi-aquatic rodents,
such as beavers.

Acknowledgements

We thank T. Gallagher, K. Majewski, and A. Zahara
for field assistance. In addition, R. Clark, N. Michel.
and D. Oswin provided helpful revisions and suggest-
ed emergence trap modifications. A grant to C. A. Mor-
rissey from the Natural Sciences and Engineering Re-
search Council of Canada supported this research.

2014

Documents Cited (marked * in text)

Link, R. 2005. Living with wildlife: Muskrats. Washington
Department of Fish and Wildlife. Olympia, Washington.
Available: http://wdfww.wa.gov/living/muskrats.pdf
(accessed 4 March 2014).

Water Security Agency. 2013. Water Security Agency antici-
pating higher water levels from Alberta rainfall. Advisory
report, 20 June. Water Security Agency, Moose Jaw, Sas-
katchewan. Available: https://www.wsask.ca/About-WSA
/Advisories-/201 3/June/Water-Security-Agency-Anticipat
ing-Higher-Water-Levels-From-A Iberta-Rainfall
(accessed 4 March 2014).

Literature Cited

Beer, J. R., and R. K. Meyer. 1951. Seasonal changes in the
endocrine organs and behavior patterns of the muskrat.
Journal of Mammalogy 32(2): 173-191.

Bucci, L. A. 2009. Anthropogenic environmental change and
habitat occupancy by riparian muskrats in a midwestern
landscape. Master’s thesis. University of Illinois at Urbana-
Champaign, Urbana-Champaign, Illinois.

Dangles, O., and B. Malmqvist. 2004. Species richness-
decomposition relationships depend on species dominance.
Ecology Letters 7: 395-402.

Leighton, A. H. 1933. Notes on the relations of beavers to
one another and to the muskrat. Journal of Mammalogy
14(1): 27-35.

NOTES

203

Lewis, P. J., M. Gutierrez, and E. Johnson. 2000. Ondatra
zibethicus (Arvicolinae, Rodentia) dental microwear pat-
terns as a potential tool for palaeoenvironmental recon-
struction. Journal of Archaeological Science 27: 789-798.

Lewis, P. J., S. Richard, E. Johnson, and W. C. Conw ay.
2002. Absence of sexual dimorphism in molar morpholo-
gy of muskrats. Journal of Wildlife Management 66(4):
1189-1196.

Marestrém, V. 1964. The muskrat Ondatra zibethicus L. in
northern Sweden. Vitrevy 2: 329-407.

Merritt, R. W., V. H. Resh, and K. W. Cummins. 1996.
Design of aquatic insect studies: collecting, sampling, and
rearing procedures. Pages 12-28 in An Introduction to the
Aquatic Insects of North America. Edited by R. W. Merritt
and K. W. Cummins. Third edition. Kendall/Hunt Publish-
ing Company, Dubuque, lowa.

Messier, F., J. A.Virgl, and L. Marinelli. 1990. Density-
dependent habitat selection in Muskrats: a test of the ideal
free distribution model. Oecologia 84: 380-385.

Mizelle, J. D. 1935. Swimming of the muskrat. Journal of
Mammalogy 16(1): 22-25.

Virgl, J. A., and F. Messier. 1992. Seasonal variation in body
composition and morphology of adult muskrats in central
Saskatchewan, Canada. Journal of Zoology 228(3): 461—
477.

Received 19 November 2013
Accepted 4 January 2014

Vol. 128

THE CANADIAN FIELD-NATURALIST

Pygmy Shrew (Sorex hoyi) in Montana East of the Rocky Mountains
with Comments on its Distribution across the Northern Great Plains

PAUL HENpRICKS!:? and SUSAN LENARD?

!Philip L. Wright Zoological Museum, Division of Biological Sciences, University of Montana, Missoula, Montana 59812
USA

°708 2nd Street, Helena, Montana 59601 USA

3Corresponding author: paul.hendricks@umontana.edu

Hendricks, Paul, and Susan Lenard. 2014. Pygmy Shrew (Sorex hoyi) in Montana east of the Rocky Mountains with comments
on its distribution across the northern Great Plains. Canadian Field-Naturalist 128(2): 204-206.

Range maps for Pygmy Shrew (Sorex hoyi) show a large hiatus over much of the northern Great Plains between the Rocky
Mountains and eastern North Dakota. We report a new record of the Pygmy Shrew in northeastern Montana, review previous
records for the state and adjacent regions bordering Montana to the north and east, and suggest that the range boundary in the
northern Great Plains be redrawn farther south to include all of Montana north of the Missouri River. This is consistent with
the known range of the Pygmy Shrew in eastern North Dakota and South Dakota, where the species has been documented
only north and east of the Missouri River, although records are still lacking from north of the Missouri River in northwestern
North Dakota and adjacent regions of Canada. Pygmy Shrews will probably be found at additional localities in prairie regions
of Canada adjacent to Montana, most likely in association with prairie pothole wetlands, river bottom riparian vegetation, and

hardwood draws.

Key Words: distribution; Montana; northern Great Plains; prairie potholes; Pygmy Shrew; Sorex hoyi; shrews; habitat

The Pygmy Shrew (Sorex hoyi) occurs across boreal
and subarctic North America, with populations in the
United States also extending south along the Rocky
Mountains in the west and the Appalachian Mountains
in the east (Long 1974; Diersing 1980; Hall 1981).
Pygmy Shrews occur in a variety of microhabitats, such
as bogs, marshes, dry grassy clearings, mesic forested
habitats, and associated riparian corridors (Long 1974;
van Zyll de Jong 1983).

In Montana, Pygmy Shrews were first reported in
1937 from the Rocky Mountain region in the western
third of the state (Koford 1938), and most additional
specimens have been reported west of or near the con-
tinental divide (Setzer 1952; Hoffmann ef al. 1969:
Key 1979; Foresman 1999). More than 150 individuals
have been captured at about 29 locations in Montana
over at least 160 000 pitfall trap—nights (Hoffmann er
al. 1969; Allen et al. 1997; Foresman 1999; Carson et
al. 2006*; Dorak et al. 2012*), indicating that Pygmy
Shrews may be relatively common at some locations,
but uncommon to’rare over much of their range within
the state. In montane western Montana, Pygmy Shrews
tend to occupy mesic sites in a variety of habitats typ-
ical of other portions of their North American range;
these include sagebrush, riparian areas, marshy or bog-
gy sites, and most coniferous forest types except those
at the highest elevations (Foresman 1999, 2012).

Most distribution maps for the Pygmy Shrew show
a large hiatus in the southern part of its range that
projects northward between the Rocky Mountains of
western Montana and southern Alberta in the west to
eastern North Dakota and South Dakota in the east
(Long 1974; Diersing 1980; Hall 1981; van Zyll de
Jong 1983; Naughton 2012). This is consistent with
the characterization of the Pygmy Shrew as a species

with boreal and montane affinities (Hoffmann and
Jones 1970; Jones et al. 1983). Thus, the Pygmy Shrew
is inferred to be absent from most prairie regions of
the northern Great Plains east of the Rocky Mountains.
In this note, we describe a new specimen of Pygmy
Shrew captured in the prairie region of eastern Mon-
tana and discuss this finding in the context of previous
records of the species from the northern Great Plains.

On 11 September 2012, we captured a female Pygmy
Shrew in a Museum Special snap trap (Woodstream
Corp., Lititz, Pennsylvania, USA) from a trap line set
in a stringer of riparian Eastern Cottonwood (Populus
deltoides) adjacent to an irrigation canal 2.9 km north
of the Missouri River (48°3'36"N, 106°14'16.79"W,
622 m elevation), 10.5 km southeast of Nashua, Val-
ley County, Montana (Figure 1, site 4). The line con-
sisted of single snap traps baited with peanut butter
and Sherman live traps (H. B. Sherman Inc., Talla-
hassee, Florida, USA) baited with rolled oats and bird
seed at 10 stations. The specimen was prepared as a
study skin (UMZM 20382; museum acronyms follow
the Global Registry of Biodiversity Repositories [http:
//grbio.org]) and deposited in the University of Mon-
tana, Philip L. Wright Zoological Museum. This was
the only Pygmy Shrew captured during 940 trap-nights
of effort at 46 sites. It was the first record of the species
from Valley County and the fifth specimen reported
in Montana from prairie regions east of the Rocky
Mountains. Also captured in this trap line were Deer
Mice (Peromyscus maniculatus) and White-footed
Mice (P. leucopus).

Pygmy Shrew specimens previously found in east-
ern Montana (Figure 1, sites 1-3) have been associated
with mostly treeless prairie pothole regions north of the
Missouri River. The first record (site 1) was a female

2014

bases =

Alberta

250
EE ————— ees iO meters

NOTES

205

Saskatchewan 49°N

North Dakota

45°N
Wyoming

I
~~
°
x

©
Q
r=
$

}
in

FiGuRE |. Distribution of Pygmy Shrew (Sorex hoyi) in Montana. Numbered sites are locations where Pygmy Shrews have
been found in prairie regions of the northern Great Plains: (1) Widgeon Slough, Sheridan County; (2) Wild Horse
Lake, Hill County; (3) Goose Lake, Sheridan County; (4) 10.5 km southeast of Nashua, Valley County. Sources for
unnumbered locations are Koford (1938), Setzer (1952), Hoffmann et al. (1969), Key (1979), Foresman (1999), and
specimens in the University of Montana, Philip L. Wright Zoological Museum (UMZM). The continental divide is

shown in bold.

(UMZM 19026) captured by Dennis Flath and John
Ciralli in a snap trap on 23 July 1977 near Widgeon
Slough (48°58'22.79"N, 104°15'7.19"W, 663 m eleva-
tion) in Sheridan County, about 170 km east of the
Valley County site. The second record (site 2) was a
partial skull (UMZM 18672) recovered from a raptor
pellet on 20 July 2000 near Sage Creek at Wild Horse
Lake (48°59'5.99"N, 110°10'26.4"W, 853 m elevation)
in Hill County, about 315 km west of the Valley Coun-
ty site (Hendricks 2001). The third record (site 3)
included two unsexed individuals (UMZM 20394, the
other specimen misplaced) captured by Jenny Flesch
in a pitfall trap and a snap trap at one trap station on
22 August 2005 in a marshy area near Goose Lake
(48°47'31.2"N, 104°2'56.4"W, 625 m elevation) in
Sheridan County, also about 170 km east of the Valley
County site.

Although few in number, the distribution of Pygmy
Shrew records from eastern Montana suggests that
this species is widely distributed across the northern
Great Plains in Montana north of the Missouri River,
but in localized wetland and riparian habitats. Exten-
sive small-mammal trapping throughout eastern Mon-
tana south of the Missouri River (Lampe et al. 1974;
Matthews and Swenson 1982; Carson ef al. 2006*;
Dorak et al. 2012*) has resulted in no additional records

for this shrew species. We suggest that future range
maps for the Pygmy Shrew in Montana include the
entire region north of the Missouri River.

The southern border that we propose for the range
of the Pygmy Shrew across the northern Great Plains
appears to match, more or less, the maximum extent
of the Laurentide ice sheet during the late Pleistocene
(Clayton and Moran 1982). This distribution of the
Pygmy Shrew in eastern Montana is also consistent
with documented locations in the eastern half of North
Dakota and eastern South Dakota, where the species
has been found only north and east of the Missouri
River (Long 1974; Hall 1981; Jones e¢ a/. 1983; Hig-
gins et al. 2002), often in association with marshy
prairie potholes and lakeshores and Green Ash (Frax-
inus pennsylvanica) hardwood draws (Gruebele and
Steuter 1988; Backlund 1995) in formerly glaciated
regions of the Drift Plains and Missouri Coteau. We
expect that Pygmy Shrews will be found at addition-
al locations in eastern Montana and adjacent prairie
regions north of the Missouri River in Canada and
northwestern North Dakota as more surveys are under-
taken for shrews in appropriate wetland, hardwood
draw, and riparian habitats in this area of the northern
Great Plains.

206

Acknowledgements

Our small-mammal surveys in northeastern Mon-
tana were part of a larger vertebrate survey conducted
for the Assiniboine and Sioux Tribes of the Fort Peck
Indian Reservation, funded under Tribal Government

Resolution 26-571-2012-03. We thank Jeanne Spaur of

the tribal government for soliciting and coordinating
the funding of our survey, which was managed through
the auspices of the Montana Natural Heritage Program.
We are most grateful to Claudine Tobalske of the Mon-
tana Natural Heritage Program for producing the map.
An earlier draft of our paper benefited from the com-
ments of two anonymous reviewers. All vouchers are
deposited in the Philip L. Wright Zoological Museum
(UMZM), University of Montana, Missoula.

Documents Cited (marked * in text)

Carson, S., A. Messer, R. Rauscher, and S. Story. 2006.
Statewide small mammal and Montana Fish, Wildlife and
Park’s lands vertebrate inventory project: 2006 final report.
Montana Department of Fish, Wildlife and Parks, Helena,
Montana. 22 pages.

Dorak, B., J. Stewart, R. L. Rauscher, D. Waltee, and A.
Begely. 2012. Milk River conservation and restoration state
wildlife grant, native species monitoring final report 2008—
2011. Montana Fish, Wildlife and Parks Region 6, Glasgow,
Montana. 65 pages plus appendices.

Literature Cited

Allen, K. L., D. Flath, and T. Weaver. 1997. Small mammal
capture efficiencies among three trap types. Intermountain
Journal of Sciences 3: 1-6.

Backlund, D. C. 1995. New records for the Dwarf Shrew,
Pygmy Shrew, and Least Shrew in South Dakota. Prairie
Naturalist 27: 63-64.

Clayton, L., and S. R. Moran. 1982. Chronology of late Wis-
consinan glaciation in middle North America. Quaternary
Science Reviews |: 55-82.

Diersing, V. E. 1980. Systematics and distribution of the
Pygmy Shrews (subgenus Microsorex) of North America.
Journal of Mammalogy 61: 76-101.

Foresman, K. R. 1999. Distribution of the Pygmy Shrew,
Sorex hoyi, in Montana and Idaho. Canadian Field-
Naturalist 113: 681-683.

Foresman, K. R. 2012. Mammals of Montana. Second edi-
tion. Mountain Press, Missoula, Montana. 429 pages.
Gruebele, M. J., and A. A. Steuter. 1988. South Dakota
records of Pygmy and Arctic shrews: response to fire.

Prairie Naturalist 20: 95-98.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Hall, E. R. 1981. The Mammals of North America. Second
edition, Volume I. John Wiley & Sons, New York, New
York. 600 pages.

Hendricks, P. 2001. A significant new record of the Pygmy
Shrew, Sorex hoyi, on the Montana—Alberta border. Cana-
dian Field-Naturalist 115: 513-514.

Higgins, K. F., E. D. Stukel, J. M. Goulet, and D. C. Back-
lund. 2002. Wild Mammals of South Dakota. Second edi-
tion. South Dakota Department of Game, Fish and Parks,
Pierre, South Dakota. 278 pages.

Hoffmann, R. S., and J. K. Jones, Jr. 1970. Influence of
late-glacial and post-glacial events on the distribution of
recent mammals on the northern Great Plains. Pages 35—
394 in Pleistocene and Recent Environments of the Central
Great Plains. Edited by W. Dort, Jr. and J. K. Jones, Jr.
Department of Geology, University of Kansas Special Pub-
lication 3. University Press of Kansas, Lawrence, Kansas.

Hoffmann, R. S., P. L. Wright, and F. E. Newby. 1969. The
distribution of some mammals in Montana I. Mammals
other than bats. Journal of Mammalogy 50: 579-604.

Jones, Jr., J. K., D. M. Armstrong, R. S. Hoffmann, and C.
Jones. 1983. Mammals of the Northern Great Plains. Uni-
versity of Nebraska Press, Lincoln, Nebraska. 379 pages.

Key, C. H. 1979. Mammalian utilization of floodplain habi-
tats along the North Fork of the Flathead River in Glacier
National Park, Montana. M.S. thesis. University of Mon-
tana, Missoula, Montana. 151 pages.

Koford, C. B. 1938. Microsorex hoyi washingtoni in Mon-
tana. Journal of Mammalogy 19: 372.

Lampe, R. P., J. K. Jones, Jr., R. S. Hoffmann, and E. C.
Birney. 1974. The mammals of Carter County, southeastern
Montana. Occasional Papers of the Museum of Natural
History, University of Kansas 25: 1-39.

Long, C. A. 1974. Microsorex hoyi and Microsorex thomp-
soni. Mammalian Species 33: 1-4.

Matthews, W. L., and J. E. Swenson. 1982. The mammals of
east-central Montana. Proceedings of the Montana Acad-
emy of Sciences 41: 1—13.

Naughton, D. 2012. The Natural History of Canadian Mam-
mals. Canadian Museum of Nature and University of Tor-
onto Press, Toronto, Ontario. 784 pages.

Setzer, H. W. 1952. Pigmy Shrew, Microsorex, in Montana.
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van Zyll de Jong, C. G. 1983. Handbook of Canadian mam-
mals. 1. Marsupials and Insectivores. National Museum of
Natural Sciences. Ottawa, Ontario. 210 pages.

Received 4 January 2014
Accepted 29 January 2014

Book Reviews

c q , j 3 J a+ la are oc i 1 . a) Ii i. ;
on Review Editor s Note: We are continuing to use the current currency codes. Thus Canadian dollars are CAD, U.S.
dollars are USD, Euros are EUR, China Yuan Remimbi are CNY, Australian dollars are AUD and so on.

ZOOLOGY

Animals of the Serengeti and Ngorongoro Conservation Area

By Adam S. Kennedy, and Vicki Kennedy. 2014. Princeton University Press, 41 William Street, Princeton, NJ, USA, 08540-

5237. 152 pages, 18.50 USD, Paper.

Birds of the Serengeti and Ngorongoro Conservation Area
By Adam S. Kennedy. 2014. Princeton University Press, 41 William Street, Princeton, New Jersey, USA, 08540-5237. 224

pages, 18.50 USD, Paper.

The Serengeti-Mara ecosystem is world-renowned
for its massive herds of migratory wildebeest, zebra,
and other grazers, encompassing approximately 25,000
km? of northern Tanzania and adjacent Kenya. The
bulk of the ecosystem lies within Serengeti National
Park and the Ngorongoro Conservation Area and adja-
cent game reserves, with the remainder in the Maasai
Mara of Kenya. Immediately to the south of the Seren-
geti ecosystem are the Ngorongoro Highlands includ-
ing the famous Ngorongoro Crater. Following up on the
2012 success of their guides on the animals and birds
of the Maasai Mara, the Kennedy’s have recently re-
leased Wildlife of the Serengeti and Ngorongoro Con-
servation Area and Birds of the Serengeti and Ngoron-
goro Conservation Area. Former safari camp managers
and now professional guides, Adam Scott Kennedy and
his wife Vicky bring their photographic talents, expe-
rience, and first-hand knowledge of the Serengeti,
Ngorongoro, and surrounding areas together in this
new pair of field guides. Although these two new
field guides enter a relatively crowded market, there is
a niche for them, particularly for visitors on their first
East African safari.

Animals of the Serengeti and Ngorongoro Conser-
vation Area is a bit of a misnomer since it only treats
mammals and a few reptiles, with birds in a separate
volume and nothing at all on amphibians, fish, or inver-
tebrates. It does cover 70 mammal species, including
most of the species likely to be encountered on safari
in Serengeti and Ngorongoro, or for that matter else-
where on Tanzania’s “Northern Circuit”. A notable
exception is the Oryx, which was has been observed
just east of Serengeti National Park in the Loliondo
area near Lake Natron. The guide also excludes bats,
shrews, and most of the 40 or so species of rodents
known from the Serengeti. No great loss, for although
important components of the ecosystem, these small
mammal groups are difficult to see and even harder to
identify when not in the hand. The Animals field guide

also covers 18 of the reptiles most likely to be observed
on safari or near Serengeti area lodges. It includes such
obvious choices as Nile Crocodiles and Black Mam-
bas, but also brightly-coloured agamas and nocturnal
geckos. Surprisingly, neither the Pancake Tortoise nor
the Egyptian Cobra were included, both striking species
that I have seen by day in the Serengeti.

The guide is focussed on the Greater Serengeti area
and has a brief introductory section with maps and text
on the general geography and ecology of the area, in-
cluding the great migration, as well as suggested locales
for great wildlife viewing. While I might quibble with
some of the details, the authors have done an admirable
job distilling this information into a concise introduc-
tion that I think it will help visitors place what they see
into a larger ecosystem context and ultimately deep-
en their appreciation.

The guide is packed with 146 colour photos, show-
ing both males and females for sexually dimorphic
species, and even young where space permits. In addi-
tion to great photographs, what sets this guide apart
from many traditional field guides is the light and ac-
cessible tone of the text. Each species has a half-page
to 4-page treatment, with information on identification,
preferred habitat, diet, and habits. Although total length
and shoulder height are provided, I do wish that aver-
age weight (or a range) was also provided for each
mammal species. For better-known species, there are
interesting notes on their distribution in the Serengeti
and Ngorongoro, as well juicy tidbits on their ecology
or behaviour. For example, the reader is warned of the
honey badger’s tendency to go for the genitals when
confronting humans. Perhaps apocryphal, it makes for
good reading nonetheless and is a part of safari lore.
The guide also has informative sidebars with insight
and observations provided by local Tanzanian guides,
the “Super Six”. It is a bit like having an experienced
flesh-and-blood guide along with you.

207

208

Common and scientific names used in the guide
generally follow those used by IUCN, and alternative
names typically provided where necessary to avoid
confusion. Welcome additions included for each spe-
cies are the Kiswahili and Maasai names (good luck
with the pronunciation!), and the etymology for some
of the more interesting names is also provided. The
species are presented in a “simple order” whose under-
lying rationale escapes me, and I think standard taxo-
nomic order would have been preferable. Fortunately,
there is a detailed Table of Contents listing all species,
as well as both a detailed and a short index at the back,
so it isn’t difficult to find a particular species.

Although some may prefer to read it back home, this
field guide is compact and light enough (14 « 21 cm
and only 320 g) to bring on safari. Wildlife sightings are
often fleeting, in poor light, or obscured by vegetation,
so having a field guide handy to confirm identification
is helpful, particularly for lesser-known species. Even
if you don’t pull out it out while on game drives, hav-
ing the guide handy to refer to in the evening or other
down time will help reinforce the names of unfamiliar
species observed during the day. If you are particularly
keen about herps, I would also recommend getting a
copy of Spawls et al.’s 2006 Reptiles and Amphibians
of East Africa (240 pages) that covers 230 of the more
commonly encountered species reptile and amphibian
species.

With over 500 documented species, Serengeti Na-
tional Park and the Ngorongoro Conservation Area are
recognized as Important Bird Areas by Birdlife Inter-
national, and the Serengeti Plains are also an Endemic
Bird Area. Novice birders may find this diversity daunt-
ing, and traditional field guides covering all of East
Africa, with 600 pages and 1400+ species, can be over-
whelming. Adam Scott Kennedy’s Birds of the Serengeti
and Ngorongoro Conservation Area is intended to
make this area’s birdlife more accessible by using pho-
tographs and limiting the guide to 264 of the more
commonly observed species. I was surprised by several
omissions such as the spectacular Southern Red Bishop
and the endemic Grey-crested Helmet-shrike, and was
puzzled by the inclusion of a sidebar on the Dodo.
Fortunately, it makes note of the endemic Fischer’s
Lovebird, Usambiro Barbet, Grey-breasted Spurfowl,
and Rufous-tailed Weaver, which are relatively easy to
find in the Serengeti.

The book is arranged by nine “habitats”, with birds
typically found in a particular habitat grouped togeth-
er, rather than by standard taxonomic order as in most
field guides. The habitats are described in the introduc-
tion and include: plains; marsh & water; woodland

THE CANADIAN FIELD-NATURALIST

Vol. 128

scrub & garden; acacia scrub; village birds; forest &
crater highland; air; night; and Lake Victoria specials.
The intent is to help novices more rapidly locate the
likely bird species in the guide based on where it is ob-
served. This approach has its limitations due to poorly
defined or overlapping habitats, especially for widely
distributed species such as the Grosbeak (Thick-billed)
Weaver, which was included as a “Lake Victoria Spe-
cial” but is also found in the Ngorongoro highlands and
beyond. I also found it frustrating, for example, to have
the hornbills spread out on pages 37, 119, 169, and 183.
Fortunately, the habitats are colour-coded in the guide,
and there are numerous cross-references to similar-
looking species that may occur in other habitats.

The quality of the photographs is truly impressive,
as is the artistic and technical skill evident in the lay-
out. There are typically 2-3 species per page (with full
page accounts for some of the more charismatic or
spectacular species), and the backgrounds of the numer-
ous images are seamlessly blended, so that it often
appears as if birds were actually side by side in their
natural habitat. Both sexes are often shown, and even
some immatures. The informative and accessible spe-
cies’ accounts have tips on identification and similar
species, and touch upon distinctive songs or calls, be-
haviour, ecology, and other points of interest. Scientific
names are not provided in the main text (they are in an
index at the back), but most novice birders often have
a hard enough time remembering the common name,
let alone the Latin one. KiSwahili names are sadly not
provided (Helmeted Guineafowl the lone exception),
but perhaps should have been, at least for the more dis-
tinctive species such as the Ostrich and vultures. Al-
though it is not a bird-finding guide, a bit more infor-
mation on local distribution/abundance or particular
birding “hot-spots” would also have been appreciated.
For example, the cliffs of Olkarien Gorge at the eastern
edge of the ecosystem are well-known (at least in some
circles) as a critical nesting site for Riippell’s Vultures.

This field guide is “aimed at all levels of birding
ability” and it is definitely suitable for novice birders
and for those on a safari primarily for mammals and
larger wildlife. The guide only covers about half of the
species known from the Serengeti and Ngorongoro
Conservation Area, so keen birders will likely want to
bring a more comprehensive field guide on their trip.
Nonetheless, the quality of the photographs and text
would make it a fine addition to field naturalist’s library
with an interest in East Africa. :

Dr. ROBERT F. FOSTER

Northern Bioscience, 363 Van Horne Street, Thunder Bay.
ON, Canada, P7A 3G3 .

2014

Rare Birds of North America

Book REVIEWS

209

By Steve N. G. Howell, lan Lewington, and Will Russell. 2013. Princeton University Press, 41 William Street, Princeton, NJ,

USA, 08540-5237. 448 pages, 35.00 USD, Cloth.

The avid lister will need to wear a bib for this book
as it contains all the species they will drool over. The
authors include all the species seen in North America
five times or less since the 1950s. These are the real
rarities or “mega-ticks” that move the combat birders
to drive for hours to add them to their lists.

Rarities come at us from three directions. Some
sweep in on winds from Europe, others overrun their
territory from the south and some blow over from east-
ern rim of Asia. Not surprisingly the east coast (New-
foundland to Maine), the states that border Mexico
and Alaska’s outer islands account for a high propor-
tion of these strays. Record committees have to evaluate
new sightings, taking in to account the bird’s migratory
tendencies, records of movements in other countries
as well as the evidence provided by the observer. For
example, if a European finch is found in Newfound-
land, the committee will consider if the species has a
history of wandering, is it being seen in Iceland or
Greenland and is it a popular cage bird. The authors
have taken all the accepted rarity records to 2011 plus
a few later sightings. For this book the authors have
stretched these limits and include some species that
have questionable provenance, yet may be true records
for a coverage of 262 species. For example, the authors
discuss the records of the Common Shelduck, a bird
usually dismissed as an escapee by the ABA, on the
grounds that it is possible some may be true vagrants.
They do not include species that breed annually even
though they may be vagrants for most of the continent
(Eastern Yellow Wagtail). The book has a North Amer-
ican focus so it does not include many rare Canadian
birds (such as Thick-billed Kingbird) as these are fre-
quently inhabitants of the southern States.

Before reading the species accounts the buyer needs
to read the introduction, as it contains much thought-
provoking information. There are discussions on migra-
tion and vagrancy, the influence of weather, mis-ori-
entation and disorientation and dispersal. The authors
introduce the concept of false vagrancy — birds that are
rarely seen but are probably regularly occurring species
(European Storm Petrel, Red-footed Booby). These
birds are in areas that simply lack coverage. The intro-
duction also covers topography moult and aging; com-
plex topics that need to be considered for any vagrant.

[ was surprised by the author’s use of Light-mantled
Sooty Albatross instead of the currently accepted
Light-mantled Albatross (Phoebetria palpebrata). They
also include the Eurasian Hen Harrier (Circus cyaneus)
as a separate species to the Northern Harrier (Circus
[cvaneus] husonicus), a split | have not seen accepted
elsewhere. (Although I have always thought they were
different species, based on their behaviour). There is an
error in the index for Little Shearwater as the quoted

page 98 is devoted to paintings of Thalassarche alba-
trosses,

The book is beautifully illustrated by lan Lewington.
The paintings are technically accurate, but also capture
the specie’s attitude effectively. For example, Lewing-
ton catches the funny, hunched posture of swallows, the
perky look of a Hoopoe and the grace of the frigate-
birds. Most birds are shown in different positions or
plumages as appropriate. The artist frequently includes
common, look-alike species (like Common and Spot-
ted Sandpipers, Little and Snowy Egret). I have seen
about half of the birds in this book, just not in North
America. So I really enjoyed looking at illustrations as
they evoked pleasant memories. This alone made the
book worthwhile for me.

One of my most recent lifers was the Himalayan
Bluetail or Orange-flanked Bush-robin. The vagrant
listed in this book is the Red-flanked Bluetail once
considered a subspecies Jarsiger c. cyanurus breeding
in northern Asia with 7. c. rufilatus breeding in the
Himalayas. The Himalayan Bluetail is now considered
by some as T. rufilatus, a distinct species. Given the
aggressive attention to detail by records committees as
carried into this book I was surprised not to see this
possibility considered. The more northern Red-flanked
Bluetail, if accepted as a split, is a long distance migrant
rather than a short-distance altitudinal migrant so it is
the more likely vagrant. The plates are lovely though
and they did bring up my views of this pretty little bird
along the roads around Darjeeling.

Clearly this is a must buy book for the passionate
list keepers, particularly those who are strongly com-
petitive. But there are good reasons for the rest of us
owning this book. When examining the details for a
rare species the authors need to consider the differences
to common birds. For example, separating the Euro-
pean Storm Petrel from the more abundant Wilson’s
Storm-petrel. These careful comparisons teach me
points of identification that I have overlooked for years.
I await the return of our American Coots so I can check
the under-tail coverts, something I have never done on
the thousands of coots I have seen. I will look for the
pale orange bill base on the Green-winged Teal (My
photo of a male does not show this, but it may be the
angle. I searched the net images and this is a very, very
difficult to see field mark. Good luck if you find a
vagrant Garganey). | will look for the black mark at
the base of the wings of a Great Blue Heron. Maybe I
will be able to see the plain primaries on an American
Woodcock if I use a better flashlight. It is an education
of field observation just to read this book. It is great
value for the price and a highly recommended pur-
chase for all birders.

Roy JOHN

2193 Emard Crescent, Beacon Hill North, Ottawa, ON, Cana-
da, K1J 6K5

210

Deer

THE CANADIAN FIELD-NATURALIST

Vol. 128

By John Fletcher. 2014. Reaktion Books Ltd., 33 Great Sutton Street, London, UK, EC1V ODX. 207 pages, 19.95 USD,

Paper.

As | read author John Fletcher’s book, Deer, |
couldn’t help but thinking of some of the deer that |
have seen in my lifetime. White-tailed deer are over-
abundant throughout much of Northeast U.S., but
interestingly, not particularly common where I live
(Cape Cod, Massachusetts), so encounters are espe-
cially memorable. My son and I commonly position
our trail-cam in different places to try and capture pic-
tures of the elusive white-tailed deer among other ani-
mals such as eastern coyotes/coywolves, fishers, and
otters inhabiting Cape Cod. I have fond memories of
seeing a white-tailed buck in a clear-cut stand and moose
in ponds in the White Mountain National Forest, New
Hampshire on family camping trips; moose in Baxter
State Park Maine is where my son saw his first wild
moose; Yellowstone National Park introduced me to
mule deer and elk (or wapiti as Fletcher refers to them
to not confuse them with moose since Alces alces is
called elk in the Old World), as well as bison, prong-
horn, bighorn sheep, and mountain goats, ungulates
that are related to deer. Although my professional inter-
est involves studying predators, mainly coywolves, I
am always interested in seeing and learning about the
various deer species that I encounter.

The back cover of Fletcher’s book introduces the
reader to his book and is worth quoting here: “The
Celts called them “fairy cattle” and the Greeks asso-
ciated them with the hunter goddess Artemis, but for
most people today, deer are seen as cute, like Bambi,
or noble, like the Monarch of the Glen. They can be a
danger when we’re driving at night, or they can simply
be a tasty venison burger. But while we may not often
eat humble pie—an actual pie filled with deer organs—
deer still appear in religion and mythology, on coats of
arms, in fine art, and in literature ranging from The
Yearling to Harry Potter and The Chronicles of Narnia.
In Deer, veterinarian and deer farmer John Fletcher
brings together the cultural and natural history of these
dignified animals. Fletcher traces the evolution of deer,
explaining why deer grow and cast aside their antlers
each year and describing their symbolism in various
cultures throughout history. He divulges the true story
of Rudolph and Santa’s other reindeer and explores the
role deer have played as prized objects of the hunt in
Europe, Asia, and America. Wide-ranging and richly
illustrated, Deer provides a fresh perspective on this
graceful, powerful animal that will appeal to hunters
and gatherers alike.”

The above description accurately articulated the
meaning of the book and what I learned from it. The
book is well illustrated with 93 pictures, 66 of those in
colour. Fletcher introduces the 40 species of wild deer
(including elk/red deer, moose, roe deer, white-tailed
deer, and caribou/reindeer), or Cervidae family, found

throughout the world. Fletcher makes it clear that there
are many kind of deer (like moose) even if they aren’t
the proto-typical “deer”, such as the white-tailed deer
in North America. The first chapter provides a good
taxonomic introduction to both Old and New World
deer. While Fletcher divides them into the two regions
it can be a little confusing as some species such as elk/
wapiti, Cervus elaphus (meaning pale-coloured rump
by Native Americans; note this same species is called
red deer in Europe) and North American moose (con-
fusingly called elk in the Old World) occur circum-
polar. That being said, the first chapter does a good
job of introducing the cast of characters of the book,
with red deer being the species most often referred to
throughout the text.

In the introduction, Fletcher noted that the book is a
cross fertilization between science and the arts. That
was certainly the case as the book mostly focussed on
human uses of and interactions with deer. I learned
some interesting things such as: (1) grazers are gen-
erally gregarious and are adapted to resist high infes-
tations of parasites whereas browsers concentrate their
food and have little ability to combat them; (2) deer are
strongly linked to human symbolism such as regener-
ation (i.e., males re-growing a set of antlers each year),
renewal, power, longevity, royalty, and justice; (3)
many of the early “noble” people of the Old World
(e.g., kings/queens) were associated with romantic
images of deer, such as putting collars on wild red deer
stags with inscriptions like the date and person putting
the collar on the deer (those collars would later be re-
trieved by others who saw the message from the roy-
alty that inscribed them); (4) there is a close association
between the legends of Robin Hood and deer such that
he and his comrades would steal/poach deer from the
rich to feed the poor, and this scene is in every movie
on the subject; (5) red deer imported to New Zealand
have become the first domesticated livestock in 5,000
years; and (6) many deer species will eat meat if given
the chance. My favourite quote from the book came on
page 61 when Fletcher referred to the image of deer
as “ephemeral wild animals briefly glimpsed which
contributes to their mystery and allure, fuelling their
symbolic role.”

John Fletcher lives in Scotland where he runs a deer
farm, which is a private estate with an enclosed deer
herd living within its confines. His background and
location clearly influenced the book as it was heavily
biased toward Old World (Europe) deer species. In fact,
in chapter 1, where he provided an introduction to the
family of deer, the Old World species were discussed
three times more than the New World deer (about 15
versus 5 pages). Furthermore, many of his accounts of
cultural symbolism, such as converting deer into sha-

2014

mans, came from Old World examples and much less
from the New World. At times I had difficulty follow-
ing the many places and names (especially kings,
queens, and other royalty) to which Fletcher discussed,
This probably isn’t too surprising since I have lived
in the United States my entire life and find the whole
concept of nobility and classification of different classes
of people bizarre, and almost imaginative (e.g., “George
Walpole, third Earl of Orford”). On page 142, Fletcher
even acknowledged the complexity of symbolism by
noting, “For us in the twenty-first century, the world
of medieval and Renaissance symbolism seems eso-
teric and confusing.” Even though many of the passages
from the book were intended for or likely to be under-
stood by only a small number of people with a special-
ized knowledge or interest, the thoroughly illustrated
book really helped bring some of the images that he
discussed to life. For me, the pictures made the differ-
ence in enjoying, rather than struggling to finish, the
book. Finally, Fletcher’s use of British terms such as
hind and hart (for female and male red deer) is con-
fusing and takes a little adjusting to. I actually had to
Google “hind” (and other terms) to understand what he
was referring to.

Fletcher did attempt to make up the Old World bias
by devoting an entire chapter to North America deer.
However, the one issue I had with that chapter and with
the rest of the book in general, was that the book was
overwhelmingly about human utilization of deer and
less on deer biology and ecology. The author mentioned
numerous times about hunting being a very necessary
thing in today’s world and sort of belittled the “urban
population” that is against such things — little mention
was made of society’s evolving relationship with nature
from utilization and romance to coexisting with and
aesthetically enjoying wildlife among us. While I cer-
tainly agree that deer can become overabundant and hu-
man hunting is a necessary tool in certain instances, I
felt that the book was too preachy on that topic. His
quote on page 182 was probably accurate but sound-
ed much like state wildlife departments here in North
America, “As human urbanization and commuting be-
come the norm, deer numbers grow (due to lack of
hunters) and the image of deer as rare and precious
mystical creatures risks being gradually displaced by
perceptions of deer as suburban nuisances.” Why not
recognize that many people enjoy seeing them on a

Dolphin

Book REVIEWS 211

daily basis and that it enriches our lives even if they
can pose a hazard to us? That isn’t to say that deer hunt-
ing is not a major influence of societies from historical
to present times, but it would have been refreshing to
not view deer only as a game animal and commodity.

Fletcher had many chances to draw attention to other
aspects of the Cervid family. For instance, there was
essentially no discussion of predators. In fact, carni-
vores were mentioned a mere eight times in the book
with most being in reference to wolves. Yet all of these
were in passing and did not describe the importance of
predators to both the evolution and management of deer
species. It is well noted that even though humans have
relied on deer over the course of our species’ relative
short existence, predators and prey have co-evolved
with each other over a much longer period of time and
the reason we see the fleetness of white-tailed deer,
the robustness of bison and moose, and the regalness of
wapiti is in part due to evolving defenses from preda-
tors such as wolves. I found it wrong that there would
be no chapter devoted to the need to restore predators
to landscapes where deer numbers are rising due to de-
clines in human hunting. As the book currently stands,
I think a better title would have been “Of Deer and
Men”, to steal the adage from Barry Lopez’s famous
1978 book on wolves. To my point, at the end of the
book Fletcher lists some associates and websites and
all are related to deer farms and venison partnerships
(i.e., to be used as food); no sources were given on deer
ecology and biology.

Deer is an interesting and well written book that
details the symbolism and the close association that
humans and deer have had over centuries. Anywhere
that people have lived, humans have made use of some
of the 40 species of deer found worldwide. There are
definitely things in this book that everyone will learn
for the first time and that alone makes the book worthy
of reading. And there are also many wonderful images
that really bring the chapters to life. As long as one
accepts the Old World (European) bias of the author,
and the relative lack of description of deer biology ver-
sus the focus on deer folklore, symbolism, and human
utilization of deer, this will be an enjoyable book to
read.

JONATHAN WAY
89 Ebenezer Road, Osterville, MA, USA, 02655

By Alan Rauch. 2014. The University of Chicago Press, 1427 East 60th Street, Chicago, IL, USA, 60637. 207 pages, 16.02

USD, Paper.

As Book Review Editor I receive unsolicited books.
If I think the subject is specialised and I know an expert
who writes well, I will ask them to review the book.
Otherwise I send a notice to the 50 or so reviewers I
have on file. About half of these people are regular re-

viewers, working on a book or more per year. Occa-
sionally no one wants to volunteer, so, if I think I have
sufficient knowledge, | will review it myself.

It has been my privilege to encounter dolphins on
many occasions, most recently the Gangetic Dolphin in

the Koshi river of Nepal and the Pink and Grey River
Dolphins in the Amazon. Therefore when this book sat
unclaimed for two weeks | decided to review it myself.
Before I got to the end of the first page I was entranced.

This little book has only six chapters, covering zool-
ogy, species, mythology, behaviour, dangers and as cul-
ture icons. In addition to the usual references (including
websites) there is a two-page time-line from 1349 to
present, showing our human interaction with this mam-
mal.

The zoology chapter covers the fundamental biology
of current dolphins. It summarises the evolution of
these marine mammals from their deer-like ancestors
over about 50 million years. The author follows their
change from a four-legged and nose bearing animal to
the flippered and blow-hole bearing mammal we know
today. He covers the often weird ideas of early natu-
ralists and the fish-mammal confusion. Rauch describes
the adaptations that dolphins have made to live under-
water — in their shape, vision and the internal body
changes.

The chapter on species covers only enough dolphins
to establish their diversity. From the whale-sized Orca
to the 1.4 m Hector’s dolphin and the more oddball
river dolphins, the author describes those species that
establish the boundaries of dolphin appearance.

Mythology discusses the real or fanciful stories that
have arisen about dolphins. These range from dolphins
with saw backs that slice open crocodile bellies, to the
origin of “le Dauphin” of France and the numerous tales
of dolphins rescuing humans. The story of Pelorus Jack,
a Risso’s Dolphin, has an intriguing twist, taking it from
real to myth.

The chapter on Intelligence, Social Behaviour and
Echolocation gives us an insight into the dolphin’s
world. It also tells how little we really know. In partic-

THE CANADIAN FIELD-NATURALIST

Vol. 128

ular the author explains their wonderful ability to use
echolocation. He also spends time recounting their “in-
telligence” although it seems futile to measure a marine
mammal’s brain power using human, land-based, think-
ing.

Apart from the natural dangers that dolphins face,
we humans add other risks. A very small portion of hu-
mans directly kill dolphins for food. More frequently
they die enmeshed in fishing nets set for tuna. Even
those that get released from these nets are so injured
that they die later. Pollution adds to their stresses. As an
apex predator their fat contains pesticides, fire retar-
dants and other carcinogens. They have been captured
for zoos, research and even the military.

Dolphins have been intertwined with human culture
for millennia. Our perspective has ranged from fero-
cious killers to children’s soul mates. We have glorified
them as surrogate humans and eaten them as fish. There
are dolphin mascots, dolphin door handles, dolphin
helmets and an amazing array of dolphin decorative art.
We have kept dolphins as “pets” and performers. Yet
none of this equates to seeing dolphins in the wild,
behaving as dolphins should.

The author explores the role of dolphins in the media,
from movies to comic books. He is not complimentary
of the way we have portrayed these mammals. We tend
to glorify these animals as smart, semi-humans while
ignoring their real and fascinating lives.

I enjoyed reading all the chapters and felt the author
presented information in an unbiased manner that was
thought provoking. Dolphins is a fascinating and well
written book and I am delighted that I was forced to
“volunteer.”

Roy JOHN

2193 Emard Crescent, Beacon Hill North. Ottawa. ON,
Canada, K1J 6K5

Wild Again: The Struggle to Save the Black-footed Ferret

By David Jachowski. 2014. University of California Press, 155 Grand Avenue, Suite 400, Oakland, CA, USA, 94612-3758.

256 pages, 34.95 USD, Cloth.

As I read author David Jachowski’s book, Wild
Again, | envisioned:a time when the Great Plains were
undisturbed by human activity and all wildlife existed
in their historical pre-European abundance, including
black-footed ferrets and their prairie dog prey. This
would be a time before diseases, such as plague and
distemper, ravaged species that had not co-evolved with
those ailments. Instead, my vision included large areas,
many thousands of acres, of prairie dog colonies that
had grazing animals, such as pronghorn and bison, on
it and predators, like wolves and grizzly bears, follow-
ing them.

Jachowski offers an engaging, personal account of
his involvement in black-footed ferret (Mustela nig-
ripes) recovery efforts. Once feared extinct, rediscov-
ered in Meeteetse, WY in 198] (page 31-35), and still

one of North America’s rarest mammals, this small
weasel exemplifies the ecological, social, and political
challenges of conservation in the West, including the
risks involved with intensive captive breeding and rein-
troduction to natural habitats. As the jacket cover of the
book indicates, the author draws on more than a decade
of experience working to save the ferret to base his
writings, and his unique perspective and informative
anecdotes reveal the scientific and human aspects of
conservation as well as the immense dedication re-
quired to protect a species on the edge of extinction.
By telling one story of conservation biology in practice
—its routine work, triumphs, challenges, and inevitable
conflicts—Wild Again gives readers a greater under-
standing of the conservation ethic that emerged on the
Great Plains as part of one of the most remarkable re-

2014

covery efforts in the history of the Endangered Species
Act. | couldn’t agree anymore with that accurate state-
ment and essential summary of the volume.

I greatly appreciated the personal focus of Wild
Again which the author unabashingly states both in
the jacket cover as well as in the Prologue section.
Jachowski comments that the book is written for every-
one and is not intended to be a comprehensive, techni-
cal review of ferrets (page vii), yet the 18 page Further
Readings chapter at the end makes it very authoritative
and comprehensive, in my opinion. One gets to learn
of the highs and lows of being a field biologist and
many of the author’s experiences could be substituted
for any animal, such as little funding, political influ-
ence and interference, difficult living situations, and
self-sacrifice beyond most jobs. In my research on east-
erm coyotes/coywolves, I have often been criticized for
being too personal and liking my study subjects, so it
was very refreshing to read the commentary on conser-
vation biology being a blend of advocacy and science
(page 29). May be our politicians, including within
wildlife agencies, will one day better appreciate that
scientists are people too and should be able to offer their
professional as well as personal opinions without fear
of repercussions on their job and career security.

| was amazed that an author around my age (mid to
late 30s) could have such of a developed and eloquent
writing style, in addition to a well balanced perspec-
tive, as he wove many diverse accounts from personal
to professional stories into one wonderful, easy-to-read
adventure. For example, his beginnings as a child help-
ing researchers at his father’s Patuxent field research
station, his late-night bar episodes, and his personal
relationship issues of living on the Great Plans in iso-
lation are captivating and mentioned throughout numer-
ous sections of the text. And the descriptions of spot-
lighting ferrets at night, mapping prairie dog burrows
during the day, detailing nuances of ferret behaviour
(such as females testing males during coitus, page 97),
trying to captive breed and increase ferret numbers, and
conducting research in many different regions, from
Montana to Mexico, made the over 200 page hardcov-
er a real page turner. This is saying something consid-
ering that book sales and traditional book reading for
my generation (and in general) are on the decline as the
internet and information age produces news items (e.g.,
global warning, poaching, habitat loss) that seemingly
change every few minutes, and as Jachowski explains
(page 210), this makes it difficult for even professional
conservation biologists to figure out what to protect and
how to react to such large volumes of details.

While readers may be turned off by the apparent
negative tone and unfortunate outcomes that are often
discussed in Wild Again, such as plague killing off
prairie dog colonies and people poisoning prairie dogs
right at the edge of ferret recovery areas, I strongly
believe that these occurrences must be documented and
included as these are the personal and professional
experiences of many wildlife biologists the world over.
There certainly aren’t always positive endings in the

Book REVIEWS

world of wildlife research as evidenced by the fact that
even after 22 years (1991-2013) and releasing more
than 3,000 captive reared ferrets at 19 reintroduction
areas, the black-footed ferret is only about a quarter of
the way to being numerically recovered (page 212),
which involves at least 30 self-sustaining adults inhab-
iting 10 different locations. Given that the main prey of
ferrets, prairie dogs, are candidates for Endangered
Species Act listing, it is not surprising that they still face
a long and uphill climb, even with advocates like Jach-
owski trying to help them along.

The author (page 211) nicely concludes with a dis-
cussion of the direct, utilitarian benefits to humankind
from conserving biodiversity, such as food, medicine,
and water quality, yet he argues that sentimentality is
another reason why we preserve nature and animals
like ferrets and the prairie dogs that they depend on.
Jachowski stresses that it is perhaps our most selfless
act to try to preserve something when it is not in the
strictest sense physically, energetically, or evolutionary
beneficially to do so (page 211). Even though it is the
law in the US to protect endangered species, | also
believe that defending and helping to preserve a con-
troversial animal is courageous given the usual hatred,
prejudice and paranoia displayed to many endangered
or politically challenging animals (see page 175-177).
While I was shocked to learn that prairie dog poison-
ing is still allowed and practised in the US, and find it
pathetic that this is a publically funded activity to this
day, I found it entirely predictable that entrenched (and
I would argue hypocritical) politicians will fight on the
behalf of ranchers and farmers to continue such expen-
sive, environmental destructive, and indefensible activ-
ities. In fact, Jachowski explained (page 175) that in
South Dakota, hatred of prairie dogs still divides the
community as easily as religion or politics does in other
parts of the country.

The manuscript has only a few, minor grammatical
errors, and most descriptions seem accurate and real-
istic for the locations discussed. There are pictures dis-
persed throughout the book which adds character and
enables the reader to see prairie dogs, black-footed fer-
rets, and some of the locations where the author con-
ducted research. However, not knowing the area that
well, and doing what the author says most people do
(i.e., flying over the plains from the forests [of subur-
bia] where I live to the mountains of places like Yel-
lowstone Park where I vacation), I wish there were
more maps to see the different places that the author
visited and studied ferrets in (there is a map on page
158 but it is of four territories of female ferrets in his
SD study area). Other than the lack of maps, my only
other inconsequential complaint was really just three
further questions that were not answered in the text but
which I pondered: (1) Because the book was so com-
prehensive, personal, and well articulated, | wanted to
know why Jachowski was no longer with the ferret
team. | did find it entirely predictable for him to publish
his findings after his decade plus stint studying ferrets,
as expressing his views while still employed would

have likely (and most unfortunately) predisposed him
to ridicule and possible even job firing as he described
occurring for other people advocating for ferrets (e.g.,
page 160). (2) What about his wife mentioned in the
acknowledgments (page 216)? He discussed other rela-
tionships he had, and how he chose the grasslands over
the woman he loved (page 5), so how did he meet her?
And (3), how was he able to move around so much
during his job, such as to the Conata Basin of South
Dakota from the UL bend in Montana. And why did
he have to pay out of pocket for things, such as hous-
ing in SD, if he was working for the federal govern-
ment? I wonder if he worked a part-time (seasonal) job
with the federal government, known as a 1039 term
position where one cannot work more than 6 months in
a year, and then volunteered the rest of the time. These

OTHER

Ecology

THE CANADIAN FIELD-NATURALIST Vol.

three questions would have really rounded out the book,
which essentially covered from his childhood to the
present (2013, I believe). }

As you can probably tell, if | am critical of such
trivial details that means I really was enthralled with
Wild Again and was able to comprehend every major
section and storyline that happened within. I recom-
mend it for anyone who is interested in wildlife, species
recovery, and stories and adventures of this highly en-
dangered and endemic North American predator, the
black-footed ferret. It is a delightful read and well worth
the time and money to learn about this little known and
studied predator, and the grassland ecosystems and
prairie dogs that it relies upon.

JONATHAN WAY

89 Ebenezer Road, Osterville, MA, USA, 02655

By Michael L. Cain, William D. Bowman, and Sally D. Hacker. 2014. 3 Edition. Sinauer Associates, Inc., Publishers, Sun-
derland, MA, USA, 01375-0407. 596 pages, 134.95 USD, Cloth.

The third edition to an already useful text contains
some very important changes and improvements from
the two earlier editions. The author’s goal for this edi-
tion was to improve the usefulness of the text for both
students and instructors and as with previous editions
the focus in the undergraduate student. Aside from
the text being updated, I will highlight the significant
changes as well as provide a general overview of the
text.

Ecology is composed of 25 chapters arranged into
7 Units to cover the complete depth of the field of ecol-
ogy. The Units represent the scale and scope of ecology:
individual, population, community, ecosystem, and
global. In addition, there are units on the major eco-
logical interactions of competition, predation, parasi-
tism, and mutualism/commensalism, and evolutionary
ecology. Natural selection forms the overlying basis for
the entire text.

The content of this edition includes a new chapter
on behavioural e¢ology which brings into focus the
importance that behavior plays in populations and the
structure of communities as well as interactions. It in-
cludes not only the ways that organisms live in aggre-
gations but also foraging and mating behavior with the
context always focused on the evolution of the behav-
iour as an important life history strategy to enhance
fitness.

Several important features are worthy of note. First
of all, the authors have included exercises that require
analyzing data. This is not only an essential skill in
ecology but also aids in developing critical thinking
skills. The authors have continued to provide case stud-
ies of important concepts but have also added “Con-
nections in Nature” which shows how chapter concepts

relate to the concepts in other chapters.

Pertinent to one of our greatest ecological challenges,
this edition has a web feature “Climate Change Con-
nections” which provides some useful material to relate
ecological concepts to issues surrounding climate
change. Most chapters have of this edition have exam-
ples relating to climate change.

An example of another new feature are the “Ecolog-
ical Toolkit” boxes. These boxes highlight important
techniques used by ecologists. Among the interesting
ones are stable isotope analysis and estimation of pop-
ulation abundance and population growth rates.

The supplemental online content appears to be of
high quality but at the time of review (05/2014) the
publisher only has material for one chapter (4) avail-
able for examination. There appears no indication that
this content will require a subscription although cer-
tainly this could change once the site is completed.

This edition has many other features that make it an
improvement to an already good text. I would strongly
recommend using this text if you are not already using
the second edition and if you are, | recommend upgrad-
ing to this edition. If you want a personal text that can
be used to facilitate self-learning of ecology or updat-
ing your ecological knowledge, this text is recommend-
ed. I can also recommen4¢ this edition as a reference if
you wish to have an ecology text on your shelf. The
addition of online content, especially if it remains free
as noted above, enhances the value.

ROGER D. APPLEGATE

Tennessee Wildlife Resources Agency, Nashville, TN, and
Department of Biology and School of Environmental Stud-
ies, Tennessee Tech University, Cookeville, TN. USA

2014

Book REVIEWS

The Once and Future Great Lakes Country: An Ecological History

By John L. Riley. 2013. McGill-Queen’s University Press, 1010 Sherbrooke West. Suite 1720, Montreal, QC, Canada, H3A

2R7. 516 pages, 39.95 CAD, Cloth.

Wow. Check out this book if you are craving a read
with vast dimensions. I don’t mean the physical book,
either. That, in hardcover, has a comfortably satisfying
heft like your favourite dictionary or rock purloined
from nature. The cover alone is breathtaking, an oblique
aerial view of a landscape in orange and gorgeous water
blue. The book doesn’t identify the place, and I couldn’t
either. Even if it is cover art, to provide an aerial photo
with no label is the perfect way to torture a geogra-
pher, or at least to cause one to spend a lot of time
scanning around on Google Earth. Instead I enlisted
friends to the case via Facebook. We think, thanks to
Sean Blaney and Becky Whittam, that it’s Matchedash
Bay looking north to Georgian Bay.

I will declare my bias. | know many reading this
join me in calling the Great Lakes country home, our
lives framed by these lakes. The Bruce Peninsula and
Lake Huron shoreline, for instance, have scoured our
childhood memories. The Waterloo region was the
landscape and ecology lab through university. Then
we were in thrall to the thrum of bird migration over
the Lake Erie shore. Ian Tamblyn’s Woodsmoke and
Oranges embodies our time with Lake Superior. John
Riley’s work is rich in time, drilling into prehistory and
telescoping into the future. We can place it on the same
library shelf as other ecological and environmental his-
tories, both regional and broader in scope, but if offers
quite a bit of cultural introspection too, an element you
might not always find elsewhere.

At the time of publication, Riley was a senior sci-
ence advisor for the Nature Conservancy of Canada.
He begins this story on his own land where with his
family he occupies an old farm (there are thousands
like it, he says) in the Great Lakes basin. From there he

builds outward in both time and geography to follow
this remarkable region through so many passages. It
is very well-written. It pulled me along, with loads of
fascinating information, analysis and evocation of land-
scapes, wildlife, people and peoples. Human beings are
the significant force of ecological change for much of
the book’s span, so their history and influence are sen-
sitively portrayed. Riley skillfully stitches past with
present and future so that the reader never forgets the
relevance of any one to the other. The book also is thor-
oughly referenced and noted, with much reliance on
first person accounts and other authors.

Just to whet your curiosity, | offer these chapter titles:
“Land Beyond Memory: Before 1500”; “Wilding the
Land with War: the 1700s”; “Taming the Unforested:
Prairies, Alvars, Barrens, Cliffs, Bogs and Fens” and
“Restoration: A New Native Landscape”. I am a fool
for prehistory and early history, so those chapters held
me more tightly than the later ones. The final chapters
contain large amounts of optimism, making the case
that the current trend is actually toward conservation
rather than the opposite. The optimism puzzles me, but
I will take it because I so thirstily want it.

I can recommend this book without reservation. You
may or may not take issue with the author’s points of
view on some subjects, or interpretation of historical
records (there is a heck of a lot to consider!) but it is
finely presented for your inspection. As well, Riley’s
writing voice is utterly human and companionable.
The hardcover edition is lovely to have. I bought the
e-book as well, for ease of searching by word.

BEV MCBRIDE
574 Tweedsmuir Ave., Ottawa, ON, Canada, K1Z 5P2

New TITLES

Prepared by Roy John

+ Available for review * Assigned

THE CANADIAN FIELD-NATURALIST

Vol. 128

Currency Codes — CAD Canadian Dollars, USD U.S. Dollars, EUR Euros, AUD Australian Dollars.

ZOOLOGY

A Checklist of North American Amphibians and
Reptiles: The United States and Canada. Volume 1
— Amphibians. By M. J. Fouquette, Jr, and Alan
Dubois. 2014. Xlibris LLC, 1663 Liberty Drive, Suite
200, Bloomington, IN, USA, 47403. 613 pages, 34.95
USD, Cloth, 29.95 USD, Paper. (Request for a review
copy refused by publisher.)

* Animals of the Serengeti. By Adam S. Kennedy, and
Vicki Kennedy. 2012. Princeton University Press, 41
William Street, Princeton, NJ, USA, 08540-5237. 152
pages, 27.95 USD, Paper.

Beetles of Eastern North America. By Arthur V.
Evans. 2014. Princeton University Press, 41 William
Street, Princeton, NJ, USA, 08540-5237. 560 pages,
35.00 USD, Paper.

* Birds of the Kenya’s Rift Valley. By Adam S.
Kennedy. 2014. Princeton University Press, 41 William
Street, Princeton, NJ, USA, 08540-5237. 264 pages,
29.95 USD, Paper.

* Birds of the Serengeti. By Adam S. Kennedy. 2012.
Princeton University Press, 41 William Street, Prince-
ton, NJ, USA, 08540-5237. 176 pages, 27.95 USD,
Paper.

The Thing with Feathers: The Surprising Lives of
Birds and What They Reveal About Being Human.
By Noah Strycker. 2014. Penguin Group (USA), 375
Hudson Street, New York, NY, USA, 10014-3657. 304
pages, 18.37 USD, Cloth.

The Sibley Guide to Birds, Second Edition. By David
Sibley. 2014. The Knopf Doubleday Group, 1745
Broadway, New York, NY, USA, 1001. 624 pages,
40.00 USD, Cloth.

The ROM Field Guide to Butterflies of Ontario. By
Peter Hall, Colin Jones, Antonia Guidotti, and Hub-
ley Brad. 2014. Royal Ontario Museum, 100 Queen’s
Park, Toronto, ON, Canada, M5S 2C6. 64 pages, 29.99
CAD, Cloth.

* Wild Again - The Struggle to Save the Black-
Footed Ferret. By David S. Jachowski. 2014. Uni-
versity of California Press, 2120 Berkeley Way, Berke-
ley, CA, USA, 94704-1012. 256 pages, 34.95 USD,
Cloth.

How Snakes Work: Structure, Function and Behay-
ior of the World’s Snakes. By Harvey B. Lillywhite.
2014. Oxford University Press, North Kettering Busi-
ness Park, Hipwell Road, Kettering, Northamptonshire,
UK, N14 1UA. 224 pages, 35.00 GBP, Cloth.

* Wildlife of the Caribbean. By Herbert A. Raffaele,
and James W. Wiley. 2014. Princeton University Press,
41 William Street, Princeton, NJ, USA, 08540-5237.
304 pages, 19.95 USD, Paper.

OTHER

*Amphibian Conservation - Global Evidence for
the Effects of Interventions. By Rebecca K. Smith,
and William J. Sutherland. 2014. Pelagic Publishing,
P.O. Box 725, Exeter, UK, EX1 9QU. 279 pages, 29.99
GBP, Paper.

Experimental Evolution and the Nature of Biodi-
versity. By Rees Kassen. 2014. Roberts and Company,
4950 S. Yosemite Street, F2 #197, Greenwood Village,
CO, USA, 80111. 288 pages, 45.00 USD, Cloth.

The Analysis of Biological Data. By Michael Whit-
lock, and Dolph Schluter. 2014. Roberts and Compa-
ny, 4950 S. Yosemite Street, F2 #197, Greenwood Vil-
lage, CO, USA, 80111. 704 pages, 123.75 USD, Cloth.

Ecology — Third edition. By Michael L. Cain, Wil-
liam D. Bowman, and Sally D. Hacker. 2014. Sinauer
Associates, Inc., 23 Plumtree Road, P.O. Box 407, Sun-
derland, MA, USA, 01375-0407. 648 pages, 129.95
USD, Cloth.

Whale-watching: Sustainable Tourism and Ecolog-
ical Management. By James Higham, Lars Bejder, and
Rob Williams. 2014. Cambridge University Press, Uni-
versity Printing House, Shaftesbury Road, Cambridge.
UK, CB2 8BS. 110.00 CAD, Cloth.

News and Comment

Upcoming Meetings and Workshops

International Society for Behavioral Ecology (ISBE) Conference 2014

The 15" International Behavioral E cology Congress
to be held 31 July — 1 August 2014 by Hunter College
of the City University of New York (CUNY) and the
behavioral ecology research community based at var-

ious CUNY campuses and New York University.
Registration is currently open. More information is
available at http://www.isbe2014.com.

Conservation and Biology of Tortoises and Freshwater Turtles Annual Symposium

The 12" annual symposium on the Conservation and
Biology of Tortoises and Freshwater Turtles to be held
4-7 August 2014 in Orlando, Florida. The meeting,
sponsored by Zoo Med Laboratories, Inc., is co-hosted
by the Turtle Survival Alliance and the IUCN Tortoise
and Freshwater Turtle Specialist Group (TFTSG).

Ecological Society of America Meeting 2014

The 99" annual meeting of the Ecological Society of
America to be held 10-15 August 2014 at the Sacra-
mento Convention Center in Sacramento, California.
The theme of the meeting is “From Oceans to Moun-

The meeting represents the largest gathering of non-
marine turtle biologists in the world and provides an
unmatched opportunity for networking and strategiz-
ing turtle conservation. More information is available
at http://www.turtlesurvival.org/get-involved/con fer-
ence#.U16FgldB870.

tains: It’s All Ecology” and registration is currently
open. More information is available at http://esa.org
/am/.

Northeast Partners in Amphibian and Reptile Conservation Meeting 2014

The 15" annual meeting of the Northeast Partners
in Amphibian and Reptile Conservation (NEPARC)
to be held 13—15 August 2014 Allegany State Park,
Salamanca, NY. NEPARC is an active, diverse, and
inclusive partnership dedicated to the conservation of

International Ornithological Congress 2014

The 26" International Ornithological Congress (IOC)
to be held 18-24 August 2014 at Rykkio University in

amphibians and reptiles and their habitats throughout
northeastern North America. Poster abstract deadline
is 11 July 2014. More information is available at http://
www.northeast parc.org/meetings/index.htm.

Tokyo, Japan. More information is available at http://
[oc26.jp.

American Ornithologists’ Union/Cooper Ornithological Society/Society of Canadian

Ornithologist Meeting 2014

Annual meeting of the American Ornithologists’
Union (132™ Stated Meeting), the Cooper Ornitholog-
ical Society (84'" Stated Meeting), and the Society of
Canadian Ornithologists (32" Stated Meeting) to be
held 23-27 September 2014 at the YMCA of the Rock-
ies in Estes Park, CO. The meeting will feature 5 days

of workshops, contributed scientific papers, posters,
and invited speakers, including symposia that look at
exciting new approaches and results in ornithology.
More information is available at https://www.bird-
meetings.org/aoucossco2014/default.asp.

Raptor Research Foundation Conference 2014

Raptor Research Foundation 2014 Conference to
be held 24-28 September 2014 at the Emerald Beach
Hotel in Corpus Christi, Texas. Co-Hosts are the Caesar
Kleberg Wildlife Research Institute at Texas A&M

THE CANADIAN FIELD-NATURALIST

University, Kingsville and Hawk Watch International.
More information is available at http:// www.raptor-
researchfoundation.org/conferences/cur
rent-conference.

Raptor Workshop: Introduction to Raptor Field Techniques

Raptor Services, LLC presents 5-day workshops to
be held in Stevens Point, Wisconsin by Gene Jacobs
of the Linwood Springs Research Station. This intro-
ductory level field course is designed to instruct stu-
dents in a full-range of field techniques used in the
study of raptors. The workshop is offered every June
with an emphasis on learning breeding season field
techniques and again in August, September, and Octo-
ber, with an emphasis on migration and winter research
techniques. Receive first-hand experience working

Retirement of Associate Editor C. D. Bird

After serving The Canadian Field-Naturalist as
Associate Editor for almost 40 years, longer than any
of his contemporaries except for ornithologist Tony
Erskine, Dr. Charles Durham Bird has stepped down
in early 2014. He primarily reviewed botanical submis-
sions but also some in entomology and general natural
history. First appointed in 1975 when Lorraine Smith
was journal editor, he has continued without interrup-
tion through the subsequent editorial terms of Francis
Cook and Carolyn Callaghan. In recognition of his con-
tributions he was made an Honorary Member of The
Ottawa Field-Naturalists’ Club in 2005 (see Canadian
Field-Naturalist 119(4): 614), and has received numer-
ous Alberta awards for his contributions. Charley was
born in 1932, a son of Ralph Durham Bird (1901—
1972, see tribute in 1972 Canadian Field-Naturalist
86(4): 393-399). The senior Bird was a prairie ento-
mologist with Agriculture Canada, widely known for
his classic monograph Ecology of the Aspen Parkland,
(1961).

Although born in Oklahoma when his father was
teaching at The University of Oklahoma, Charley was
raised in Manitoba, where he developed early inter-
ests in botany from his mother (Lois Gould, 1905—
1959), and entomology from his father. He obtained a
BSc from University of Manitoba in 1956, MSc from
Oklahoma University, Stillwater, in 1958, and PhD
from Oklahoma State University in 1960. His thesis for
the latter was on vegetational and waterfowl changes

with: live raptors, capturing, handling, banding tech-
niques, broadcast call surveys, tree climbing, rappel-
ling, blood sampling and more. Summer Sessions: 2-6
June and 23— 27 June; Fall sessions: 25-29 August
and 13-17 October 2013, and two weekend sessions
5,6,7,13,14, September. Cost is $450 and space is
limited (6-8 students per workshop). More informa-
tion is available at http://www.raptorresearch.com
/workshop.htm.

in P.F.R.A. reservoirs in west-central Canada based on
surveys for Canadian Wildlife Service in the summers
of 1956-1959. In 1960-1962 he held a National Re-
search Council Postdoctoral Fellowship at the Univer-
sity of Alberta, became an Assistant Professor of
Botany in 1962, Associate Professor in 1967, and Full
Professor in 1974. He and his wife Ann purchased a
farm near Mirror, Alberta, in 1975 and he took early
retirement from the university in 1979. In 1992 they
sold the farm and retired in Erskine, Alberta.

He has contributed over 300 articles, notes, book re-
views, and reports, notes in print and electronic media,
primarily on mosses, lichens, liverworts and Lepidop-
tera but also including tributes, conservation, and ex-
tended family history items. He has collected over
37,000 insects for eventual deposit in collections of the
University of Alberta and 36,000 plants largely already
deposited there or in various other herbaria. Presently,
Charley is continuing his work on the Lepidoptera of
natural areas in south-central Alberta as well as his
research on family genealogy. In addition, he is con-
ducting inventory work of the biota of the J.J. Collett
Natural Area and contributing regularly to their web-
site. Like Tony Erskine, he has agreed to review occa-
sional future papers submitted to The Canadian Field-
Naturalist whenever his expertise and perspective are
particularly needed.

FRANCIS R. COOK

2014

Obituary — Farley Mowat 1921-2014

An ardent, outspoken conservationist, fervent Cana-
dian, and world acknowledged northern literary suc-
cess, Farley Mowat, has died at his home at Port Hope,
Ontario, 8 April 2014. He is survived by his wife Claire.

Mowat showed an affinity for natural history, writ-
ing, and controversy early (see Cook 2003), starting as
a teenager with a bird column in the Saskatoon Star-
Phoenix in the mid 1930s as he recounted in Born
Naked (1993). After some initial success, this was ter-

minated when he submitted an enthusiastic account of

the sexual activities of some of his subjects that was
considered too lurid for newspaper public of the time.

Mowat seemed destined to a career as a scientist
when he went to the University of Toronto and took his
first collecting expedition with fellow zoology students
Frank Banfield and H. Hord to Saskatchewan in the
summer of 1939 (as recalled in Otherwise, 2008). Ban-
field covered mammals and Mowat and Hord birds.
and sold their collections to the Royal Ontario Museum
to finance the trip. Both Mowat and Banfield subse-
quently enlisted in Canadian army and served over-
seas in World War II but wrote up their field results and
published in The Canadian Field-Naturalist. Charac-
teristically, Banfield submitted his account before going
overseas and Mowat only after his return.

The latter was apparently the first and last formal
scientific paper Mowat wrote, but far from his last
expedition or literary publication. After the war he was
field assistant in the Northwest Territories to the leg-
endary and irascible North American mammalogist
and naturalist, Francis Harper. A clash of personalities
soon led Farley to undertake his own explorations sep-
arately (Harper later extracted a reciprocal promise that
neither would mention the other in their respective
future writing, a promise also extracted from Mowat
by later field companions for their lifetimes).

A research proposal led to his hiring by the Cana-
dian Wildlife Service for the four-researcher Caribou
survey which Banfield headed. Later, however, Ban-
field was ordered to fire Mowat by the chief of the
Wildlife Service due to local complaints and lack of
advance formal approval for some activities.

Cut loose, Mowat turned to writing — notably Peo-
ple of the Deer (1952) and Never Cry Wolf (1963),
on Mowat’s conclusions on the plight of some native
northern people and on the life and human persecution
of wolves, respectively. Although early reviewers of
both were captivated by the writing style and subjects,
later in depth reviews by researchers familiar with
areas, animals, and people, were generally hash due to
frequent wanderings from fact in both. Pre-eminent
among the critics were the prominent botanist Erling
Porsild (see Duthan 2012) and mammalogist Frank

News & COMMENT

219

Banfield (see Morris 1990) whose reviews appeared in
1952 (The Beaver June: 47) and 1964 (The Canadian
Field-Naturalist 78: 52-54), respectively. The reaction
to the former is extensively covered by Dathan 1997:
569-594; the latter drew a response to the CFN Editor,
reportedly from the wolf “Uncle Albert”, defending
Mowat’s account (The Canadian Field-Naturalist 78:
206).

Urged on initially by the resourceful promoter and
publisher of Canadian writers, Jack McCelland (see
King 1998), Mowat produced over 40 books which
were widely translated worldwide with sales of more
than fourteen million copies on topics ranging from his
experiences in WWII (particularly in And No Birds Sing
1979 — perhaps his finest writing), pets, boats, whales.
overharvest of ocean resources (Sea of Slaughter 1984),
a biography of Dian Fossey (1987), and serial portions
of autobiography.

Mowat never finished his degree and was lost to for-
mal science but his writings gained and entertained a
huge audience of readers as he was above all else a
really good story-teller with an appealing message. Al-
though he often admitted a tendency not to let the facts
get in the way of a good story, his writings made an
impact by focusing on basic truths as he saw them
and his delight in goring beaurocracy. By adding to a
growing public focus on wildlife and conservation they
helped promote pressure for increased governmental
protection at all levels and influenced young scientists
to research careers on the problems faced by the crea-
tures with which we share the earth. His appreciation
of northern fauna and his defense of wolves gained a
large appreciative audience far beyond North America
particularly in the country which also has vast arctic and
boreal areas, the Soviet Union, which he later also was
invited to visit, thus providing material for some sub-
sequent books.

See also:

Banfield, Frank A. 1942. Notes on Saskatchewan mam-
mals. Canadian Field-Naturalist 55(8): 117-123.

Cook, F. R. 2003. [Book review] Farley: The Life of Farley
Mowat By James King. Canadian Field-Naturalist 117(3):
509-510;

Dathan,Wendy. 2012. The Reindeer Biologist: Alf Erling
Porsild, 1901-1997. University of Calgary Press, Calgary,
Alberta.

Morris, R. D. 1998.Obituary Alexander William Francis
Banfield: 1918-1996. Journal of Mammalogy 79(1): 364—
369.

Mowat, F. M. 1947. Notes on the birds of Emma Lake, Sas-
katchewan. Canadian Field-Naturalist 61(3): 105—115.
King, R. 1998. Jack: A Life with Writers. McCelland and

Stewart, Toronto, Ontario.

FRANCIS R. COOK

Editor’s Report for Volume 127 (2013)

Mailing dates for issues in volume 127:
Gl), Lémly201s

(Drs 2BEOstober 2018

(3) 4 December 2013

(4) 14 January 2014

In an effort to get caught up in the production sched-
ule, we published 8 issues of The Canadian Field-
Naturalist over a period of 14.5 months (3 October 2012
— 14 January 2014). Not surprisingly, submission rates
have increased concomitantly over the past few years;
a 56% increase since 2011. To ensure that we will keep
up with the higher number of submissions, we have in-
creased our team of Associate Editors.

The Canadian Field-Naturalist remains an important
forum for dispersing science on the distribution, taxon-
omy, biology, behaviour, ecology, and conservation of
species across a broad range of taxa in North America.
In volume 127, several new records were published of
species of mammals, fish, birds, insects, lichen, plants,
plesiosaurian, and ichthyosaur in particular regions of
Canada and one new record for the country. Some of
the other varied topics include the biology of rare, rein-
troduced, and invasive species, improved methods for
detecting species at risk, impacts of road infrastructure
and traffic on wildlife, effects of management actions
on the conservation of rare habitats, interspecific inter-
actions, species surveys in under-surveyed regions, and
analyses of datasets spanning many decades. As some
habitats and species undergo increased pressure from
the effects of anthropogenic activity, the utility of CFN
to conservation will increase via the provision of sci-
entific information based on field studies. The values
that led to the creation of CFN in the 1800s continue
to be relevant today.

Trina Rytwinski, Assistant Editor, edited content,
acted as an occasional Associate Editor, and acted as
Editor-in-Chief for two months. Elizabeth Morton
proofed and edited manuscripts; Wendy Cotie typeset
galleys, provided corrections for page proofs, and cre-
ated pdfs; Roy John, Book Review Editor, requested
books for review, selected reviewers, edited submitted

reviews, and prepared the new titles listings; Jay Fitz-
simmons, Journal Manager, managed financial ac-
counts, issue mailing, and requests for back issues, con-
ducted journal promotion through Twitter and other
means, and provided digital content to subscribers.
William Halliday prepared the Index with proofreading
by Trina Rytwinski and Dan Brunton. Sandra Garland,
webmaster of the Ottawa Field-Naturalists’ Club, post-
ed tables of contents, abstracts, and pdfs on the CFN
section of the OFNC website. Our Associate Editors
managed manuscripts, provided reviews and recom-
mendations, and guided authors through the revisions
process. The Publication Committee, chaired by Dan
Brunton and consisting Paul Catling, Jay Fitzsimmons,
Sandra Garland, Tony Gaston, Karen McLachlan-
Hamilton, Frank Pope, and Jeff Saarela effectively guid-
ed the operation of the journal. We are indebted to our
very dedicated and effective team.

A summary of the distribution of memberships in
the Ottawa Field-Naturalists’ Club and subscribers to
The Canadian Field-Naturalist for 2013 is provided in
Table 1. The number of articles and notes in Volume
127 is summarized in Table 2 by topic; totals for book
reviews and new titles is given in Table 3; and the dis-
tribution of content by page totals per issue is provided
in Table 4. Manuscripts (excluding book reviews, no-
tices and reports) submitted to The Canadian Field-
Naturalist totalled 84 in 2013. Of these, 73 were accept-
ed for publication, and 30 were published in volume
127, along with 16 manuscripts submitted in 2012 and
revised in 2013. A total of 25 Articles, 18 Notes, 3 Tri-
butes, and 3 Essays were published in 2013.

The following Associate Editors managed, assessed
and reviewed manuscripts published in volume 127
(number of manuscripts in parentheses): R. Anderson,
Canadian Museum of Nature, Ottawa ON (4); C. Cal-
laghan, Luskville QC (3); P. M. Catling, Agriculture
and Agri-Food Canada, Ottawa ON (3): F. R. Cook,
Emeritus Research Associate, Canadian Museum of Na-
ture, Ottawa ON (5); J. Foote, Algoma University, Sault
Ste. Marie ON (4); S. Jung, Yukon Government, White-

TABLE 1. The 2013 circulation of The Canadian Field-Naturalist (2012 in parentheses). Compiled by Jay Fitzsimmons from mail-

ing list for 127(4).

ee ee ae

Subscriber Type Canada USA Other Total
Memberships

Family & Individual 89 (89) 8 (8) | (1) 98 (98)
Subscriptions

Individual 79 (81) 28 (30) 3 (3) 110 (114)

Institutional 127 (2) 159 = (164) 28. (2%) 313 (314)
Total 295 (292) NOS (202) S261) 522 (525)

2014

TABLE 2. Number of research and observation articles and
notes published in The Canadian Field-Naturalist, Volume
127, by major field of study.

Subject Articles Notes Total
Mammals 7 7 I4
Birds 6 5 1]
Amphibians and Reptiles | | 2
Fish 2 | 3
Plants 3 2 5
Insects 2 2 4
Other + 0 4
Se ee eee eer
Total 25 18 43

TABLE 3. Number of reviews and new titles published in
the Book Review section of The Canadian Field-Naturalist,
Volume 127, by topic.

SSeS 79750—*"—————————

Reviews New Titles
Zoology 23 47
Botany 3 4
Environment 4 2
Miscellaneous y| 28
Children | 0
Total 38 81

horse YT (3); D. F. McAlpine, New Brunswick Museum,
St. John NB (5); D. W. Nagorsen, Mammalia Biological
Consulting, Victoria BC (5); C. Renaud, Canadian Mu-
seum of Nature, Ottawa ON (3); T. Rytwinski, Carleton
University, Ottawa ON (3); J. Saarela, Canadian Museum
of Nature, Ottawa ON (7).

The following referees reviewed manuscripts pub-
lished in volume 127 (number of manuscripts reviewed
>I in parentheses): Becky Abel, The Trumpeter Swan
Society, Plymouth MN; Ken Abraham, Ontario Ministry
of Natural Resources, Peterborough ON; Paul Abram,
Carleton University, Ottawa ON; Jim Atkinson, Uni-
versity of Guelph, Guelph ON; Shannon Berch, BC
Ministry of Environment, Victoria BC; Sean Blaney,
Atlantic Conservation Data Centre, Sackville NB; Ed-
ward Bork, University of Alberta, Edmonton AB; Ernie
Brodo, Canadian Museum of Nature, Gatineau QC;
Ronald J. Brooks, University of Guelph (retired),
Guelph ON; Dan Brunton, Ottawa ON; Angela Bur-
kinshaw, Alberta Environment and Sustainable Re-
source Development, Red Deer AB; Naomi Cappuc-
cino, Carleton University, Ottawa ON; Kevin Cash,
Environment Canada, Ottawa ON; Dr. Stephen Clay-
den, New Brunswick Museum, Saint John NB; Philip
A. Cochran, Saint Mary’s University of Minnesota,
Winona MN; Daniel Cristol, The College of William &
Mary Williamsburg VA; Dr. Rick A. Cunjak, University
of New Brunswick, Fredericton NB; Fredrik Dalerum,
University of Pretoria, Pretoria South Africa; Margaret
F. Docker, University of Manitoba, Winnipeg MB;
Erica Dunn, scientist emeritus, Environment Canada,

EDITOR’S REPORT 2

bo

TABLE 4, Number of pages per section published in 7he
Canadian Field-Naturalist, Volume 127 (2013), by issue.

=aeaBe—eaNjeeeoso————————————————————————

Issue
| 2 3 4 Total
Editorials 0 0 | 0) |
Articles 56 72 59 2D) oe 17,
Notes 19 14 12 20) 65
Book Reviews* 15 9 10 12 46
News and Comment,

Reports** 6 5 18 nm Se
Tributes 6 0 0 14 20
Index ~ - 10 10
Total 102 + 100 100 92 394

*Includes reviews and new titles listings
** Includes CFN Editor’s report, Minutes of the OFNC Annual
Business Meeting, and OFNC Awards report.

Ottawa ON; Tony Erskine, Sackville NB (2); Kris
Everatt, University of Pretoria, Pretoria South Africa:
Lenore Fahrig, Carleton University, Ottawa ON; Dianne
Fahselt, University of Western Ontario, London ON:
Jennfier Foote, Algoma University, Saulte Ste. Marie
ON; Jan Freiwald, University of California, Santa Cruz
CA; Marcel Gahbauer Stantec, Montreal QC; John A.
Gilhen, Nova Scotia Museum of Natural History, Hal-
ifax NS; Steve Hager, Augustana College; Rock Island
IL; Sarah Hambleton, Agriculture and Agri-Food Can-
ada, Ottawa ON; Mark Hanson, Fisheries and Oceans
Canada, Moncton NB; Lee Hastie, University of Aber-
deen, Aberdeen Scotland; Mark Hipfner, Environment
Canada, Delta BC; Howard M. Huynh, Texas Tech
University, Lubbock TX; Lawrence Igl, U.S.G.S. North-
ern Prairie Wildlife Research Center, Jamestown ND;
Danny Ingold, Muskingum University, New Concord
OH; Todd Katzner, West Virginia University, Morgan-
town WV; Charles Krebs, Professor Emeritus, Uni-
versity of British Columbia, Vancouver BC (2); Karl
Larsen, Thompson Rivers University, Kamloops BC:
Cori Lausen, Wildlife Conservation Society Canada,
Kaslo BC; Zoe Lindo, University of Western Ontario,
London ON; Eric Lofroth, Victoria BC; Scott McBur-
ney, Atlantic Veterinary College, University of Prince
Edward Island, Charlottetown PEI; Dr. Nick McCann,
Minnesota Zoo, Apple Valley MN; Dave Mech, Uni-
versity of Minnesota, St. Paul MN; Dr. David A. Meth-
ven, University of New Brunswick, Saint John NB;
Randall Miller, New Brunswick Museum, St. John NB;
James Miskelly, Royal BC Museum, Victoria BC;
Ron Moen, University of Minnesota, Duluth MN; Eric
Moise, University of Western Ontario, London ON;
Anders Pape Moller, Université Paris-Sud, Orsay Cedex
France; Ralph Morris, Brock University (retired), St.
Catharines ON; Marion Munroe, Nova Scotia Provin-
cial Museum, Halifax NS; Jan Murie (retired), Uni-
versity of Alberta, Edmonton AB; Dr. Clay Nielson,
Southern Illinois University, Carbondale IL; Michael
Oldham, Ontario Ministry of Natural Resources, Peter-

i)
NM
Nw

borough ON (3); Dr. Jack Parmalee, Agriculture and
Agri-Food Canada, Ottawa ON; Brent Patterson, On-
tario Ministry of Natural Resources, Peterborough ON
(2); Brian Peer, Western Illinois University, Macomb
IL; Paul M. Peterson, National Museum of Natural His-
tory, Smithsonian Institution, Washington DC; Roger
Powell, North Carolina State University, Raliegh NC;
Dr. Gilbert Proulx, Alpha Wildlife Research & Man-
agement Ltd., Sherwood Park AB; Rick Rosatte, On-
tario Ministry of Natural Resources, Peterborough ON;
Claude Samson, Parks Canada, Quebec City QC; Gau-
tam Sarath, USDA, Lincoln NE; B. Christian Schmidt,
Canadian Food Inspection Agency, Ottawa ON; Cory
Sheffield, Royal Saskatchewan Museum, Regina SK;
Dr. Becky Sjare, Fisheries and Oceans Canada, St.
John’s NL; Dr. Rick Starr, Moss Landing Marine Lab-
oratory, Moss Landing CA; David Strayer, Cary Insti-
tute of Ecosystem Studies, Millbrook NY; Don Suther-
land, Natural Heritage Information Centre, Ontario
Ministry of Natural Resources, Peterborough ON;
Zachary A. Sylvain, Colorado State University, Fort
Collins CO; Dr. John Therhane, University of New
Brunswick, St. John NB; Karen Vanderwolf, New
Brunswick Museum, St. John NB, and Canadian Wild-

THE CANADIAN FIELD-NATURALIST

Vol. 128

life Federation, Ottawa ON; David Walter, Royal Al-
berta Museum, Edmonton AB; Dr. Jane Watson, Van-
couver Island University, Nanaimo BC; Wayne Weller,
Niagara Falls ON; Jim Wilson, Saint John NB; Linda
Wires, University of Minnesota, Duluth MN; Graham
Young, Manitoba Museum, Winnipeg MB.

The journal was printed at Gilmore Printers, Ottawa.
Thanks to Gilmore for overseeing production and print-
ing. I am grateful to President Fenja Brodo and Council
of the Ottawa Field-Naturalists’ Club for their support
of the journal. I am also grateful to all of the individual
subscribers and authors who encouraged our team as
we strive to provide a high-quality scientific journal
on natural history. A special thank you to Liz Morton for
her years of service as copy editor, and to Leslie Cody
for producing the Index. Finally, we thank our fami-
lies for being patient and supportive throughout many
evenings and weekends of working on the journal.

CAROLYN CALLAGHAN
Editor in Chief

and

TRINA RYTWINSKI
Assistant Editor

TABLE OF CONTENTS (concluded) Volume 128, Number 2

Notes (continued)

Muskrat (Ondatra zibethicus) interference with aquatic invertebrate traps
MICHAEL C. CAVALLARO, ANSON R. MAIN, and Curisty A. MORRISSEY

Pygmy Shrew (Sorex hoyi) in Montana east of the Rocky Mountains with comments on its distribution
across the northern Great Plains PAUL HENDRICKS and SUSAN LENARD

Book Reviews

ZOOLOGY: Animals of the Serengeti and Ngorongoro Conservation Area — Birds of the Serengeti and
Ngorongoro Conservation Area — Rare Birds of North America — Deer — Dolphin — Wild Again: The
Struggle to Save the Black-footed Ferret

OTHER: Ecology — The Once and Future Great Lakes Country: An Ecological History

New TITLES

News and Comment

International Society for Behavioral Ecology (ISBE) Conference 2014
Conservation and Biology of Tortoises and Freshwater Turtles Annual Symposium
Ecological Society of America Meeting 2014

Northeast Partners in Amphibian and Reptile Conservation Meeting 2014
International Ornithological Congress 2014

American Ornithologists’ Union/Cooper Ornithological Society/Society of Canadian Ornithologist Meeting
2014

Raptor Research Foundation Conference 2014

Raptor Workshop: /ntroduction to Raptor Field Techniques
Retirement of Associate Editor C. D. Bird

Obituary — Farley Mowat 1921—2014

Editor’s Report for Volume 127 (2013) CAROLYN CALLAGHAN

Mailing date of the previous issue 127(4): 26 March 2014

2014

200

204

207
214
26

THE CANADIAN FIELD-NATURALIST Volume 128, Number 2

Articles

Effectiveness of stream sampling methods in capturing non-native Rusty Crayfish (Orconectes rusticus)
in Ontario Scott M. Reip and JANE DEVLIN

Estimating breeding bird survey trends and annual indices for Canada: how do the new hierarchical
Bayesian estimates differ from previous estimates?
ADAM C. SMITH, MARIE-ANNE R. HUDSON, CONSTANCE DOWNES, and CHARLES M. FRANCIS

Acoustic monitoring of migratory birds over western Lake Erie: avian responses to barriers and the
importance of islands CLAIRE E. SANDERS and DANIEL J. MENNILL

Parental care by lone male Ferruginous Hawks (Buteo regalis), Rough-legged Hawks (Buteo lagopus),
and Great Horned Owls (Bubo virginianus) was limited to providing food
JosEF K. SCHMUTZ, MARTIN A. GERARD, GORDON S. Court, and R. WAYNE NELSON

Apparent widespread decline of the Boreal Chorus Frog (Pseudacris maculata) in eastern Ottawa
Davip C. SEBURN, KARI GUNSON, and FREDERICK W. SCHUELER

Activity and diet of bats in conventional versus organic apple orchards in southern Michigan
BRENNA L. LONG AND ALLEN KURTA

Decline in breeding of the Great Black-backed Gull, Larus marinus, and the Herring Gull, L.
argentatus, on Boot Island, Nova Scotia, 1986 to 2010
COLIN M. MACKINNON and ANDREW C. KENNEDY

Distribution and abundance of baling twine in the landscape near Osprey (Pandion haliaetus) nests:
implications for nestling entanglement
RENEE SEACOR, KAYHAN OsSTOVAR, and MARCO RESTANI

Cues used by predators to detect freshwater turtle nests may persist late into incubation
JULIA L. RILEY and JACQUELINE D. LitzGus

Notes
A Gray Wolf (C anis lupus) delivers live prey to a pup L. DAviD MECH

Assessing capture success of small mammals due to trap orientation in field—forest edge habitat
DANIEL M. WoLcott, MADISON R. ACKERMAN, and MICHAEL L. KENNEDY

Effect of food patch discovery on the number of American Crows (Corvus brachyrhynchos) using
a flight lane WILLIAM LANGLEY

2014

11]

119

135

145

154

158

165

es

ie,

189

19]

(continued on inside back cover)

ISSN 0008-3550
MIX

Paper from
responsible sources

eee FSC® C100205

The CANADIAN
FIELD-NATURALIST

Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

Volume 128, Number 3 July-September 2014

The Ottawa Field-Naturalists’ Club

FOUNDED IN 1879

Patron
His Excellency the Right Honourable David Johnston, C.C.,C.M.M., C.O.M., C.M.
Governor General of Canada

The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these
fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-

ments of high quality for living things.

Ronald E. Bedford
Edward L. Bousfield
Charles D. Bird
Fenja Brodo

Irwin M. Brodo
Daniel F. Brunton
Michael D. Cadman

J. Bruce Falls

President: Fenja Brodo

18‘ Vice-President: vacant

2"4 Vice-President: Jeffrey H. Skevington
Recording Secretary: Annie Bélair
Treasurer: Ken Young

Paul M. Catling
Francis R. Cook
Bruce Di Labio
Anthony J. Erskine

Jay Fitzsimmons

Honorary Members

John M. Gillett
Peter W. Hall
Christine Hanrahan
C. Stuart Houston
Theodore Mosquin
Robert W. Nero

2014 Council
Daniel F. Brunton Don Hackett
Carolyn Callaghan David Hobden
Barbara Chouinard Diane Kitching
Owen Clarkin Diane Lepage

Barry Cottam Ann MacKenzie

E. Franklin Pope
Joyce M. Reddoch
Allan H. Reddoch
Dan Strickland
John B. Theberge
Sheila Thomson

Karen McLachlan-Hamilton
Lynn Ovenden

Rémy Poulin

Henry Steger

Eleanor Zurbrigg

lan Davidson

To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate
P.O., Ottawa, ON, K1Z 1A2, or e-mail: ofnc@ofnc.ca. For information on Club activities, go to www.ofnc.ca

The Canadian Field-Naturalist

The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this
journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.

Website: www.canadianfieldnaturalist.ca/index .php/cfn

Editor-in-Chief: Dr. Carolyn Callaghan Assistant Editor: Dr. Trina Rytwinski
Copy Editor: Sandra Garland Typographer: Wendy Cotie
Journal Manager: Dr. Jay Fitzsimmons: subscriptions @canadianfieldnaturalist.ca or via postal address above.

Book Review Editor: Roy John
Charles D. Bird
Paul M. Catling

Francis R. Cook
Jennifer R. Foote

Associate Editors: Anthony J. Gaston
Thomas S. Jung
Donald F. McAlpine

Garth Mowat

David Nagorsen
Claude B. Renaud
Marty Obbard

Jeffery M. Saarela
David C. Seburn
Jeffrey H. Skevington

Chair, Publications Committee: Daniel F. Brunton

All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by e-mail:
editor@canadianfieldnaturalist.ca or postal mail (with disk): 611 Chemin Cregheur, Luskville, QC JOX 2GO0; (819) 455-1087:
Book-review correspondence should be sent to the Book Review Editor by e-mail: rjohn@rogers.com or postal mail: 2193
Emard Crescent, Ottawa, ON, K1J 6K5

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$20 (student) includes an online subscription to The Canadian Field-Naturalist. Members can receive printed issues of CEN for
an additional $30 per volume (four issues). For further details, including discounts for members of the Canadian Museum of Nature,
see http://ofnc.ca/member.php. The club’s regional journal, Trail & Landscape, covers the Ottawa District and regional Club
events and field trips. It is mailed to Ottawa area members, and is available to those outside Ottawa on request. It is available to
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Cover: Lake Charr (Salvelinus namaycush) in spawning condition on Great Bear Lake, NT August 2007. Photo: Andrew Muir
See Note on anadromous migration Harris et al. on pages 260-264 in this issue.

MCZ
LIBRARY

. NOV 12 2014

The Canadian Field-Naturalist HARVARD

SU NIVERSITY
July-September 2014

Volume 128, Number 3

Home Range, Movements, and Denning Chronology of the Grizzly Bear
(Ursus arctos) in West-Central Alberta

KAREN GRAHAM!:? AND GORDON B. STENHOUSE!

lis Researc s ss
‘Foothills Research Institute Grizzly Bear Program, 1176 Switzer Drive, Hinton, Alberta T7V 1V3 Canada
“Corresponding author: kgraham@foothillsri.ca

Graham, Karen, and Gordon B. Stenhouse. 2014. Home range, movements, and denning chronology of the Grizzly Bear (Ursus
arctos) in west-central Alberta. Canadian Field-Naturalist 128(3); 223-234.

An understanding of the natural history of the Grizzly Bear (Ursus arctos) is important for recovery planning. We present data
on home range size, movements and denning chronology collected using Global Positioning System (GPS) collars on Grizzly
Bears in west-central Alberta. Mean annual kernel estimates for adult (1034 + 656 (SD) km?) and subadult (1298 + 1207 km?)
males were larger than those for females with cubs of the year (213 + 212 km?) and lone adult females (337 + 176 km?) but not
different from sub-adult females, females with yearlings, or females with > 2-yr old cubs (P > 0.05). Mean rates of movement
among female age-reproductive classes were different from each other (Z, < 2.70, P > 0.05) but not different from sub-adult
males (Z, < 2.70, P > 0.05). Rates of movement of adult males were significantly different only from those of females with cubs
of the year (Z, = 3.94, P= 0.001). The greatest amount of movement occurred in June and the least in October. Bears traveled
fastest in the morning and evening and slowest at night. Pregnant females had the longest denning period (175 days, + 16 days
SD). No difference was detected in denning duration among the remaining five age-sex—reproductive classes (P > 0.05). GPS
collars provided large location datasets from which accurate home range estimates, hourly movement rates, and precise denning
dates were determined. Examining similarities and differences in the basic biology of Grizzly Bears from various locations will

improve our understanding of the plasticity of this species and the potential impacts of habitat and climate change.

Key Words: Grizzly Bear; Ursus arctos; home range; movement; denning chronology; GPS collar; Alberta

Introduction

Many populations of the Grizzly Bear (Ursus arctos)
are at some level of endangerment, and a better under-
standing of the species’ basic biology is important to
aid conservation efforts (Servheen 1993; Alberta Sus-
tainable Resource Development 2008*). The collection
and collation of biological data from across the range of
the Grizzly Bear provides an opportunity to compare
current data with those in past studies as well as studies
from other areas, so that we might better understand the
plasticity and adaptability of this species. Increased
knowledge of its basic biology may also increase our
ability to predict impacts of landscape change, habitat
modification, and climate change on Grizzly Bear pop-
ulations.

The purpose of this paper is to present information
on Grizzly Bear home range size, movements, and den-
ning chronology in west-central Alberta from bears
equipped with Global Positioning System (GPS) collars
in west-central Alberta. We compare age—sex classes
in terms of home range size, movements, and denning
chronology. Further, we compare movements during
different months and different times of day.

Despite numerous studies of the Grizzly Bear in
west-central Alberta addressing specific research top-
ics using GPS collar data (¢.g.., Nielsen et al. 2002,
2003, 2004, 2006, 2010; Munro ef al. 2006; Berland

et al. 2008; Roever et al. 2008, 2010; Graham et al.
2010; Cristescu et al. 2011; Northrup et al. 2012; Stew-
art et al. 2012), basic biological data on home range,
movements and denning chronology have not yet been
presented. Earlier studies using Very High Frequency
(VHF) radio collars have provided estimates of home
range size, movements, and denning chronology:
however, improvements in the quality and quantity of
location data made possible by GPS collar technology
should increase our understanding of these character-
istics. We compare our results with those from earlier
studies in Alberta that relied on VHF technology, as
well as studies throughout the circumpolar range of the
Grizzly Bear.

Study Area

We focused on two genetically distinct Grizzly Bear
populations in west-central Alberta (Proctor ef al. 2012):
the Yellowhead population unit (YPU; 53°14'S3"N,
117°25'12"W) and the Grande Cache population unit
(GCPU; 44°10'19"N, 77°13'44"W). The location of the
YPU includes southern Jasper National Park in the
Rocky Mountains (Figure |). The GCPU is located di-
rectly north of the YPU and covers the northern part
of Jasper National Park and the Wilmore Wilderness
area, both in the Rocky Mountains (Figure |). Eleva-
tions are highest (up to 2700 m) in the western portion

225

224

THE CANADIAN FIELD-NATURALIST

Vol. 128

ALBERTA

BRITISH COLUMBIA

Collar locations ]
Major highways
Alberta boundary
Protected areas

GB Population Units
Grande Cache pop unit

Yellowhead pop unit

0 25 3 a
ees Kilometers

ALBERTA

~

50 75

FIGURE |. Location data for Grizzly Bears (Ursus arctos) collected using Global Positioning System collars from 1999 to
2010 for the Yellowhead and Grande Cache population units in west-central Alberta, Canada.

of both areas and decline eastward to approximately

900 m. This elevation gradient results in a diversity of

ecosites (Beckingham ef al. 1996*) including alpine
subalpine meadows; forests dominated by coniferous
species consisting of Lodgepole Pine (Pinus contorta),
White Spruce (Picea glauca), Black Spruce (P. mari-
ana), or Englemann Spruce (P. englemannii); mixed
forests comprising conifers with Trembling Aspen
(Populus tremuloides) or Balsam Poplar (P. balsam-
ifera); and wet meadow complexes. Both locations in-
clude large cleared areas, seismic lines, pipelines, and
roads associated with timber harvesting, oil and gas
exploration and development, and open-pit coal min-

ing. A number of recreational activities also occur with-
in these areas, including hunting, trapping, use of all-
terrain vehicles, camping, hiking, and mountain biking.
A regulated spring hunt for American Black Bears
(Ursus americanus) occurred during the study. A lim-
ited-entry hunt for Grizzly Bears occurred during
spring 1999-2005, but ceased in 2006.

Methods
Captures

Grizzly Bears in the YPU and GCPU were captured
and radio-collared from 1999 to 2010 and 2003 to 20] 0,
respectively. Capture methods included the use of cul-

2014

vert traps, leg-hold snares, and aerial darting from a
helicopter. In 2010, we terminated the use of snares
because of potential long-term capture impacts (Cat-
tet ef al. 2003, 2008). From 2010, capture techniques
involved aerial darting and culvert traps with satellite
alarm systems, designed to minimize the amount of
time bears were confined in the trap. Aerial darting via
helicopter occurred in open areas, such as alpine/sub-
alpine meadows and logged areas. Most snares and cul-
vert traps were placed in forested areas where aerial
darting was not feasible. Sites were usually < 100 m
from a road or other access feature type; however, some
sites were accessible only by helicopter. Capture pro-
tocols were approved by the Canadian Council on Ani-
mal Care for the safe handling of bears and approved
annually by the University of Saskatchewan and the
Government of Alberta animal care committees.

Grizzly Bears were fitted with a GPS radio collars,
including Simplex, Tellus, or Tellus Satellite collars
(Televilt [pow Followit], Lindesberg, Sweden); or Ad-
vanced Telemetry Systems (ATS; Isanti, Minnesota,
USA) collars. All collars emitted a unique VHF radio
signal to locate the bear. From 1999 to 2005, Simplex
collars were used and typically programmed to obtain
a location every 4 h during the non-denning period.
Collars deployed in the spring were retrieved in the fall
of the following year (1.5 years later) using a remote
drop-off mechanism, which allowed retrieval without
the need for recapture. Improvements in Tellus collars
and battery life occurred in 2004. The new collars were
able to provide hourly locations during the non-denning
period over 1.5 years. Remote data downloads were
possible for all versions of Televilt collars, allowing
data to be obtained even if the collar was not retrieved
at the end of its life. In 2010, one bear was collared
with a Tellus Satellite GPS collar, and data from this
collar were remotely obtained from a service provider.
ATS collars were used from 1999 to 2008. As they
did not have the remote data retrieval option, collars
were retrieved using a remote drop-off mechanism at
the end of the battery life, typically 8 months. Grizzly
Bears were also fitted with a VHF ear tag transmitter
(ATS), so that they could be located for collar removal
in case the collar failed.

GRAHAM AND STENHOUSE: GRIZZLY BEAR MOVEMENTS AND DENNING

A premolar tooth was extracted from captured bears
and sent to a commercial laboratory for age determi-
nation based on cementum annuli counts (Matson e/
al. 1993*). Age classes used in our analysis included
adult (= 5 years) and independent sub-adult (2 to < 5
years). Bears younger than 2 years were not collared,
but were given an ear tag transmitter, for possible relo-
cation and capture in successive years. We recorded
whether captured females were accompanied by cubs
of the year (COY), yearlings, or older cubs (= 2 years
old).

Home range

We calculated annual 100% minimum convex poly-
gons (MCP) and 95% kernel home ranges using the pro-
gram ABODE (Laver 2005*) in a Geographic Infor-
mation System (GIS). We determined both MCPs and
95% kernels (Worton 1987) to allow comparisons with
other studies. Fixed biweight kernels (Sliverman 1986)
were calculated using a volume contouring method.
We used a least-squares smoothing factor (Seaman and
Powell 1996) and a grid cell size of 300 m?. Kernels
were standardized using the unit variance method (Sil-
verman 1986).

We included a home range estimate only if data loca-
tions were available from May to October to ensure that
the entire year was represented (Arthur and Schwartz
1999: Belant and Follmann 2002; Girard et al. 2002).
In addition, each home range estimate required a mini-
mum of 100 days of location data or it was excluded
from the analysis (Arthur and Schwartz 1999; Belant
and Follmann 2002). If a bear generated more than one
year of location data for the same age or reproductive
class, only the data for the year with the greatest num-
ber of locations were used. We determined mean 100%
MCP and 95% kernel home range estimates for seven
age—sex—reproductive classes: female with COY, fe-
male with yearlings, females with > 2 year olds, lone
adult females, sub-adult females, sub-adult males, and
adult males (Table 1). Only the 95% kernel estimates
were statistically compared among the seven age—sex—
reproductive classes.

Movement rates
We used Visual Basic in Access Microsoft (version
2003) to determine the distance between successive

TABLE 1. Home range estimates, calculated as mean 100% minimum convex polygons (MCP) and 95% kernels, for Grizzly

Bears

(Ursus arctos) in west-central Alberta, by age-sex-reproductive class based on Global Positioning System collar

data. Kernel estimates for classes with different numbered superscripts are statistically different (P < 0.05) based on non-

parametric multiple comparison tests.

Age-sex— n

reproductive class

Female with COY 1] 370 (250,
Female with yearling 9 735 (822,
Female with > 2 year old i 722 (464,
Sub-adult female 14 732 (376,
Lone adult female 22 615 (311,
Sub-adult male .

Adult male

Mean annual 100% MCP
(SD, range), km?

2152 (1469, 509-4993)
1824 (1006, 336-3154)

Mean annual 95% kernel
(SD, range), km?

119-1025) 213 (212, 62-808)!
267-2904) 472 (527, 136-1848)!
202-1554) 494 (428, 105-1397)!"
222-1447) 394 (231, 86-836)!
200-1260) 337 (176, 107-706)!

1298 (1207, 3544282)
1034 (656, 203-2071)

EE eee

locations and calculate hourly movement rates. We
chose |-h time units because these were our most fine-
scaled temporal data and hourly readings were possi-
ble after 2004 using improved collars that allowed a
collar to last 1.5 years at an hourly fix rate during the
non-denning period. We only measured movements
outside the denning period because previous research
has shown that Grizzly Bear movements are reduced
before den entry (Nelson e¢ al. 1983; Friebe e¢ al. 2001;
Manchi and Swenson 2005) and immediately after den
exit (Craighead and Craighead 1972; Nelson ef al.
1983; Schwartz et a/. 2010). To exclude the denning
period, we removed data locations within 500 m of
known den sites and within an average of 7 days of
den entry and exit dates (see below). Also, the move-
ment of many Grizzly Bears is reduced for up to a
month after a capture event (Cattet et a/. 2008); thus,
location data collected within 30 days of a capture were
also removed from analysis. For movement rates, we
pooled females with yearlings and older cubs into one
class (females with yearlings+) to increase sample size
and produce six age-sex—reproductive classes.

Denning chronology

We determined den entry and exit dates and time in
the den from collar data. Collars were programed to
acquire at least one location every day during the den-
ning period because the manufacturer recommended
keeping the batteries active rather than shutting them
off completely for months at a time. Typically, when
bears entered their dens, GPS collars were unable to
acquire a location even though an attempt was made;
therefore, the day the collar consistently stopped re-
cording locations was considered to be the day the
bear entered the den. Den exit dates were determined
in a similar fashion, as the day the collar began to sig-
nal locations consistently again in the spring. Time in
the den was calculated based on data from bears with
known entry and exits dates for the same denning peri-
od.

Statistical analysis

We tested for normality and homoscedasticity using
a Shapiro-Wilk test for normality and a Bartlet’s test
for equal variances as well as visual examination of
standardized normal probability plots. When data were
not normal, appropriate transformations were applied or
nonparametric analyses were used. We used a Krusal-
Wallis test to determine whether mean kernel sizes dif-
fered among age—sex—reproductive classes and sub-
sequent nonparametric multiple comparison tests to
determine which means were significantly different
from others (Zar 1984). We graphed the hourly move-
ment rates for each age-sex—reproductive class by hour
of the day and month of the year. We used a mixed
effects multiple linear regression on log transformed
hourly movement rates with bears as the random effects
factor to determine whether movement rates differed
across the fixed effect factors of age-sex—reproductive
class, activity period, and month. We then conducted

THE CANADIAN FIELD-NATURALIST

Vol. 128

multiple comparisons of the marginal means for the
fixed effect factors using a Bonferroni adjustment to
determine where differences occurred. We used
ANOVA to compare den duration across age—sex—
reproductive class followed by a Tukey post-hoc mul-
tiple comparison to determine which classes were dif-
ferent from others. Statistical analyses were conducted
using Stata SE (v. 12.0 for Windows; StataCorp LP,
College Station, Texas). We used an alpha level of 0.05.

Between 1999 and 2010, we captured 40 females
and 31 males from the YPU and 18 females and 36
males from the GCPU. A total of 251 capture events
took place. Individual bears were captured on aver-
age 2.5 + 1.8 times in the YPU and 1.4 + 0.8 times in
the GCPU. A total of 53 Grizzly Bears from the YPU
and 46 from the GCPU were collared. Collars func-
tioned on bears from the YPU for an average of 476
+ 445 days, range = 13-2025 days) and 346 days on
bears from the GCPU (SD 396 days, range 3—2072
days). In total 659 744 GPS locations were collected
during this period.

Home range size

We estimated 97 annual 100% MCP and 95% kernel
home range sizes for 59 Grizzly Bears, using location
data collected from 1999 to 2010. We included 21 bears
more than once because their age or reproductive class
changed over time. A mean of 1691 locations (SD 1962,
range 203-9804) were used in the MCP and kernel esti-
mates. The mean kernel size across the seven age—sex—
reproductive classes differed significantly (H, = 32.31,
P < 0.001; Table 1). Kernel sizes of adult and sub-
adult males were not significantly different from each
other (P > 0.05, Table 1) or from those of sub-adult
females (P > 0.05), females with yearlings (P > 0.05)
or females with > 2-year-old cubs (P > 0.05), but they
were significantly larger than females with COY (adult
males: 0, = 4.56, P < 0.05; sub-adult males: QO. = 4.82,
P < 0.05) and lone adult females (adult males: QO, =
3.33, P< 0.05; sub-adult males: O, = 3.64, P < 0.05).
There were no differences in home range size among
the five female age—reproductive classes (P > 0.05,
Table 1).

Movements

Hourly location data from 39 Grizzly Bears in the
YPU and GCPU provided 87 959 hourly movement
rates. Except for females with COY, mean movement
rates for all other age—sex—reproductive classes tend-
ed to be greatest in June; the greatest movement rates
for females with COY occurred in August (Figure 2).
Movement rates generally declined after June, although
adult males showed an increase in September and Oc-
tober (Figure 2). Movement rates by hour of the day
showed a bimodal pattern, with four distinct activity
periods (Figure 3). Bears travel slowly at night (2200-—
0500), quicker during the morning (0600-1100) and
evening (1700-2100), and moderately in the afternoon
(1200-1600). These four activity periods were used in
the regression analysis.

2014 GRAHAM AND STENHOUSE: GRIZZLY BEAR MOVEMENTS AND DENNING

Zal
600
wok
¢ ‘
500 —— 7‘
4 al : ‘
4 ,
o” NS
* e | ~~ srenerases
= 400 reese
E
7)
e AdF_coy
5 —t— AdF
o _yearling+
E 300
$ —— SubF
=
2 200 ---@--- SUBM
—<—-AdM
100
0
May Jun Jul Aug Sep Oct
Month

FiGURE 2. Mean rates of movement by month for six age-sex—reproductive classes of Grizzly Bears (Ursus arctos) in west-
central Alberta. AdF_coy = females with cub of the year, AdF_yearling+ = females with yearling or older cub, SubF
= sub-adult females, AdF = lone adult females, SubM = sub-adult males, AdM = adult males.

TABLE 2. Results for the fixed effect factors (age-sex—reproductive class, month and activity period) used in the multiple
linear regression analysis to explain hourly movement rates (grouped by bear). Lone adult females, the month of May and
the morning activity period were reference categories. Data were log transformed to accentuate heteroscedasticity.

Age-sex— Standard
Factor reproductive class Coefficient deviation Zi 12 ed 95% confidence interval
Class Female with coy —0.0742 0.0128 =5.79 0.000 —0.0993 —0.04905
Female with yearling+ 0.0419 0.0105 B96 0.000 0.0211 0.0626
Sub-adult female 0.1293 0.0098 18222) 0.000 0.1101 0.1485
Sub-adult male 0.0157 0.0415 0.38 0.704 —0.0655 0.0970
Adult male 0.0839 0.0390 2AIS 0.032 0.0074 0.1604
Month June 0.1291 0.0099 13.03 0.000 0.1097 0.1486
July 0.1858 0.0091 20.46 0.000 0.1680 0.2036
August 0.1578 0.009] 17.28 0.000 0.1399 0.1756
September 0.0680 0.0093 7.29 0.000 0.0497 0.0863
October —0.1326 0.009] —14.55 0.000 =Va505 —0.1147
Activity Afternoon —0.1606 0.0068 PSS) 0.000 —0.1740 -0.1473
period Evening 0.0828 0.0068 ly 0.000 0.0694 0.0961
Night —0.5367 0.0062 =souLy) 0.000 0.5489 —0.5245
2.0634 0.026] 79.09 0.000 2.0122 2.114

Intercept
oo nn OC—————a—

Age-sex—reproductive class (six classes), activity movement rate occurred in June and the slowest in
period (morning, afternoon, evening, and night) and — October. Likewise, all four activity periods were signif-
month (May to October) were significant predictors of icantly different from each other (Z, < 94.88, P < 0.01
hourly movement rates (Tables 2 and 3). Comparison for all six comparisons; Table 5). Bears moved fastest
of marginal means indicated that all 6 months were _ in the evening followed by the morning and afternoon,
significantly different from each other (Z, < 40.57, PS with the slowest mean movement rate occurring at
0.01 for all 15 comparisons; Table 4). The fastest mean night.

228 THE CANADIAN FIELD-NATURALIST Vol. 128

=

=

£

r AdF_coy
= —+«— AdF_yearling+
o

é —— SubF

£ AdF

c

s ---@--- SuUDM

= - ¢@-AdM

O 123 4 5 6 7 8 9 10 41 12 13 14 15 16.47 18 19 20 23 27 2
Hour of the day

FiGurE 3. Mean rates of movement by hour of the day for six age-sex—reproductive classes of Grizzly Bears (Ursus arctos)
in west-central Alberta. AdF_ coy = females with cub of the year, AdF_yearling+ = females with yearling or older
cub, SubF = sub-adult females, AdF = lone adult females, SubM = sub-adult males, AdM = adult males.

TABLE 3. Results for bear as the random effect factors used in the multiple linear regression analysis to explain hourly movement
rates. Data were log transformed to accentuate heteroscedasticity.

Random effects for bear Estimate Standard deviation 95% confidence interval
Standard deviation of intercept 0.1117 0.0139 0.0874—0.1426
Standard deviation of residuals 0.6903 0.0016 0.6871—0.6935

TABLE 4. Mean hourly movement rates of Grizzly Bears (Ursus arctos) in west-central Alberta by month. Post-hoc comparisons
with Bonferroni adjusted P values are provided.

Mean movement

Month n- rate, m/h (SD)* June July August September October

May 10 657 287 (487.7) Zig 3.08 Z, = 20.46 Z, = 17.28 Lig Tee) AAS
P<0.001 P<0.001 P<0.001 P<0.001 P<0.001

June 10 298 391 (602.3) Zh = (0) 3S) Ly = Sho Z,= S63 Z=-DB88
P<0.001 2 O100T P<0.001 P<0.001

July 17 429 346 (473.5) Loe=—3.38 L,=—15 14 Z,=—40 57
P<0.001 P<0.001 P<0.001

August 18 132 343 (478.9) Zea: Lg——s vor
P<0.001 P<0.001

September 15 461 320 (497.7) 2.2505
P<0.001

October 15 982 251 (497.8)

Cee
*SD = standard deviation.

2014

GRAHAM AND STENHOUSE: GRIZZLY BEAR MOVEMENTS AND DENNING 229

TABLE 5. Mean hourly movement rates of Grizzly Bears (Ursus arctos) in west-central Alberta by activity period. Post-hoc
comparisons with Bonferroni adjusted P values are provided.
.35.0R?COoaewNoqooqoqonWw—~——eee

Activity period n Mean movement rate, m/h (SD)* Afternoon Evening Night
Morning 22 526 403 (541.8) Z,=-23.55 Z,=12.17 Z,=-86.19
: P< 0.001 P< 0.001 P< 0.001
Afternoon 18 863 296 (447.3) Z, = 34.28 Z, = 57.29

é P< 0.001 P< 0.001
Evening 19 032 448 (563.7 Z, = —94.88
P< 0,001

Night 27588 185 (420.7)

SoS 0—="0—_—=™——6sSOeooeoeo—
*SD = standard deviation.

TABLE 6. Mean hourly movement rates of Grizzly Bears (Ursus arctos) in west-central Alberta, by age—sex—reproductive class.
Post-hoe comparisons with Bonferroni adjusted P values are provided. Non-significant (P > 0.05) differences are in bold.

eee
V———OOOO——————eeeeeeee

Age-sex— Mean Female Lone
reproductive movement rate, with Sub-adult adult Sub-adult Adult
class n m/h (SD)* yearling+ female female male male
Female with COY 7 354 191 (317.8) = 928 Zy= 12:69 7B ba AS) Z,= 2.12 Lom aot
P<0.001 P<0.001 P<0(0.001 P=0.515 P=0.001
Female with yearling+ 14 645 297 (449.6) ZO Zo 9.96 Z, = —0.62 Z,— 1.06
P<(0.001 P=0.001 P= 1.000 P= 1.000
Sub-adult female 15 048 366 (506.4) L3= B22) “Zia 75-4
P<(0.001 P=0.102 P= 1.000
Lone adult female 28 160 292 (459.6) Z, = 0.38 Z,=2.15
P=1.00 P=0.47
Sub-adult male 7 584 342 (522.3) Li 23
P=0.128
Adult male 15 168 408 (649.7)
*SD = standard deviation.
Den entry Month of the year Den exit Mean denning duration
Age Sex Reproductive class No.bears Oct Nov Dec Jan Feb Mar Apr May No. bears No. days No. bears sD
es dizsdizsdiesdicsdiasdizs 4
as WH = =
Adult F With cubs 21 || 16 140 13 15.6
some me | 7 e 4 ®
Adult F __Alone/unknown 17 me || | 14 147 13 0.0
Sub-adult M Na 8 5 134 5 0.0
Aduit M Na 13 11 144 10 0.0

FiGuRE 4. Den entry and exit dates by quarter month and age-sex—reproductive class for Grizzly Bears (Ursus arctos) in
west-central Alberta, determined from Global Positioning System collar data from 1999 to 2010. Small black
squares indicate mean den entry or exit dates, light shading indicates the range of entry and exit dates and dark shad-
ing indicates when all bears were in their dens. Den time was longer for pregnant females than for the other five
classes (Tukey post-hoc multiple comparison test; P < 0.002) and is indicated by an asterisk. Females with cubs rep-
resent females accompanied with any aged cub at den entry.

P > 0.05) while sub-adult males were not different from
any of the age-sex—reproductive classes (Z, < 2.70,
P > 0.05). Females with COY had the slowest move-

Adult males and sub-adult females had the fastest
mean movement rates (408 m/h and 366 m/h, respec-
tively) but adult and sub-adult males had the largest

standard deviations (650 and 522, respectively; Table
6). Comparisons of marginal means (Table 6) showed
that adult males were significantly faster than females
Wwith:COy (Z,—= 3.95; P=0.001) but not different from
the remaining age-sex—reproductive classes (Z, < 2.03;

ment rate, followed by lone adult females, females with
yearlings, and older and sub-adult females; all which
were significantly different from each other (Z, > 3.94,
P<.01).

Denning

Different age-sex classes exhibited different den-
ning chronology (Figure 4). Pregnant females entered
dens first, in early November, followed by lone adult
females, females with cubs of all ages and adult males
in close succession. Sub-adult males and females
entered their dens last, in late November (Figure 3).
Conversely, Grizzly Bears exited their dens in the re-
verse order with sub-adult males and females emerg-
ing first, in early April, followed by adult males, fe-
males with yearlings or older, lone adult females, and
finally, females with COY, in late April. Denning dura-
tion varied among age—sex—reproductive class (F’,
= 8.78: P< 0.001 ). Pregnant females had the longest
denning duration, significantly longer than lone adult
females (94) <= —4.04, P = 0.002), females with year-
lings and older (q,. ,= —4.99, P < 0.001), adult males
(4606 = ~4-17; P = 0.001), sub-adult females (q,, , =
—5.04, P< 0.001), and sub-adult males (G60. =—-4.27;
P = 0.001). There were no significant differences in
length of denning duration among the other age—sex—
reproductive classes (P > 0.05). Pregnant females spent
the longest time in dens, (approximately 48% of the
year), whereas sub-adults of both sexes spent the least
amount of time in dens, approximately 38% of the year)
(Figure 3).

Discussion

The age and sex of a Grizzly Bear are important fac-
tors influencing home range size and rates of move-
ment throughout their circumpolar range (Pearson
1975*; Blanchard and Knight 1991; Mace and Waller
1997; McLoughlin ef al. 1999; Dahle and Swenson
2003a; Stevens and Gibeau 2005; Ciarniello et al. 2009;
Edwards ef al. 2009). Adult males typically have large
home ranges and move quickly, probably, in part,
because they are searching for reproductive females
(Blanchard and Knight 1991; Dahle and Swenson
2003a; Krofel et a/. 2010). Our results support this
searching behaviour: the fastest movements by adult
males occurred in June, which corresponds to the peak
mating period in our area (Stenhouse ef a/. 2005).
Large ranges of movement by adult males have also
been explained by individuals travelling more in search
of food to support:a large body mass (McNab 1963;
McLoughlin et al. 1999) and this could also be the rea-
son for the increase in movement rates observed among
adult males in the fall during our study. Fall is the peri-
od of hyperphagia (Nielsen et a/. 2004); thus, adult
males may travel more at this time in search of food
to fatten up before denning. Adult males may also be
actively gathering information on other bears within
their home range for the following year (Dahle and
Swenson 2003a). The relatively fast movements of
females with yearlings and older cubs compared with
those of lone adult females may also be a result of
searching for food to satisfy the energy demands of the

THE CANADIAN FIELD-NATURALIST

Vol. 128

female and her growing cubs (Blanchard and Knight
1991).

The slow movements of females with COY have
been attributed to the reduced mobility of the COY and
an avoidance of males to reduce possible infanticide
(Blanchard and Knight 1991; Dahle and Swenson
2003a,b; Steyaert 2012). However, home range size of
females with COY was not significantly different from
that of other female age-reproductive classes. This is
in contrast to findings for Grizzly Bears in Sweden
(Dahle and Swenson 2003a), but similar to those in
the Northwest Territories (McLoughlin et al. 1999).
Dahle and Swenson (2003b) found that spring ranges
of females with COY in Sweden were small, but ex-
panded once the breeding season was over. Although
we did not examine seasonal ranges, our mean monthly
movement rates suggested that movements increased
each month until August when rates were similar to
those of lone adult females. We cannot say whether
movement rates in May were slow because of limited
COY mobility or to avoid infanticidal males, but home
ranges of females with COY likely expand throughout
the summer until they are similar in size to those of
other female age—reproductive classes.

Our home range estimates for male and female sub-
adults were based on whether the bear was independent
and < 5 years of age. Because we did not distinguish
between dispersing and philopatric individuals, our esti-
mates likely included dispersing bears and, therefore,
did not conform to the traditional definition of a home
range (Burt 1943). Because sub-adult males disperse
farther than sub-adult females (Blanchard and Knight
1991; Zedrosser et al. 2007), the likely inclusion of dis-
persing males resulted in a large home range estimate
and standard deviation and could explain why our esti-
mate was four times larger than the home range report-
ed for philopatric sub-adult males in Sweden (Dahle er
al. 2006).

The large variation in hourly movement rates by
adult and sub-adult males and the conservative nature
of the Bonferroni adjustment (Garcia 2004) could have
resulted in missing significant differences (Type II
error). The mean movement rate of sub-adult males was
not different from the other age-sex—reproductive class-
es, and adult males’ rate was only different from that
of females with COY. Other researchers have docu-
mented differences in movement rates between the
sexes (Blanchard and Knight 1991, McLoughlin eg al.
1999). Perhaps of more interest is the large variability
we found in male movement rates. Further work to look
at movements by males and site visits to areas where
different movement rates occurred are needed to under-
standing the observed differences.

The bimodal activity pattern displayed by Grizzly
Bears in our study was similar to that reported for
British Columbia (McCann 1991), Montana (Aune and
Kasworm 1989*), and Europe (Roth 1983: Roth and

2014

Huber 1986; Moe er al. 2007), but the mid-day inac-
tive period was longer and appeared to be the main rest
period for the European bears. In contrast, male Griz-
zly Bears in Wyoming were active throughout the night
and rested in mid-afternoon while females showed a
pattern similar to our adult females (Holm e¢ al. 1999),
Movement can be influenced by many factors includ-
ing season, cover, temperature, food availability, age,
sex, bear density, and human activities (Aune and Kas-
worm 1989*; McCann 1991; Dahle er a/. 2006). In the

future, the ability to compare movements with data of

similar quality and quantity within a population over
time and across populations may help us to better un-
derstand factors that influence movement rates.
Three studies conducted more than a decade ago
within our study area also reported home range esti-
mates (Pearson and Nolan 1976*; Russell et al. 1978*,
1979*; Horejsi and Stegenga 1981*; Horejsi and Slat-
ter 1982*; Horejsi and Raine 1983*; Nagy et al. 1989*;
Nagy and Haroldson 1990). These studies, which relied
on VHF collars to obtain a location on a weekly or bi-
weekly schedule, provided the foundation of our under-
standing of Grizzly Bear biology in Alberta. VHF-based
home range estimates were typically based on < 50
locations and bear locations were likely missed when
the animal had moved outside the survey area (Russell
et al. 1979,* Dahle and Swenson 2003a, Collin et al.
2005); therefore, these studies likely underestimated
the true home range size (Arthur and Schwartz 1999;
Girard et al. 2002). However, similar to our results,
their data showed that male bears were fast moving,
MCPs for males were larger than those for females,
and females with COY had the smallest MCPs.
Denning is believed to have evolved as a mechanism
to endure periods with little food (Manchi and Swen-
son 2005); however, the triggers that cause a Grizzly
Bear to enter and leave a den are not fully understood
(Friebe et a/. 2001). On a large scale, latitude explains
some of the variability in denning period across the
Grizzly Bear’s range (Manchi and Swenson 2005): in
Sweden (Manchi and Swenson 2005) and the Canadian
north (McLoughlin e¢ al. 2002) Grizzly Bears denned
10-30 days longer than in west-central Alberta, where-
as, in Yellowstone, den time was 10—30 days shorter
(Judd et al. 1986). However, latitude alone cannot ex-
plain all the differences in denning period as some
Grizzly Bears on Kodiak Island, Alaska, do not den at
all (Van Daele et al. 1989), and Grizzly Bears in British
Columbia, at a latitude similar to that of our study area,
denned more than 50 days longer (Ciarniello et al.
2005). Heavy snowfall (Craighead and Craighead 1972;
Servheen and Klaver 1983; Friebe e¢ al. 2001; Manchi
and Swenson 2005) has been associated with den entry
for some populations, while declines in food supply
may have influenced den entry in others (Clevenger ef
al. 1990: Ciarniello et al. 2009). Precise denning dates,
along with local weather and snow conditions, body

GRAHAM AND STENHOUSE: GRIZZLY BEAR MOVEMENTS AND DENNING

condition, and food supply information, are needed to
improve understanding of den ecology within and across
populations (K. Pigeon, unpublished data).

Denning duration has been related to a bear’s fat
stores prior to denning, its surface-volume ratio, den
type, and reproductive status (Craighead and Craighead
1972; Schwartz et al. 1987; Ciarniello et al. 2005;
Friebe et al. 2001; Manchi and Swenson 2005). A long
denning duration for pregnant females has been doc-
umented for Grizzly Bears throughout their circum-
polar range (Craighead and Craighead 1972; Ballard
et al. 1982; Ciarniello et al. 2005; Friebe et al. 2001;
Manchi and Swenson 2005). Early den entry by preg-
nant females may have more to do with reproductive
physiology than environmental factors (Hissa ef al.
1994; Friebe et al. 2001), and late den exit by females
with COY may be a result of waiting until travel con-
ditions are suitable for very young bears (Craighead
and Craighead 1972). Small, young male bears had a
longer denning duration than old, large males in south-
ern Sweden (Manchi and Swenson 2005), while sub-
adult females had a shorter denning duration than adult
females (Friebe et a/. 2001). We did not find a differ-
ence in denning duration among our sub-adults and
adults. However, the sample size for sub-adult males
was small and post-hoc multiple comparisons can in-
crease the chance of conducting a Type II error (Zar
1984). Therefore, it is possible that we failed to recog-
nize a significant difference among some age—sex—
reproductive classes. Further work to examine the den-
ning biology of Grizzly Bears in Alberta is currently
underway.

Denning dates from the VHF studies that overlapped
our study area were available for Jasper National Park
(Russell et a/. 1979*) and the Wapiti River area (Hore-
jsi and Raine 1983*). These dates were often inexact,
and denning duration was difficult to determine when
spring and fall monitoring flights were missed due to
inclement weather. However, even with inexact dates,
researchers conducting the VHF-based studies detect-
ed a chronological order for entering and exiting dens
that was similar to our findings decades later (Russell
et al. 1979*; Horejsi and Raine 1983*). GPS collars
allowed us to determine precise denning dates and
should allow researchers to identify changes in denning
duration that may be important for detecting impacts
of climate change on Grizzly Bear den biology in the
future.

Conservation implications

GPS collars allowed biologists to collect large data-
sets on movements of individual bears, to determine
accurate estimates for home range, and to determine
hourly movement rates and precise denning dates.
These data permit biologists to compare the biology of
the Grizzly Bear across its range and test hypotheses
not possible with VHF data. Although VHF-based stud-
ies provided home range estimates, they could under-

estimate the true area and movement data were limited.
Denning chronology by age-sex—reproductive class
was recognized in VHF-based studies, but precise dates
were lacking. VHF-based studies required biologists
to locate their animals visually, which often involving
dangerous low-level flying, but offered opportunities
for direct observations. GPS collars provide accurate
and abundant data, but ethological observations are
often minimal. Observations of individual animals and
site visits to GPS locations are crucial for interpreting
GPS telemetry data and should be an important com-
ponent of any research project that employs GPS col-
lars on wildlife.

Acknowledgements

Our study was supported financially by the numer-
ous partners of the Foothills Research Grizzly Bear
Program, which includes oil and gas industry partners,
forest industry partners, the Alberta government, and
the Government of Canada, as well as various non-
governmental organizations. We thank the bear biolo-
gists and veterinarians who captured and collared Griz-
zly Bears over the years and those who collected GPS
data on the ground and in the air. We also recognize the
many highly skilled pilots who continued to bring us
home safely after each data collection flight. We thank
Julie Duval for maintaining our ever-growing Grizzly
Bear database. Thank you to Karine Pigeon, PhD Can-
didate at Laval University, in association with the
Foothills Research Institute for her comments in the
denning section of this paper. Thanks to T. Jung and an
anonymous reviewer for the helpful and thoughtful
suggestions regarding drafts of this paper.

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Aune, K., and W. Kasworm. 1989. East front Grizzly Bear
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20 and%20Kasworm%201989.pdf.

Beckingham, J. D., I. G. W. Corns, and J. H. Archibald.
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THE CANADIAN FIELD-NATURALIST

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Received 13 February 2013
Accepted | October 2013

Mortality of Common Eider, Somateria mollissima (Linnaeus, 1758),

and other Water Birds during two Inshore Oiling Events in Southeastern
Newfoundland, 2005 and 2006

GREGORY J. ROBERTSON! », ScoTT G. GILLILAND!, PIERRE C. RYAN!, JOHANNE DUSSUREAULT!, KYRAN
POWER-, and BRUCE C. TURNER!

lif pracy ae : ‘ne Ci. ~

C anadian Wildlife Service, 6 Bruce Street, Mount Pearl, Newfoundland and Labrador AlN 4T3 Canada
“P.O. Box 21, St. Mary’s Bay, Newfoundland and Labrador AOB 1E0 Canada

‘Corresponding author: greg .robertson@ec.ge.ca

Robertson, Gregory J., Scott G. Gilliland, Pierre C. Ryan, Johanne Dussureault, Kyran Power, and Bruce C. Turner. 2014.
Mortality of Common Eider, Somateria mollissima (Linnaeus, 1758), and other water birds during two inshore oiling
events in southeastern Newfoundland, 2005 and 2006. Canadian Field-Naturalist 128(3): 235-242.
Although the waters off Newfoundland harbour millions of wintering marine birds, chronic marine oil pollution has been
repeatedly reported. Unusually high numbers (hundreds) of oiled birds were noted following two events in March 2005 and
April 2006 in southeastern Newfoundland. Common Eiders (Somateria mollissima [Linnaeus, 1758]) were the main victims in
the first event, with at least 1400 affected, based on retrieval of carcasses and aerial surveys. The April 2006 event affected
19 species; Common Eiders were again the most numerous with a minimum of 337 birds oiled. Among the Common Eiders
affected in both events, most were the northern type, including the borealis (C. L. Brehm, 1824) subspecies and presumed
intergrades between borealis and dresseri Sharpe, 1871. Coupled with the legal harvest, these oiling events may have had an
effect on the wintering Common Eider population. Alcids, other sea ducks, loons and gulls were also oiled, but in low numbers
(< 100); thus, their populations were not likely affected by these events.

Key Words: Oil pollution; Common Eider; Somateria mollissima; water birds; Avalon Peninsula; Newfoundland

Introduction spills. To help understand potential effects on popula-
The cold, productive waters off Newfoundland and _ tions, we present information on oiled carcasses to
Labrador, Canada, are home to large populations of — assess the population structure and affinities of birds
breeding and wintering marine birds (Lock et al. 1994). _ killed. Finally, we evaluate the potential effect of these
However, chronic oiling problems have been document- __ two oiling events on eiders and other water birds win-
ed in the coastal waters of Newfoundland (Wiese and __ tering in Newfoundland.
Ryan 2003), likely because of the large numbers of
ships that transit these waters. Although murres (Uria Study Area and Methods
spp.) and Dovekies (Alle alle [Linnaeus, 1758]) are © March 2005 incident
most severely affected by oiling, other species, includ- On Saturday, 26 February 2005, wildlife officials
ing sea ducks, are also found (Wiese and Ryan 2003). and pollution prevention officers received reports that
Among the coastal wintering species, the Common _ oiled eiders were being taken during the legal hunt.
Eider (Somateria mollissima [Linnaeus, 1758]) is the Further reports were received on Monday, 28 February
most numerous and also an important game bird in 2005, the final day of the open hunting season for sea
Newfoundland (Wendt and Silieff 1986; Gilliland and — ducks in Newfoundland. On | March, an experienced
Robertson 2009). Unlike pelagic seabirds, which tend observer (S.G.G.) used a Canadian Coast Guard heli-
to be widely distributed, Common Eiders may form — copter (BO-105, Boélkow, Stuttgart, Germany) to sur-
dense flocks, and a single oiling event can affect a large vey the east coast of the Avalon Peninsula from Cape
component of the population. Oil spills affecting Com- St. Francis to Cape Race (Figure 1) and count the num-
mon Eiders have occurred in Atlantic Canadian waters __ ber of eiders at risk in the area and visibly oiled eiders.
in the past (Brown et al. 1973). The survey began and ended at Cape St. Francis, begin-
Two significant events involving hundreds of oiled ning at 1030 and ending at 1345. Survey conditions
coastal birds occurred from late February to early April — were excellent: light easterly winds, clear sky, and a
2005 and April 2006 in southeastern Newfoundland (a __ temperature ot) Cc. The flight track and all observa-
third event in November 2004 involving mostly pelagic tions were recorded using United States Fish and Wild-
species is described elsewhere [Robertson ef al. 2006]). life Service Global Positioning System Voice Software.
In this paper, we present information from a variety of During the initial fight, from north to south at 500 m
surveys conducted to assess the geographic scope of above sea level, the size and location of all large flocks
these incidents and estimate the number of eiders and _ of eiders and Black-legged Kittiwakes (Rissa tridactyla
other water birds at risk and actually oiled from these _ [Linnaeus, 1758]) were recorded. On the return flight,

i)
ee)
‘Nn

Avalon Peninsula =

47°N

Cape St/Mary’s * bi

Ecological Reserve's: V0"

wane

Nov. 2004

March 1970

THE CANADIAN FIELD-NATURALIST

Vol. 128

| Cape
|St. Francis

(LU }

@ / e

4° St. John’s 4

1 | Blackhead

A,
| f
_|

Witless Bay Islands
Ecological Reserve

, St. Michael's
March 2005

Ferryland

FIGURE 1. Extent of two oiling events in southeastern Newfoundland, March 2005 (solid line) and April 2006 (dashed line).
The locations of two other events mentioned in the text are also shown: Irving Whale, March 1970 (arrow: Brown et
al. 1973) and November 2004 (dotted line; Robertson et al. 2006).

the focus was on birds that may have been oiled. This
survey was conducted from south to north at 35 m
above sea level, and the number of eiders loafing on
beaches and rocks was recorded; in some cases oiling
was visibly apparent on these birds. The number and
location of Bald Eagles (Haliaeetus leucocephalus
[Linnaeus, 1766]) that appeared to be scavenging oiled
eiders was also noted.

On 6 March 2005, a second survey was conducted
specifically to count the number of eiders that appeared
oiled. This survey included simultaneous air and ground
counts to correct for observer bias in estimating flock
sizes (see Bordage er al. 1998) and to determine wheth-
er eiders were indeed oiled. Fifteen ground count sites
were identified at 4 locations: Cape St. Francis, Black-
head, St. Michael’s, and Ferryland (Figure 1). On 5
March 2005, the primary helicopter surveyor (S.G.G.)
visited the ground sites with each observer, who would
be counting eiders the next day, to ensure that the same
geographic areas were being assessed simultaneously
from the air and the ground. Observers on the ground

counted eiders every 30 minutes at their site. As it was
apparent that eiders flushed with the arrival of the heli-
copter, the previous count was used. Helicopter survey
protocols were similar to those used on the return
portion of the 1 March survey, except a 206 LR (Bell
Helicopter Trextron Canada Ltd, Mirabel, Quebec,
Canada) from Universal Helicopters (Goose Bay,
Newfoundland and Labrador) was used for the portion
of the flight from Logy Bay (near St. John’s) to Cape
Bonavista (48°42'N, 53°05'W) to the north. The flight
ran south to north and took place from 0850 to 1430.
Survey conditions again were generally good: light
winds, clear sky, and —12°C:; however visibility was
restricted by local snow squalls. The ratio of ground
count to aerial count was calculated for each site where
birds were present, and the mean of those ratios was
used to correct the aerial counts.

From 26 February to 7 April 2005, crews searched
beaches daily for oiled bird carcasses. In addition,
birds (and parts) were received from the public, mostly
hunters, in late February.

2014

We examined all carcasses received, recording the
species, sex, age where possible, degree of scavenging,
and degree of oiling. We measured body mass and com-
monly recorded morphometrics (wing, tarsus, and cul-
men length) of intact carcasses. For Common Eiders,
we also measured aspects of the bill used in subspecific
identification (Mendall 1986) and assessed visually
whether the bird was borealis, dresseri Sharpe, 1871,
or an intergrade between these subspecies based on bill
shape and colouration (Goudie er a/. 2000). Given the
uncertainty of distinguishing the borealis subspecies
from intergrades, the category “northern-type common
eider,” which included borealis and intergrades, was
used (Gilliland and Robertson 2009). To determine
whether oiled eiders had lost body condition before
being found, we measured the body mass of 60 Com-
mon Eiders shot between December 2004 and Febru-
ary 2005 near Point Lance, Newfoundland (near Cape
St. Mary’s) to compare with the mass of oiled carcass-
es. Even though this sample was collected earlier in
the winter than the oiled birds, they should be compa-
rable as northern Common Eiders do not show strong
seasonality in body mass dynamics during the winter
(Jamieson et al. 2006).

April 2006 incident

On 9 April 2006, reports were received from the pub-
lic of oiled birds, including gulls, auks, and Long-tailed
Ducks (Clangula hyemalis [Linnaeus, 1758]) sighted
along the southeast coast of the Avalon Peninsula,
near Cape Pine and the town of Trepassey (Figure 1).
Response teams were deployed to examine beaches
from 10 to 18 April. On 12 April, two observers (S.G.G.
and G.J.R.) conducted a shoreline survey in a 206 LR
helicopter to look for oiled sea ducks, following pro-
tocols similar to those described for the March 2005
survey. Skies were clear and survey conditions were
very good.

As the birds affected by this oiling event were a
varied mix of coastal and pelagic species, a minimum
number for each affected species was derived from a
variety of sources: for waterfowl, the birds counted on
shore on 12 April and any carcasses collected before
that date were summed; for species not well assessed
by aerial surveys, the number of dead carcasses collect-
ed, plus reliable observations of oiled birds from areas
outside the beaches regularly surveyed were summed.
Birds were measured as described above, but a visual
assessment of the subspecific status of Common Eiders
was not conducted.

Results
March 2005 incident

We estimated that on | March 2005, there were ap-
proximately 42 000 Common Eiders along the eastern
Avalon Peninsula of Newfoundland; this represented
4 raw count and was not corrected for observer bias.
Assuming that all eiders that were hauled out on beach-
es and rocks were oiled, we estimated the number of

ROBERTSON ET AL.: OILED EIDERS IN NEWFOUNDLAND

P|

oiled eiders at 1165, Major concentrations of birds were
located on islands, which are relatively sparse along
this coast, with most on the larger islands in the Witless
Bay Islands Ecological Reserve (Figure |). In addition
to the eiders, we observed at least two oiled Long-tailed
Ducks, and one dead Northern Gannet (Morus bas-
sanus [Linnaeus, 1758]). During the 6 March 2005
helicopter survey, we counted 537 oiled eiders along
the eastern Avalon Peninsula. Once again, major con-
centrations were found on the islands of the Witless
Bay Islands Ecological Reserve, and extended further
north to include Cape St. Francis (Figure 1). In addition
to the eiders, we observed at least two oiled Long-
tailed Ducks.

We successfully conducted simultaneous air and
ground counts at four locations (Table 1). The ground
counts were slightly higher than the corresponding aer-
ial counts (ratio 1.07, standard error [SE] 0.14); using
this ratio to adjust for errors in the aerial counts, we esti-
mated that 577 (SE 75) eiders may have been affected
by oil on 6 March. Based on other casual observations
of oiled eiders at sites that were not detected by the
helicopter, we believe this is an underestimate.

TABLE 1. Simultaneous aerial and ground counts of oiled
Common Eiders (Somateria mollissima) at four loafing loca-
tions, 6 March 2005, in Newfoundland.

Location (site) Ground count Aerial count

Cape St. Francis 84 85
Blackhead 6 0
St. Michael’s
Site | | l
Site 2 0 0
Site 3 0 0
Site 4 22 9
Site 5 0 0
Cribbies 2 2
Ferryland
Site |
Site 2
Site 3
Site 4 0
Site 5 0 0
Site 6 l |
Total 121 108

Most of the carcasses retrieved during this incident
were Common Eiders, although carcasses of another
five species were also retrieved (Table 2). In addition,
another four species were seen oiled: Purple Sand-
piper (Calidris maritima {Brunnich, 1764]), Common
Loon (Gavia immer [Brunnich, 1764]), American Her-
ring Gull (Larus smithsonianus Coues, 1862), and
Long-tailed Duck. Based on visual inspection, most
eiders found were of the northern (borealis) or inter-
mediate-northern type (males: 92%, n = 36; females:
92%. n = 25). However, using Mendall’s (1986) key,
more males were classified as dresseri (42%, n = 36).

THE CANADIAN FIELD-NATURALIST

Vol. 128

TABLE 2. Species and ages of oiled birds collected on beaches or provided by hunters after an oil spill incident in southeast-

ern Newfoundland, March 2005.

Immature Adult Unknown age Total
Common Eider (Somateria mollissima)
Female 4 29 1] 44
Male 16 2 4 48
Thick-billed Murre (Uria lomvia) ) 3 5
Dovekie (Alle alle) | |
Black Guillemot (Cepphus grylle) 2) I :

Great Black-backed Gull (Larus marinus) |

Of the males, where more specific aging based on
plumage was possible, 28 (64%) were found to be
adults, 16 (36%) were sub-adults and none were juve-
niles. Female and male oiled eiders were 15% and 16%
lighter, respectively, than eiders shot earlier the same
winter (females: oiled 1445 g, SD 133 g, n = 24; un-
oiled 1694 g, SD 138 g, n = 32; males: oiled 1600 g,
SD 218 g, n= 33, unoiled 1904 g, SD 104 g, n= 28).
Live oiled eiders were seen well into April 2005 around
the eastern Avalon Peninsula, after which eiders left the
area on migration, indicating that this species may sur-
vive for over a month after oiling.

April 2006 incident

Between 10 and 16 April 2006, 186 oiled carcasses
of 11 species were collected on the southeastern tip
of the Avalon Peninsula (Table 3, Figure 1). The most
common species recovered were sea ducks (Common
Eider 18.6%; Long-tailed Ducks 12.9%) and auks
(murres 48.4%; Black Guillemots [Cepphus grylle

(Linnaeus, 758)] 13.4%; Dovekies and Atlantic Puffins
[Fratercula arctica (Linnaeus, 1758)] 1.1% each).
During the 12 April 2006 aerial survey, 409 birds were
seen on shore and assessed as oiled: these included
320 Common Eiders, 64 Long-tailed Ducks, 12 loons,
9 murres, 3 Black Guillemots, and a White-winger
Scoter (Melanitta fusca [Linnaeus, 1758]). The extent
of the distribution of Common Eiders presumed oiled
was smaller than that seen after the 2005 event (Fig-
ure 1). Combining the aerial survey with the carcass
recovery data and adding oiled birds seen outside the
main area of focus, we estimate that a minimum of
574 birds of 19 species were affected by this spill
(Table 4).

Among recovered Common Eiders, 21 of 22 birds
were adults (95.5%), but the sex ratio was close to even
(18 females of 34 birds, 52.9%) (Table 3). Similarly,
among Long-tailed Ducks, 18 of 21 (85.7%) birds were
adults, but 20 of 24 birds were male (83.3%). Based

TABLE 3. Species and ages of oiled birds recovered on beaches in southeastern Newfoundland, April 2006.

Juvenile
Species F M U

Common Loon l ]
(Gavia immer)

eg]

Northern Fulmar
(Fulmarus glacialis)

Common Eider | 6
(Somateria mollissima)

Long-tailed Duck 3 3
(Clangula hyemalis) .

American Herring Gull

(Larus smithsonianus)

Common Murre 2 12 |
(Uria aalge)

Thick-billed Murre l 6 5
(Uria lomvia)

Murre (Uria spp.)

Dovekie (Alle alle)

Black Guillemot | |

(Cepphus grylle)

Atlantic Puffin

(Fratercula arctica)

Adult Unknown
M U F M U Total
| 2 l 6
l l
15 1] l 34
15 l 2 24
2 2
l 13 4 33
4 28 2 46
1] 1]
2 2
| 7 1S aS

Note: F = female, M = male, U = unknown.

2014

TABLE 4. Minimum numbers of birds by species affected by
an oiling incident in southeastern Newfoundland, April
2006. Numbers are totals of birds seen ashore during aerial
surveys (ducks), found before the aerial survey, found after
the survey and not counted during the aerial survey (auks),
and oiled birds reported outside the survey area.

Minimum
Species number oiled
Common Eider (Somateria mollissima) 337
Murre (Uria spp.) 98
Long-tailed Duck (Clangula hyemalis) 66
Black Guillemot (Cepphus grylle) 29
Loon (Gavia spp.)* pee
American Herring Gull (Larus smithsonianus) 7
Razorbill (Alca torda) 2
Dovekie (Alle alle) 2
Great Black-backed Gull (Larus marinus) 2
Red-necked Grebe (Podiceps grisegena) |
Northern Fulmar (Fud/marus glacialis) l
King Eider (Somateria spectabilis) ]
White-winged Scoter (Melanitta fusca) 1
Purple Sandpiper (Calidris maritima) ]
Atlantic Puffin (Fratercula arctica) ]
Ring-billed Gull (Larus delawarensis) 1
Glaucous Gull (Larus hyperboreus) 1
Total 574

*Includes Common (Gavia immer) and Red-throated Loon
(G. stellata).

on Mendall’s (1986) subspecific key, three of the 12
female Common Eiders measured were classified as
borealis, eight as intergrades, and one as dresseri. Of
the 14 males, six were classified as borealis, four as
intergrades, and four as dresseri. All five of the birds
classified as dresseri had a total culmen length within
1 mm of the range for the intergrade category, and we
suspect that these 5 birds were borealis-dresseri inter-
grades. As in the March 2005 incident, the mass of
oiled eiders was low: mean for females 1359 g (SD
136 g, range 1145-1479 g, n = 5); mean for males
1585 g (SD 153 g, range 1430-1864 g, n = 6). The
mass of oiled male Long-tailed Ducks was 527 g (SD
93 g, range 434-608 g, n = 4) and the one female
weighed 675 g.

Of the murres that could be identified to species,
41.8% (33 of 79) were Common Murres (Uria aalge
[Pontoppidan, 1763]) and 58.2% were Thick-billed
Murres (U. /omvia [Linnaeus, 1758]). The age ratios
were different for the two murre species: 51.7% of
Common Murres were older than one year, while
84.1% of Thick-billed Murres were older than one
year. The mean body mass of Thick-billed Murres
was 711 g(SD 81 g, range 596-822 g, n= 8) and that
of Common Murres was 804 g (SD 37 g, range 774-
844 9g, n = 4). These masses are 26.0% and 21.0%
lighter, respectively, than wintering murres collected
in Newfoundland in the 2000s (Thick-billed Murres
958 g, n = 113, McFarlane Tranquilla er al. [2010];
Common Murres: 1018 g, 7 = 63). A sufficient sample

ROBERTSON ET AL.: OILED EIDERS IN NEWFOUNDLAND

239

was available for measurement of adult Thick-billed
Murres: wing length 215 mm (SD 6 mm, range 206—
231 mm, n= 24); culmen 33.9 mm (SD 2.2 mm, range
30.4-37.7, n= 18).

Discussion

The Avalon Peninsula harbours important concen-
trations of wintering Common Eiders, especially later
in the winter, as birds move south in advance of sea ice
formed to the north (Goudie ef al. 2000) and in early
spring when they begin their northern migration. This
was certainly the case in March 2005, when we count-
ed 42 000 eiders along the eastern stretch of the penin-
sula. Sea ice was notably heavy on the northeastern
coast of Newfoundland in February 2005, forcing
eiders into small open water leads where they were
highly vulnerable to hunting or south into more open
water. Therefore, large numbers of eiders were ex-
pected around southeastern Newfoundland during this
winter.

Based on behaviour that was also noted by Brown
et al. (1973) by eiders involved in the /rving Whale
incident, we assumed that at least 1165 eiders were
oiled. Eiders in unoiled areas flushed well in advance
of the aircraft, but birds suspected to be oiled only
flushed on close approach or did not flush at all, as was
the case among heavily oiled birds hiding in vegetation
adjacent to the shoreline (Brown et al. 1973). There-
fore, the behaviour of oiled eiders appears to be con-
sistent: they remain close to or on shore and do not
respond to an approaching aircraft unless it is very
close. On this basis, we are reasonably confident in as-
suming that all eiders found loafing on islands were
oiled, and this likely represents a minimum number.
Based on our count of 1165 oiled birds on 1 March
2005, plus oiled eiders shot and reported by hunters, at
least 1400 eiders were assumed to be oiled, and this
number could easily be larger. Fewer (337) oiled Com-
mon Eiders were recorded after the April 2006 incident,
but again this represents a minimum.

We detected several large aggregations of loafing
eiders on the first flight in 2005 (1 March): 55% of all
the eiders encountered during this survey were located
at five sites with 75-300 birds each, while only 15%
occurred in groups of less than 10 birds. We believed
that the greatest source of error was our inability to esti-
mate accurately the number of eiders affected from the
air, and we followed this survey with a second flight
on 6 March 2005 combining independent aerial and
ground counts to correct for observer biases in flock
estimation (see Bordage ef al. 1998). However, by the
second survey, all but two of the large aggregations had
dissipated (the two largest counts of remaining birds
were 65 and 85) and the challenge was detecting indi-
viduals or small groups of eiders that were distributed
across more than 200 km of coastline. In the future, we
recommend that aerial assessments be conducted as
soon after an incident as possible to count affected birds

240

when they occur in large aggregations that are easily
detected. Ground counts or photographs should be
used to correct observer biases in flock estimation. If
the birds are widely distributed in small groups, we
suggest using double-observer approaches (see Nichols
et al. 2000) to allow more precise estimates of the
number of birds affected.

The reduction in the number of oiled eiders record-
ed between the | March and 6 March 2005 surveys is
noteworthy and may have resulted from a number of
causes. The first possibility is that birds perished in the
interim, although large numbers of carcasses were not
noted at these sites during the 6 March survey or dur-
ing beach surveys. Eiders appeared to show more “nor-
mal” behaviour at the time of the 6 March survey; thus,
more lightly oiled birds may have been in the water
foraging or loafing and would not have been assessed
as oiled. If this is the case, assessments of oiled eiders
should take place as soon as possible after an incident
and before lightly oiled birds adjust and establish a
somewhat normal behavioural routine. Clearly more
work is needed to understand the behaviour of oiled
birds in the wild so that surveys that best reflect the
numbers of birds oiled during an event can be imple-
mented.

Among oiled birds, adults predominated in both
incidents, especially the April 2006 event. This was
also noted by Brown et al. (1973) in the Irving Whale
spill, which occurred in March 1970 in the same gener-
al area as the April 2006 spill. Sex ratios were close
to one, however, which differs from the male-biased
ratio noted by Brown et al. (1973). Most Common
Eiders found in both incidents were either members of
the northern, or borealis, subspecies, or presumed inter-
grades between the northern and American, dresseri,
subspecies. In fact, most individuals were classified as
borealis-dresseri intergrades, which are found breeding
in central Labrador (Mendall 1986). Pure borealis
Common Eiders that breed in the eastern Canadian
Arctic winter mainly in Greenland, although about a
quarter of the population winters off Newfoundland
and in the Gulf of St. Lawrence (Mosbech et al. 2006).
The Common Eider population breeding in Labrador
has been growing (Chaulk et a/. 2005) and is assumed
to overwinter exclusively in eastern Canada. In iden-
tifying subspecies, we found some discrepancies be-
tween visual inspection of eiders and the use of bill
morphology to key eiders to subspecies (Mendall 1986).
When developing the key, Mendall did not have a
large selection of intergrades, especially males, to in-
clude in the analysis. Some more work on determin-
ing subspecies of Common Eiders, especially in hybrid
zones such as Labrador, is still needed.

Common Eiders are popular game birds in New-
foundland and Labrador and Greenland, and concerns
have been raised about the harvest levels, which, ac-
cording to Gilliland eg a/. (2009), ranged from 68 000
to 83 000 northern eiders annually in the two countries.

THE CANADIAN FIELD-NATURALIST

Vol. 128

This assessment of harvest levels suggested that if
6500 female borealis are taken annually in Canada, the
harvest should be sustainable. The 1400 eiders assumed
to be oiled in the March 2005 event, of which half were
female, would add at least another 11% to the overall
winter take, possibly pushing the population into de-
cline for that season (assuming oiling was in addition
to natural and hunting mortality). The fact that most
oiled eiders were adults — unlike hunted birds, which
are largely juveniles (Gilliland and Robertson 2009)
— only exacerbates the pressure this event placed on
the population.

Recovered carcasses of Common Eiders were some-
what lighter than normal wintering birds, but they were
not as emaciated as murres, and living oiled eiders were
observed for over a month after the initial oiling event.
Therefore, unlike murres and other auks, Common
Eiders appear to be able to sustain some level of oiling
and survive. Eiders are inshore feeders, foraging in
shallow intertidal and subtidal waters. When oiled,
eiders, like the auks, come to shore in large numbers,
presumably to avoid frigid waters and the risk of hy-
pothermia resulting from oiled plumage. However, food
for eiders is readily available close to shore, and eiders
can continue to dive, feed, and return quickly to shore.
In contrast, auks that come to shore are far removed
from their food source and eventually succumb to star-
vation. Even if oiled birds are unable to forage, Com-
mon Eiders, at least females, are adapted to withstand
severe loss of body mass (up to 40%) and condition
while they incubate their clutch (Parker and Holm
1990). What is not known is the long-term viability
of the eiders that were lightly oiled. Even if they sur-
vived, future reproductive attempts by these birds may
be compromised, as minute levels of hydrocarbons
have been shown to severely disrupt or inhibit repro-
duction (Leighton 1993; Stubblefield et a/. 1995), and
lingering long-term physiological impairment has been
shown in birds that were exposed to sub-lethal doses
of hydrocarbons (Alonso-Alvarez et al. 2007a,b).

The species composition of murres found in the
April 2006 incident was similar in some respecis, but
also differed from those seen in the November 2004
incident (Robertson et a/. 2006). Common Murres
made up a larger proportion of the murres recovered
in April 2006, reflecting their prevalence along the
south coast of Newfoundland, especially later in the
winter season. In spite of the proximity of Common
Murre breeding colonies to the affected area (both C ape
St. Mary’s and Witless Bay Island Ecological Reserves
are less than 100 km away), the majority of recovered
Common Murres were juveniles. On the other hand,
most Thick-billed Murre carcasses were adults. in a
proportion comparable to what would be expected in
the population at large (Wiese er al. 2004). Body mass-
es of both species were lower than expected for winter-
ing birds. However, the mass of Thick-billed Murres
was not quite as low as that of starved birds found in

2014

the “seabird wrecks” of 2007 and 2009 when beached
dead birds were found to weigh 622 g and 639 g on
average, respectively (McFarlane Tranquilla ef al.
2010), indicating that oiled birds had some remain-
ing body reserves and hypothermia, toxicity of oil, or
both, contributes to the death of oiled murres. Wing
and culmen measurements were typical of birds breed-
ing in the Northwest Atlantic and similar to those in-
volved in other oiling events (Brown ef al. 1973; Rob-
ertson ef al. 2006). Of note was the lack of Dovekies,
which are a common victim of oiling in Newfoundland
waters (Wiese and Ryan 2003; Robertson et al. 2006).
Possibly Dovekies had begun to move north on migra-
tion by the time of the incident in April 2006.

The April 2006 event was noteworthy in terms of the
wide variety of avifauna affected: no less than 19 spe-
cies, including sea ducks, alcids, loons, gulls, and even
a sandpiper and a tubenose. This impact is related to
both the timing and location of the event. Unlike many
parts of southeastern Newfoundland, the southern tip
of the Avalon Peninsula has extensive beach habitat in
addition to the rockier cliff habitat surrounding most
of the peninsula. This habitat diversity leads to a broad-
er avian community. In addition, in mid-April, many
species would be congregating for the spring migra-
tion, and others would have begun migration. The spill
was early enough that the main wintering aggregations
of both eiders and murres had not yet left the area, as
most of the eiders were of the northern types and Thick-
billed Murres dominated among the recovered murre
carcasses (both of these species breed north of New-
foundland). Both inshore (ducks, loons) and offshore
species (auks, fulmars) were affected, suggesting that
this slick likely started somewhere out from shore and
was blown inshore.

Between the April 2006 oiling incident and April
2014, there have been no notable (1.e., affecting >10
birds) inshore oiled bird events in Newfoundland from
unknown sources, although significant numbers of
oiled birds have been found associated with the M/V
Manolis L wreck on the northeast coast of Newfound-
land. Regular long-term beached bird surveys in south-
eastern Newfoundland are also beginning to show a
reduction in the number of oiled birds coming to shore
(Wilhelm ef a/. 2009). In 2005, federal legislation was
passed in Canada directed at reducing the chronic ship-
source oil pollution problem (Elmslie 2006); concomi-
tantly, additional surveillance and enforcement and
education programs were put in place to target this is-
sue. Although continued monitoring 1s warranted, the
indications are that a reduction in chronic ship-source
oiling, at least in inshore waters off southeastern New-

foundland, is occurring.

Acknowledgements

We thank the many agencies and individuals who
assisted in the response to these oiling events. In addi-
tion to collating reports and collecting carcasses, the

ROBERTSON ET AL.: OILED EIDERS IN NEWFOUNDLAND

241

Canadian Coast Guard provided helicopter support
for the various surveys. Graham Thomas, the Govern-
ment of Canada’s Environmental Emergencies Pro-
grams, and members of the Atlantic Regional Envi-
ronmental Emergencies Team provided key oversight
of these responses. Sabina Wilhelm, Francois Bolduc,
and an anonymous reviewer provided helpful com-
ments on an earlier draft of this manuscript.

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THE CANADIAN FIELD-NATURALIST

Vol. 128

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Received 23 January 2014
Accepted 10 March 2014

Impact of the 2012 Drought on Woody Vegetation Invading Alvar
Grasslands in the Burnt Lands Alvar, Eastern Ontario

PAUL M. CATLING

Environmental Health, Biodiversity, Saunders Building, Central Experimental Farm, Agriculture and Agri-Food Canada, Ottawa,
Ontario K1A 0C6, Canada; email: Paul.Catling@agr.ge.ca

Catling, Paul M. 2014. Impact of the 2012 drought on woody vegetation invading alvar grasslands in the Burnt Lands Alvar,
eastern Ontario. Canadian Field-Naturalist 128(3); 243-249,

Counts of dead and living trees in open alvar indicated that the 2012 drought had a significant impact on the Burnt Lands
Alvar in eastern Ontario, with mortality of woody species ranging from 10% to 100% at open alvar sites. The maximum age of
dead trees suggested that the most recent drought of similar impact occurred more than 35 years ago, possibly in 1974. Some of
the killed junipers were 87-90 years old. All dry periods indicated on a summer rainfall diagram may not be severe enough
to kill woody vegetation, but a drought causing an average 50% kill of woody vegetation in open alvar may have occurred at
least once every 30 years over the past century. Consequently the encroachment of woody species into alvar grassland appears
not to be a threat to this habitat. Open alvar could have existed on the landscape as isolated occurrences since early postglacial
times, if the climate fluctuated in the distant past as it has recently. With evidence for drought in a fluctuating climate to maintain
the disjunct occurrences of midwestern plant and animal species, these occurrences may be considered as reliable indicators
of a more continuous distribution of such species more than 10 000 years ago.

Key Words: alvar; Great Lakes region; eastern Ontario; drought; woody vegetation; succession; Common Juniper; Juniperus

communis; White Spruce; Picea glauca; Eastern White Pine; Pinus strobus; Eastern White Cedar; Thuja occidentalis

Introduction

Alvar grasslands are naturally open areas of thin soil
over flat limestone or marble rock with a more-or-less
sparse grass—dominated vegetation with trees absent
(Catling and Brownell 1995). Three kinds of predom-
inant alvar grassland are considered to be globally im-
periled, meaning that they are “at high risk of extinction
or elimination due to restricted range, few populations
or occurrences, steep declines, severe threats, or other
factors” (Natureserve 2014). Alvar grasslands also con-
tain a diversity of plants and animals, some of which
have been evaluated as at risk or susceptible to extir-
pation (Catling et al. 2014), reflecting the fact that these
habitats are subject to a variety of threats. Alvar habitats
are important with regard to climate change studies, and
the grasslands, in particular, are believed to be the only
existing analogues of the Picea parkland that existed in
front of the Wisconsin glacier for hundreds of thousands
of years (Webb 1987). Alvars also provide in situ pro-
tection for native crop relatives and major recreational
opportunities (Catling et al. 2014).

The survival of alvar grassland in the northern Great
Lakes region since the early postglacial period (e.g.,
Catling and Brownell 1995; Hamilton and Whitcomb
2010) was likely the result of resistance to succession
to woodland as a consequence of fire removing trees
and shrubs, periodic drought, and a seasonal cycle of
drought and flooding (Catling and Brownell 1995; Jones
and Reschke 2005). The effects of fire at a number of
sites in the Great Lakes region have been reported to
range from the creation and maintenance of grasslands
to no effect (e.g., Catling and Brownell 1998; Jones and
Reschke 2005; Catling 2009), and it has been conclud-

ed that, in some situations, fire is “not the primary fac-
tor in maintaining open condition” (Jones and Reschke
2005). The seasonal cycle of spring flooding and late
summer drought can be seen on an annual basis, but
only Stephenson and Herendeen (1986), working at
Maxton Plains in northern Michigan, have documented
the impact of periodic drought on alvar grassland over
a longer term. An opportunity to add to their observa-
tions arose with the drought of 2012 when woody veg-
etation declined on the northern part of the Burnt Lands
Alvar in the Ottawa Valley of eastern Ontario. At the
end of the summer of 2012, leaves on many of the pre-
dominant conifers had turned brown, and much of the
brown leaf material and dead branches were evident on
the trees and Juniper shrubs (personal observation).

In this article, | explore the idea that drought had a
significant effect in restricting woody growth on the
Burnt Lands Alvar through analysis of data on mortal-
ity of woody plants. In general, this work is designed to
improve understanding of the derivation and future of
alvar habitats by elucidating the factors that control spe-
cies composition and, specifically, those that relate to
the threat of forest encroachment.

Study Area

The Burnt Lands Alvar, 4 km northeast of Almonte,
Mississippi Mills, Ontario, at approximately 45.2698,
~76.1942 is a 15—km” area of provincially significant
flora and fauna (Brunton 1986*), much of which is
within Burnt Lands Provincial Park. The grasslands here
are dominated by perennial Prairie Dropseed (Sporobo-
lus heterolepis [|A. Gray] A. Gray) and annual Sheathed
Dropseed (Sporobolus vaginiflorus |Torrey ex. A. Gray]

243

244

THE CANADIAN FIELD-NATURALIS1

Vol. 128

FIGURE 1. Dead Common Juniper (Juniperus communis) in alvar grassland at the north end of the Burnt Lands Alvar, East-
ern Ontario. The shrubs were killed by the extreme drought in 2012. Photo: P. M. Catling, September 2013.

Alph. Wood var. vaginiflorus) with numerous other
prominent species characteristic of alvars also present,
including Crawe’s Sedge (Carex crawei Dewey), Small
Skullcap (Scutellaria parvula Michaux), False Pen-
nyroyal (Trichostema brachiatum L.), and Balsam
Groundsel (Packera paupercula [Michaux A. Léve
& D. Love). Other species locally common are Com-
mon Self-heal (Prunella vulgaris L.), Oxeye Daisy
(Leucanthemum vulgare Lamarck), Tall Hawkweed
(Hieracium piloselloides Vill.) and Common Viper’s
Bugloss (Echium vulgare L.). The grasslands are sur-
rounded by forest dominated by Eastern White Cedar
(Thuja occidentalis L.) with Common Juniper (Junipe-
rus communis L.), White Pine (Pinus strobus L.), and
White Spruce (Picea glauca [Moench] Voss). These
woody species are also present as individuals within
the grassland. The earlier established woody species
provide shade that reduces water loss for later colo-
nizers resulting in patches of woody plants. Through
this process of nucleation, woodland begins to replace
open alvar.

Methods

To test the hypothesis that the 2012 drought had a
significant impact on woody species, the following data
were gathered in late September 2013, one year after
the drought and at a time when the drought impact was
conspicuous with persisting dead branches and branch-
es retaining brown leaves (Figures | and 2).

Age of woody vegetation killed

The age of some of the larger woody plants killed
was determined by counting growth rings of cut dead
trees with a stereo microscope. Resolution of growth
rings was improved through the application of linseed
oil to the cut surfaces. Seven Eastern White Cedars and
three White Spruces were cut at breast height, and dia-
meter at breast height (dbh) was measured. Five years
was added to the growth ring estimates to account for
the time required to reach breast height and make esti-
mates of age more accurate. The ages of three White
Pines were estimated by counting spaces between
branches on the trunk. Three Common Junipers were
cut at the stem base and growth rings were counted.

2014

FIGURE

CATLING: DROUGHT IMPACT ON Woop\ VEGETATION INVADING ALVAR GRASSLANDS 245

7

goes)

2. Dead Eastern White Cedar (Thuja occidentalis, front right) and dead White Spruce (Picea glauc eee ;
nucleation patch with some living cedar (left) and half dead Common Juniper (Juniperus mill ph aes : :
The dead cedar and spruce were approximately 30 years old as indicated by a count of prowih sais . ah ce
were killed in 2012, when 60% of the trees in nucleation patches in the Burnt Lands Alvar, Eastern Ontario, diec

apparently due to extreme drought. Photo: P. M. Catling, September 2013.

246

Decline of woody vegetation along a limestone pave-
ment crack

Based on the amount of living material compared
with dead leaves and leafless branches, trees and shrubs
were classified as dead, half dead, or alive and mapped
along two, 25—m long cracks in the flat bedrock (lime-
stone pavement) to provide a visual indication of the
impact of the drought in an area of heavy damage. The
area examined was rectangular (20 m x 25 m). Dbh for
trees and maximum diameter of branch spread (i.e., of
the patch) for juniper shrubs were recorded and indi-
cated on the diagram (Figure 3). The dbh and shrub
diameters were also used to calculate percentage de-
cline (total dbh or patch size of dead material divided
by total dbh or patch size times 100).

Juniper decline in open grassland

The maximum diameter of more or less circular
patches of juniper was recorded in an open rectangu-
lar area 50 m =< 100 m (5000 m7?) and the extent to
which patches had died, in terms of percentage of lost
surface area, was estimated to within 5%.

General decline of woody vegetation in open grassland

At three sites separated by 0.5—1 km, the status of
all trees and shrubs in alvar grassland was recorded as
dead, half dead, or alive (see above), and dbh for trees
and maximum diameter for juniper shrubs was record-
ed. For each of the tree species and for each status cate-
gory, the number of individuals, the dbh range and total
of all dbh measurements were noted (Table 1). The
sizes of areas surveyed are indicated in Table 1. From
the data for dead tissue (total dbh for dead trees + 4 dbh
for half dead trees), a percentage decline for all trees
was calculated by dividing by the total dbh and multi-
plying by 100. A percentage decline for juniper shrubs
was calculated by dividing loss of surface area by total
surface area of patches times 100.

Documentation of the drought of 2012

Total rainfall for June, July, and August was used
to identify drought years from 1890 to present. Rain-
fall is only an approximation of drought because the
evenness of the pattern of rainfall for the period is not
considered, nor is localized convectional rain, which
may differ between a recording station and an area under
consideration. The amount of heat, which increases
water loss by evapotranspiration, is also an important
factor that is not considered here. Although it is not
an entirely reliable indication of drought, low total rain-
fall is highly correlated with severe drought impact and
is, thus, helpful in determining drought occurrences if
these limitations are taken into account.

Historical Environment Canada (2014) weather data
for the closest stations were used. Because the record
for the nearest town, Almonte, is incomplete, the record
for Ottawa (45.38333, —75.71666), 40 km east by
southeast of the study site, was used for the period
1890-2006. For 2007-2013, the potentially more accu-
rate data from the nearby village of Appleton (45.18683,

THE CANADIAN FIELD-NATURALIST

128

Vol.

FIGURE 3. Diagram showing status of woody vegetation along
two pavement cracks (stippled) at the north end of
the Burnt Lands Alvar, Eastern Ontario. The diagram
is based on a field sketch of a rectangular area 20 x
25 m and is drawn approximately to scale. Circles
are trees and shrubs, where C = Eastern White Cedar
(Thuja occidentalis), P = White Pine (Pinus strobus),
J = Common Juniper (Juniperus communis) and
white = dead, grey = half-dead, and black = alive.
The numbers for cedar and pine are diameter of the
trunk at breast height in cm; for junipers they are
diameter of the whole shrub (m).

~76.10768), 12 km south by southeast of the study
site, were used.

Results
Age of woody vegetation killed

The interpretation of growth rings was difficult in
some areas and resulted in a range of values. Harvest-
ed Eastern White Cedar was 34—40 years old with dbh
ranging from 15 to 17 cm. White Spruce was of similar
age and diameter, while White Pine ranged from 35 to
50 years old. Most dead Common Juniper shrubs were
older than the trees: 33-90 years old and 3-13 cm dbh.

P. : : Prd
014 = CATLING: DROUGHT IMPACT ON Woopy VEGETATION INVADING ALVAR GRASSLANDS = 247

TABLE aNclIFAMmant< At dac Bye eee . os , : Ce
ae =a ain of dead, half-dead and living Eastern White Cedar ( Thuja occidentalis), White Spruce (Picea glauca),
ite Pine (Pinus strobus), and Common Juniper (Juniperus communis) at three open alvar grassland sites on the Burnt

Lands Alvar, Eastern Ontario.

S..—OOOOOo—S—e

Site 1 (45 x 80 m)

Half-
Species and measures Dead dead Alive
Eastern White Cedar
Number 16 2 10
Dbh range (cm) 1-20 + 1-9.5
Total dbh (cm) 96.7 8 Bos
White Spruce
Number 2 = 4
Dbh range (cm) 5-6 3-12
Total dbh (cm) 11 - 22,
White Pine
Number 2 - —
Dbh range (cm) 2.32.4 ~ -
Total dbh (cm) 4.7 - -
Common Juniper
Number 8 l 5
Patch diameter (m) 0.82.5 06 0.3-1.8
Total of all patches (m) 10.9 0.6 3.9
All trees
Total dbh (cm) L75e7
% decline 66.3
Common Juniper
Total of all patches (m) 15.4
% decline E,

Site 2 (25 x 55 m) Site 3 (28 x 42 m)
Half Half
Dead dead Alive Dead dead _ Alive
19 2 8 12 | 19
2-16 5-14 3-14 1-18 7 15-16
176 19 67 84 7 111
— 6 — =
= = 2-10 _
ne 20 = é
_ — 2 3 | —
= — 20-28 12-15 qf -
= = 48 39 7 -
| 3 3 | 3 ]
2 3-5 2-4 | 1-3 4
2 12 9.5 ] 5 4
310.0 268.0
59.8 56.0
23.5) 10.0
34.0 35.0

Decline of woody vegetation along a pavement crack

Most of the trees and shrubs in nucleation patches
along a pavement crack were killed (Figure 3). The
larger dead cedars (those about 15 cm dbh) in these
patches were likely over 30 years old based on dbh of
the few trees of definitely known age. The overall de-
cline of trees in these pavement cracks was 80% and
that of junipers was 93%.

Juniper decline in open grassland

Fifty-eight more-or-less circular patches of Com-
mon Juniper, 0.5—1 m in diameter occurred in the sel-
ected area. The majority of these shrubs were 35—45
years old based on basal stem diameters of a few
trees of known age, and larger patches were composed
of more than one plant. In 2012, 20 of the plants had
died and many others had lost much of their foliage
(Figure 1). The total area of living Common Juniper
observed here at the start of 2012 had been approxi-
mately 274 m? and this had declined to 88 m? by the
end of 2012, representing a 68% reduction. This area
included shrubs up to 90 years old.

General decline of woody vegetation in open grassland

The overall decline of trees at three sites ranged from
56% to 66% and decline of junipers ranged from 35%
to 73%. Thus over half of the woody vegetation in the
openings was killed (Figure 2). Some of the trees at all

three sites were over 35 years old, but all age classes
and all three tree species were affected by the drought
(Table 1).

Documentation of the drought of 2012

Since 1890, the average annual summer precipita-
tion in the Ottawa area has been 255 mm. Based on
this average and the pattern, a plot for this period
(Figure 4) shows that unusually dry years (less than
160 mm of rain in June-August), approximately equiv-
alent to the 2012 drought, have occurred on average

500 fF" ~

1880 1900 1920 1940 1960 1980 2000 2020
Year

Total Rainfall (June - Aug., mm)

Figure 4. Summer rainfall (mm) from 1890 to 2013 near the
Burnt Lands Alvar, Eastern Ontario, based on records
of Environment Canada (2014).

248

every 8 years. The period from 1890 to 1906 was evi-
dently the longest drought—free period on record. The
2012 drought was the second—most extreme in terms
of limited rainfall (134 mm), the most extreme being
the drought of 1931 (112 mm).

Unusually wet years (over 380 mm of rain in June—
August) have occurred less frequently (Figure 4) than
unusually dry years, with an average of 20 years be-
tween. Before 2013 (432 mm), the last unusually wet
year occurred in 1972 (468 mm) and was the wettest
on record.

Discussion

The 2012 drought had an impact on the Burnt Lands
Alvar that was very similar to that reported for Max-
ton Plains (Stephenson and Herendeen 1986), where
woody species mortality ranged from 10% to 100% at
open alvar sites. The wide variation at Maxton Plains
and Burnt Lands Alvar may have been based on differ-
ences in surface water flow, making some sites more
susceptible to woody plant mortality than others.

The variation in the age of dead trees on Burnt Lands
Alvar may be a consequence of random differences in
times of establishment. Assuming that the same dry
sites are susceptible to consecutive droughts, and based
on the age range of 34—S0 years of the larger dead trees,
the last droughts as severe as that of 2012 occurred
52 years ago in 1960 and 38 years ago in 1974. How-
ever, some of the killed junipers were 87—90 years old
and had survived droughts for almost 100 years. Al-
though the wet years have allowed woody invasions,
the higher frequency of dry years, some severe, has
prevented encroachment of woody plants into grass-
lands and other open areas.

The drought of 2007 did not have as severe an im-
pact on woody vegetation as that of 2012 (personal
observation) despite similarly low overall rainfall (Fig-
ure 4). Possibly different rainfall patterns, less local
rain, and higher heat contributed to the much greater
severity in 2012. A drop to a critical level of dryness
may also have played a role in 2012. Although not all
dry periods indicated on the total rainfall diagram were
severe enough to kill woody vegetation, those poten-
tially capable of doing so have occurred with a high
enough frequency that a drought averaging a 50% kill
of woody vegetation in open alvar may have occurred
once every 30 years over the past century. As this may
be sufficient to maintain open alvar in many areas, it
supports the conclusion of Stephenson and Herendeen
(1986) that woody species encroachment into alvar
grassland is not a threat to open alvar habitat. Studies
of drought effects on some Scandinavian alvars, such
as Stora Alvaret on the Swedish Island of Oland, have
also documented a decline of juniper as a result of
drought (Rosen 1984), although these alvars are often
thought to have been largely created and maintained by
livestock grazing (Znamenskiy et al. 2006).

THE CANADIAN FIELD-NATURALIST

Vol. 128

By preventing encroachment of woody plants, peri-
odic drought may provide a mechanism for the long-
term persistence of species in alvars that are consid-
ered remnants of more continuous distributions in early
postglacial landscapes. These occurrences of early post-
glacial and midwestern species assemblages, including,
for example, the occurrence of the flightless leafhop-
per associated with the community—dominant Prairie
Dropseed grass (Hamilton and Whitcomb 2010), now
have a better supported theory to explain their isolation.

Although fire may have more impact on the edges of
open alvar and in the creation of open alvar from alvar
woodland (Catling 2009), or may vary in impact from
one alvar to another (Jones and Reschke 2005), drought
may prevent succession to forest on parts of open alvar
and on alvar pavements not subject to fire. Drought
may also promote fire by creating drier conditions and
by increasing fuel. Consequently these processes may
have a complementary effect in the maintenance of
open conditions on alvars, but drought alone can have
substantial impact.

Documents Cited (marked * in text)

Brunton, D. F. 1986. A life science inventory of the Burnt
Lands. Ontario Ministry of Natural Resources, Carleton
Place, Ontario, Canada. 118 pages + 3 maps.

Literature Cited

Catling, P. M. 2009. Vascular plant diversity in burned and
unburned alvar woodland: more evidence of the impor-
tance of disturbance to biodiversity and conservation.
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Catling, P. M., and V. R. Brownell. 1995. A review of the
alvars of the Great Lakes Region: Distribution, floristic
composition, biogeography and protection. Canadian
Field-Naturalist 109: 143-171.

Catling, P. M., and V. R. Brownell. 1998. Importance of
fire in alvar ecosystems — evidence from the Burnt Lands,
eastern Ontario. Canadian Field-Naturalist 112(4): 661—
667.

Catling, P. K.. P. M. Catling, J. Cayouette, and M. J.
Oldham. 2014. Canadian Alvars and Limestone Barrens:
Areas of “Special Conservation Concern” for plants?
Canadian Botanical Association Bulletin. Jn press.

Environment Canada. 2014. Accessing the data. Environ-
ment Canada, Fredericton, New Brunswick, Canada.
Accessed 20 Nov, 2013. http://climate.weather.ge.ca/.

Hamilton, K. G. A., and R. F. Whitcomb. 2010. Leafhop-
pers (Homoptera: Cicadellidae): a major family adapted to
grassland habitats. Pages 169-197 in Arthropods of Cana-
dian Grasslands. Volume 1: Ecology and Interactions in
Grassland Habitats. Edited by J. D. Shorthouse and K. D.
Floate, Biological Survey of Canada.

Jones, J., and C. Reschke. 2005. The role of fire in Great
Lakes Alvar Landscapes. Michigan Botanist 44: 13-27.
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Status. NatureServe, Arlington, Virginia, USA. Accessed
February 2014. http://explorer.natureserve.org/ranking.htm.

Rosen, E. 1984. Some short-term changes in the dynamics
of limestone grassland of South Oland, Sweden. Acta Uni-

2014

versitatis Upsaliensis Nova Acta Regiae Societatis Scien-
tiarum Upsaliensis Series V C. 3: 189-205.

Stephenson, S. N., and P. S. Herendeen. 1986. Short-term
drought effects on the alvar communities of Drummond
Island, Michigan. Michigan Botanist 25: 16-27.

Webb, T. 1987. The appearance and disappearance of major
vegetational assemblages: long term vegetational dynamics
in eastern North America. Vegetatio 69: 177-187.

CATLING: DROUGHT IMPACT ON Wooby VEGETATION INVADING ALVAR GRASSLANDS

249

Znamenskiy, S., Helm, A. and M. Partel. 2006. Threatened
alvar grasslands in NW Russia and their relationships to
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Received 6 January 2014
Accepted 14 February 2014

A Review of Colour Phenotypes of the Eastern Red-backed
Salamander, Plethodon cinereus, in North America

JEAN-DaAvip Moore!? and MARTIN QUELLET2

'Forét Québec, Ministére des Foréts, de la Faune et des Parcs, Direction de la recherche forestiére, 2700 rue Einstein, Québec,
Québec GIP 3W8 Canada

?Amphibia-Nature, 469 route d’Irlande, Percé, Québec GOC 2L0 Canada

3Corresponding author: e-mail: jean-david.moore@mffp.gouv.qe.ca

Moore, Jean-David, and Martin Ouellet. 2014. A review of colour phenotypes of the Eastern Red-backed Salamander, Plethodon
cinereus, in North America. Canadian Field-Naturalist 128(3): 250-259.

The Eastern Red-backed Salamander (Plethodon cinereus) is the most abundant salamander species in many forests of north-
eastern North America. It is well-known for its colour polymorphism, which includes eight colour phenotypes: the red-backed
(striped), lead-backed (unstriped) and erythristic morphs, as well as the iridistic, albino, leucistic, amelanistic and melanistic
anomalies. Here we review the various colorations of P. cinereus, with the objective of facilitating the identification of these
different phenotypes and of generating interest among field herpetologists and scientists reporting on this species. We also
list six previously unpublished occurrences of colour variants in this species (1 case of erythrism, 3 of iridism, | of leucism, and
| of partial leucism). To our knowledge, these cases include the first documented occurrence of iridism in the red-backed morph
of P. cinereus, and the first two mentions of this colour anomaly in the lead-backed morph from Canada.

Key Words: Phenotypes; coloration; red-backed; lead-backed: erythristic; colour morph; iridistic; albino; leucistic; amelanis-

tic; melanistic; colour anomaly; Eastern Red-backed Salamander; Plethodon cinereus: North America

Introduction

Of all North American amphibians, the Eastern Red-
backed Salamander (Plethodon cinereus) is probably
one of the most studied species, because of its wide dis-
tribution and abundance in forests of eastern Canada
and northeastern United States (Highton 1962; Burton
and Likens 1975; Petranka 1998), its ecological role
(Burton and Likens 1975; Wyman 1998), and its poten-
tial as indicator of climatic change (Lotter and Scott
1977; Gibbs and Karraker 2006) and habitat distur-
bances (deMaynadier and Hunter 1995) in forest eco-
systems. Striking colour polymorphism of this salaman-
der species has long attracted the attention of scientists,
herpetologists, and naturalists. Three pigment cell class-
es, or chromatophores, are responsible for skin colora-
tion in amphibians (Bagnara 1966; Taylor and Bagnara
1972): melanophores (brown to black pigment cells),
xanthophores (yellow and red (erythrophores) pigment
cells), and iridophores, which produce the shiny irides-
cent and reflecting aspect of the amphibian skin.

In this article, we review the state of knowledge for
each of the eight colour phenotypes known to date in P
cinereus, which include the red-backed (striped), lead-
backed (unstriped) and erythristic morphs, as well as the
iridistic, albino, leucistic, amelanistic and melanistic
anomalies. Our objective is to provide a standardized
description accompanied by photographs for the colour
phenotypes reported for P. cinereus. Characteristics use-
ful to discriminate among these different phenotypes
are summarized. The North American distribution of the
different colour morphs and anomalies in P. cinereus is
also provided. Finally, we report six previously unpub-
lished phenotype occurrences in this species (1 of ery-

thrism, 3 of iridism, | of leucism, and | of partial leu-
cism).

Colour Morphs
The red-backed and lead-backed colour morphs

The red-backed and lead-backed morphs (Figure 1)
are the two most abundant phenotypes in P. cinereus.
The ventral body of both morphs is coarsely mottled
with black and white, resulting in a distinctive “salt-
and-pepper” appearance (Figure 2; Table 1; Appendix).
Individuals with intermediate colorations or other vari-
ations between these two morphs have been observed
(e.g., Figure 3; Sipes 1964). The mid-dorsal stripe of
the red-backed morph is generally red-orange, with
gray-black borders that become mottled with white
along the lower half of the body sides. Mid-dorsal stripe
colours other than red-orange, such as shades of brown,
gray, pink, white, and yellow have also been observed
(e.g., Figures 4, 5; Bishop 1941; Test and Bingham
1948; Reed 1955; Schueler 1975: Hulse et al. 2001).
Colour variations also include individuals with stripes
that occur in disconnected sections along the body, and
others with only continuous stripes on the tail (Bish-
op 1941; Cook 1967; Petranka 1998). The dorsum of
the lead-backed morph does not exhibit stripes, and the
coloration is generally a uniformly pigmented gray-
black, but sometimes olive-gray or chestnut-brown (es.
Figure 6; Klemens 1993). The dorsal-lateral body often
exhibits some degree of iridescent or metallic. blue
(Sawyers and Novick 2011), brassy, golden, shades of
green or silver flecks (e.g., Figure 7).

Burger (1935) was the first to publish evidence that
both morphs could be found in the same brood. which

250

2014

MoorE AND QUELLET: COLOUR PHENOTYPES OF PLETHODON CINEREUS j

Nn

\BLE I. Main characteristics of the 8 colour phenotypes of the Eastern Red-backed Salamander (Plethodon cinereus).

Phenotype Dorsal stripe

Dorsal-lateral body

Red-backed Continuous or
discontinuous
red-orange stripe;
sometimes shades
of brown, gray,
pink, white or

Gray-black

Ventral body Iris
“Salt-and-pepper” Normal (dark)
appearance pigmentation

yellow
Lead-backed Unstriped Gray-black; sometimes “Salt-and-pepper” Normal (dark)
olive-gray or chestnut-brown appearance pigmentation
Erythristic Unstriped Red or orange, with or Red or orange Normal (dark)
without black mottling pigmentation
Iridistic Stripe present Generalized iridescent “Salt-and-pepper” Normal (dark)
or absent flecking (brassy, golden, appearance with or pigmentation
shades of green or silver) without iridescent
flecking
Albino Unstriped Pink-white to white Pink-white to white, Red-pink
translucent
Leucistic Unstriped Pink-white to white Pink-white to white, Normal (dark)
translucent pigmentation
Amelanistic Peach-red, orange Pink-white to white Pink-white to white, White to normal
or peach-yellow translucent (dark) pigmentation
Melanistic Unstriped Black Black, translucent Brown to black

showed that they are of the same species (Highton
1959). Both morphs can be found across the species
range, throughout southeastern Canada and the eastern
United States (Petranka 1998), but in very different
proportions (Moore and Ouellet 2014). In some areas,
both morphs are abundant, while in others, one or the
other is more abundant. Monomorphic red-backed pop-
ulations are not uncommon, while in other cases, red-
backed individuals can be scarce or absent (Thurow
1955; Highton 1959; Pfingsten and Walker 1978;
Reichenbach 1981; Fisher-Reid et a/. 2013). The red-
backed morph has been reported in five Canadian
provinces and 22 American states (Table 2), which
represents the distribution range of this species. Ken-
tucky, Minnesota, and Tennessee are at the limit of
the species’ range and the lead-backed morph of P.
cinereus has not been reported there.

In the last decades, studies have suggested that the
lead-backed morph of P. cinereus is more closely
associated with warmer climates than the red-backed
morph (Lotter and Scott 1977; Moreno 1989; Gibbs
and Karraker 2006; Anthony et a/. 2008). However, it
seems that the possible role of climate in the distribu-
tion of the lead-backed morph has never been unani-
mously recognized; some doubts still persist in the
scientific community about the validity of using this
morph as indicator of climatic changes (Angleberger
and Chinnici 1975; Pfingsten and Walker 1978; Pet-
ruzzi et al. 2006; Anthony ef al. 2008). Based on new
discoveries in the northern areas of the species’ range
and on the largest compilation ever made for this spe-
cies. Moore and Ouellet (2014) demonstrate however
that climate and geographic variables do not influence

the colour morph proportions in P. cinereus popula-
tions. Fitzpatrick et al. (2009) mentioned that the
most reasonable hypothesis to explain the existence of
visual polymorphism across a geographical and phylo-
genetic range as wide as that of P. cinereus is that selec-
tion acts directly on appearance. Local natural selection
could explain the morph frequency in given areas.

The erythristic colour morph

Erythrism (from Greek eruthros, red) refers to an
excessive production and deposition of red or orange
pigments (erythrophores). The erythristic morph of P.
cinereus (Figures 8, 9) is usually all red or orange, but
may exhibit varying degrees of black mottling on its
dorsal and lateral surfaces. The ventral surface is gen-
erally red, orange or pink (Reed 1955; Thurow 1961;
Rosen 1971). Some intermediate colorations (partial
erythrism) also exist (e.g., Figure 10). The erythristic
morph has been found in four provinces (Brown 1928;
Bleakney and Cook 1957; Cook and Bleakney 1961)
and 10 states (Reed 1908; Reed and Wright 1909; Bar-
bour 1914; Burt 1945; Matthews 1952; Pfingsten 1969;
Mueller and Himchak 1983; deMaynadier 1995; Mc-
Donald et al. 2011), but it is more frequently found
in United States than in Canada. The prevalence of
this morph in the United States can sometimes be as
high as 35-50% (Pauley er a/. 2001; Cassell and Jones
2005), but generally rarely exceeds 20% (Lotter and
Scott 1977; Tilley et al. 1982). According to Pauley
et al. (2001), the distribution of this morph would be
limited to cooler climates of glaciated areas of southern
Canada and the northeastern United States. In this con-
text, the apparent rarity of the erythristic morph in east-
ern Canada remains enigmatic. Thirty-seven erythristic

ry 9
252 THE CANADIAN FIELD-NATURALIS1 Vol. 128

FIGURE |. Red-backed (striped) and lead-backed (un- FiGURE 2. Distinctive “salt-and-pepper” appearance of
striped) morphs of the Eastern Red-backed Sala- the ventral body of a lead-backed morph of P.
mander (Plethodon cinereus) from Québec. Photo: cinereus from Québec. Photo: M. Ouellet.

M. Ouellet.

mee ee ——

FiGuRE 3. Slight colour variation of a red-backed morph of FIGURE 4. Cream coloration of the dorsal stripe of a

P. cinereus from Québec. Photo: M. Ouellet. red-backed morph of P. cinereus from New

Hampshire. Photo: D. J. Hocking.

FIGURE 5. Shade of gray coloration of the dorsal stripe of FIGURE 6. Chestnut coloration of a lead-backed morph of
a red-backed morph of P. cinereus from Ohio.
Photo: C. D. Anthony.

FIGURE 7. A lead-backed morph of FP. cinereus with blue FiGurE 8. An erythristic morph of P. cinereus from
iridophores from Maine. Photo: D. E. Swann. Québec. Photo: P. Beaupre

20 :
2014 MoorE AND OUELLET: COLOUR PHENOTYPES OF PLETHODON CINEREUS

Mm
Nn
oe)

ae 4 Se PB

, eC or eX: >of er - : : ; f

FiGuRE 9. Another example of erythrism of P. cinereus, FiGure 10. A partially erythristic morph of P. cinereus
with black mottling on the tail, from Nova Scotia. from New Hampshire. Photo: D. J. Hocking
Photo: R. Merrick. }

~ ~
. 2 Ss Ae
FiGure 11. An iridistic colour anomaly of a red-backed FIGURE 12. Iridism of a lead-backed morph of P. cinereus
morph of P. cinereus from North Carolina. from Nova Scotia. Photo: J. Gilhen

Photo: L. A. Williams.

4 ® 7 a Z ay oe — : :
FIGURE 13. An albino colour ae Y P. cinereus FicurE 14. A leucistic colour anomaly of P. cinereus
from Nova Scotia. Photo: J. Gilhen. from Québec. Photo: L. Bouthillier

FiGuRE 15. Another example of leucism of P. cinereus FIGURE 16. Dorsal view of a partially leucistic P. cinereus
from Nova Scotia. Photo: J. Gilhen. from Québec. Photo: P. Beaupre

nN
Nn
Ss

FIGURE 17. Ventral view of the previous specimen of a
partially leucistic P. cinereus, showing the
translucent aspect of the body. Photo: P.
Beaupre.

FiGure 19. An amelanistic colour anomaly of P. cinereus
from Québec. Photo: J.-D. Moore.

individuals have been reported in Canada (e.g., Piersol
1909; Gilhen 1968; Rosen 1971; Westell and Ross 1974:
Jongsma 201 2a), including a new occurrence for Qué-

bec (Figure 8). This specimen was observed in October

2005 in the La Haute- Yamaska Regional County Munic-
ipality, Québec (45°21'20"N, 72°37'18"W; datum =
NAD83). Jongsma (2012a) reported 11 erythristic indi-
viduals among 85 P. cinereus specimens in Fredericton,
New Brunswick, thus demonstrating that some sala-
mander populations in eastern Canada could neverthe-
less show a high prevalence of erythrism.

Thurow (1955) mentioned that the nature of the ery-
thristic form and its distribution suggest that genetic
rather than environmental factors are involved. Accord-
ing to his interpretation, this phenotype could result
from the action of a mutant allele that quantitatively
inhibits the development of melanin (or possibly mel-
anophores). For intermediate colorations of this morph,
the author noted the possibility of only partial domi-
nance or the involvement of more than one locus. Other
studies suggested that erythristic P cinereus may be the
Batesian mimic of the Eastern Newt (Notophthalmus
viridescens) at the elf stage (Lotter and Scott 1977:
Tilley et al. 1982; Cassell and Jones 2005), which is
known to be toxic for potential predators (Brodie ef al.
1974). This phenomenon would partly explain the high
abundance of the erythristic morph in some forests

where the red eft is present. However, the presence of

the red eft in northern forests does not necessarily imply

THE CANADIAN FIELD-NATURALIST

Vol. 128

FiGuRE 18. A partially leucistic P cinereus from Ohio.
Photo: C. D. Anthony.

FiGure 20. Another example of amelanism of P. cinereus
from Ohio (the red-eye effect is due to the pho-
tographic flash). Photo: J. G. Davis.

that the erythristic morph will be found in these ecosys-
tems (Moore et al. 2012).

Colour Anomalies
The iridistic colour anomaly

Iridism in amphibians is defined as the excessive
production and deposition of iridophore platelets. Indi-
viduals are characterized by generalized iridescent or
metallic, brassy, golden, shades of green or silver flecks
on their dorsal-lateral body. This condition has previ-
ously been described (Muchmore 1955; Hertzler 1958)
and reported for the lead-backed morph of P. cinereus
in the United States (Bogert 1952; Highton 1962,
1972; Schueler 1975; Klemens 1993; Hulse et al. 2001:
Gibbs er al. 2007). However, it was not always clear
from these documents whether the iridescent flecking
was very abundant or visible only upon close examina-
tion. Muchmore (1955) noted that microscopic exam-
ination revealed that this flecking was due to numerous
individual iridophores located around the melano-
phores. These flecks are also known to fade rapidly
on preserved specimens (Grobman 1949; Muchmore
1955; Pfingsten and Downs 1989).

We report here three previously unpublished occur-
rences of iridistic P. cinereus, including, to our know1-
edge, the first documented occurrence of iridism in
the red-backed morph and the first two mentions in
lead-backed morphs from Canada. In all these cases,
the dorsal body exhibited abundant. continuous, and

201 —
4 Moore AND QUELLET: COLOUR PHENOTYPES OF PLETHODON CINEREUS

Nm
nN
Nn

a a, tn eee : bse ,
ri 7 aes American distribution of the 8 different colour morphs and anomalies of the Eastern Red-backed Salamander
ethodon cinereus). Abbreviations: Red = red-backed morph, Lea = lead-backed morph, Ery = erythristic morph, Iri = iridistic

an ae : lets Jape tie ae
anomaly, Alb = albino anomaly, Leu = leucistic anomaly, Ame = amelanistic anomaly, Mel = melanistic anomaly.

large iridescent brassy flecks. The first individual (Fig-
ure 11) was observed in September 2012 in Ashe Coun-
ty, North Carolina (36°25'35"N, 81°35'26"W; datum
= WGS84), and the second (Figure 12) was reported
in June 1982 in Colchester County, Nova Scotia
(45°23'37"N, 63°55'04"W; datum = NAD83). The
third specimen was observed in October 2013 in the
Antoine-Labelle Regional County Municipality, Queé-
bec (46°43'03"N, 75°33'06"W; datum = NAD83).
These occurrences show that iridescent flecking is
sometimes quite abundant and may represent the dom-
inant colour in some P. cinereus individuals. Added with
cases from other studies (e.g., Muchmore 1955), they
also suggest that iridism in P. cinereus is probably more
common than previously reported, and thus deserves
to be regarded as a noteworthy colour variant. To date,
it has been reported in two provinces and 9 states

(Table 2).

The albino colour anomaly

Albinism (from Latin a/bus, white) is defined as the
total absence of pigmentation formation in the skin
and irises. The albino P. cinereus is unstriped and char-
acterized by a flesh-colored body (pink-white to white)
and red-pink irises (Figure 13). According to the liter-
ature, this anomaly has been observed three times in

Ery

Canada (Milnes 1946; Moore ef al. 2012) and three
times in the United States (Fowler 1942; Hensley 1959;
Harris 1968a), though no detailed description of the
specimens was given by Milnes (1946) nor Harris
(1968a). The only case mentioned in Québec (Moore
et al. 2012) has been redefined as an amelanistic P.
cinereus, after the photographic verification showed
normal (dark) pigmentation of the irises, a light orange
dorsal stripe and the pink-white colour of the speci-
men’s dorsal-lateral body. Similarly, the albino case re-
ported by Harris (1968b) was redefined as an amelanis-
tic, based on a published photograph (Zahl 1972).

The leucistic colour anomaly

Leucism (from Greek /eukos, white) is a defect in
the skin causing an inability to support any type of skin
pigments. Since all types of skin pigments are thus re-
duced, leucistic individuals resemble albino P. cinereus
(Figures 14, 15), except for their normally pigmented
irises. This explains why this condition has also been
defined in the past as “albinos with orbital melano-
phores” (Brame 1962), or “partial albinos” (Gilhen
1986). The leucistic anomaly has been observed four
times in Canada (Gilhen 1986 (based on the colour pho-
to NSM973-634-1); Rye 1991; Lamond 1994; Jongsma
2012b) and six times in the United States (Wilmott

256

1945; Pauley 1974; Lotter and Scott 1977 (white par-
tial albino); Mitchell and Mazur 1998; Mendyk et al.
2010). A previously unpublished occurrence of leucism
(Figure 14) was observed in September 2013 in the
Thérése-De Blainville Regional County Municipality,
Québec (45°39'28"N, 73°54'33"W; datum = NAD83).
An unpublished occurrence of partial leucism (Fig-
ures 16, 17) was also reported in September 2005 in
the Nouvelle-Beauce Regional County Municipality,
Québec (46°32'30"N, 71°00'01"W; datum = NAD83).
Leucism was considered as partial in this specimen,
due to the presence of melanophores. Two similar cas-
es have also been described in Ohio (Figure 18, Paluh
et al. 2013).

The amelanistic colour anomaly

Amelanism is a condition characterized by the lack
of melanin production. The appearance of this anomaly
on the skin depends on the remaining non-melanin pig-
ments. Although similar, amelanistic P. cinereus sala-
manders may retain more coloration than the albino and
leucistic anomalies, and are characterized by a body
without black pigmentation (Figures 19, 20). The dor-
sal stripe is typically pale in colour, usually peach-red,
orange or peach-yellow. The dorsal-lateral body is often
pink-white to white, the ventral body is sometimes
translucent, and the iris colour may vary from white to
normal (dark).

This colour anomaly has been reported five times in
eastern Canada (Gilhen 1986 (based on the colour pho-
to NSM984-108-1); Moore and Gilhen 2011; Russell
et al. 2011; see also the above section on the albino
anomaly), and twice in the United States (Davis 2009;
see also the above section on the albino anomaly).
Some authors may sometimes have considered the
amelanistic condition as partial albinism, using termi-
nology such as “albinos with xanthophores” or “partial
albino with xanthophores” and “albinos with erythro-
phores” or “partial albino with erythrophores” (Brame
1962; Harris 1970; Dyrkacz 1981).

The melanistic colour anomaly

Melanism (from Greek melas, black) is a condition
characterized by an overabundance of melanin pig-
ments (melanophores) production and deposition. Mel-
anistic individuals are usually totally black, including
the ventral body and irises. This condition is some-
what the opposite of amelanism. This anomaly in a P
cinereus was first mentioned in Québec (Moore et al.
2012). With its uniformly black and translucent belly,
the melanistic individual is relatively easy to distin-
guish from the lead-backed morph (occasionally very
dark), which has a salt-and-pepper belly appearance.
The melanistic anomaly is not new and has already
been encountered in other salamander and anuran spe-
cies, in which the ventral body is sometimes translu-
cent (Dubois, 1979).

Conclusion
Eight colour phenotypes of the woodland sala-
mander P. cinereus have been reported to date. These

THE CANADIAN FIELD-NATURALIST

Vol.

include main colour morphs as well as rarer colour
anomalies. To discriminate among these different phe-
notypes, useful characteristics are summarized in Table
1, and a dichotomous key is provided (Appendix).
Contrary to amphibian malformations that are mostly
found in young individuals and rarely in adults because
they are maladaptive (Ouellet 2000), most of the P.
cinereus specimens encountered that exhibit colour
anomalies are adults. We agree with Mendyck ez al.
(2010) that these colour conditions may be less detri-
mental for this secretive species than for other amphib-
ians, and thus more likely to be present.

We hope that this article and the greater accessibil-
ity of digital photography will encourage documenting
these atypical cases in future inventories. We suggest
using uniform colour names based on an easily-acces-
sible online list of colours (Wikipedia 2014). This list
is preferred over other guides (e.g., Smithe 1975) which
are rather more difficult to obtain. An ongoing colour
photo gallery is currently online for recognition and
comparison purposes (http://www.amphibia-nature.org
/en/projects/amphibians-reptiles/colorations-plethodon/).
Photos of unusual colorations with the date, location,
and details about the observation can be sent to the
following address: info@amphibia-nature.org. Close-
up pictures that show the colour of the dorsal stripe,
dorsal-lateral body, ventral body, and irises are pre-
ferred.

Acknowledgements

We are grateful to all photographers who kindly con-
tributed pictures of P. cinereus to illustrate this article:
C. D. Anthony (John Carroll University); P. Beaupré
(Ministére du Développement durable, de I’ Environ-
nement, de la Faune et des Parcs du Québec); L.
Bouthillier (Ministére du Développement durable, de
Environnement, de la Faune et des Parcs du Québec);
J. G. Davis (Cincinnati Museum Center); J. Gilhen
(Nova Scotia Museum of Natural History); D. J. Hock-
ing (University of New Hampshire); R. Merrick; D.
E. Swann; and L. A. Williams (North Carolina Wildlife
Resources Commission). We also acknowledge C. Gen-
est (Amphibia-Nature) for sharing her field observa-
tions, P. Galois (Amphibia-Nature) and D. Tousignant
(Forét Québec) for reviewing an earlier version of this
manuscript, J. G. Davis for some museum data contri-
bution, and J. T. Bagnara (Professor Emeritus, Univer-
sity of Arizona), F. R. Cook (Canadian Museum of
Nature), J. Gilhen, T. B. Persons (USGS Southwest
Biological Science Center) and F. W. Schueler (Bishops
Mills Natural History Centre) for helpful comments.

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Received 27 March 2014
Accepted 24 April 2014

2014

MOoRE AND QUELLET: COLOUR PHENOTYPES OF PLETHODON CINEREUS

Appendix

l.a
L.b

2.a
2.b
3.a
3.b
4.a
4.b
S.a
5.b
6.a
6.b
7.a
7.b
8.a
8.b

Dichotomous key of the 8 different colour phenotypes of the Eastern Red-backed Salamander (Plethodon cinereus).

Presence of a continuous or discontinuous dorsal stripe: 2

Absence of a dorsal stripe: 4

Presence of melanophores on the skin surface: 3

Absence of melanophores, dorsal-lateral body pink-white to white: Amelanistic anomaly (Figures 19, 20)
Generalized iridescent flecking: Iridistie anomaly (Figure 11)

No generalized iridescent flecking: Red-backed morph (Figures 1-5)
Presence of melanophores on the skin surface: 5

Absence of skin melanophores: 8

Presence of red or orange pigmentation: Erythristic morph (Figures 8-10)
Absence of red pigmentation: 6

“Salt-and-pepper” appearance of the ventral body: 7

Black and translucent ventral body: Melanistic anomaly

Generalized iridescent flecking: Iridistic anomaly (Figure 12)

No generalized iridescent flecking: Lead-backed morph (Figures 1, 2, 6, 7)
Red-pink pigmentation of the irises: Albino anomaly (Figure 13)

Normal (dark) pigmentation of the irises: Leucistic anomaly (Figures 14-18)

Notes

Long-distance Anadromous Migration in a Fresh Water Specialist: the
Lake Trout (Salvelinus namaycush)

; 4
Les N. Harris!>>, JEAN-SEBASTIEN Moore’, CHRISTOPHER G. McCDermip!:3, and HEIDI K. SWANSON

‘Arctic Aquatic Research Division, Fisheries and Oceans Canada, 501 University Crescent, Winnipeg, Manitoba R3T 2N6
Canada
“Institut de Biologie Intégrative et des Systémes, Université Laval, 1030 Avenue de la Médecine, Québec, Quebec G1 V 0A6

Canada ; -T
Clayton H. Riddell Faculty of Environment, Earth, and Resources — Environmental Science, University of Manitoba, Winnipeg,
Manitoba R3T 2N2 Canada
“Department of Biology, University of Waterloo, 200 University Avenue W., Waterloo, Ontario N2L 3G1 Canada
Corresponding author: Les.N.Harris@dfo-mpo.gc.ca

Harris, Les N., Jean-Sebastien Moore, Christopher G. McDermid, and Heidi K. Swanson. 2014. Long-distance anadromous migra-
tion in a fresh water specialist: the Lake Trout (Sa/velinus namaycush). Canadian Field-Naturalist 128(3): 260-264.

The Lake Trout, Sa/velinus namaycush, is believed to be one of the most saline intolerant salmonid species, typically com-
pleting its life wholly in fresh water. Historical observations and more recent quantitative assessments have shown, however,
that in some Arctic populations, Lake Trout can migrate to marine waters (i.e., display anadromy). In the four coastal Arctic
populations of Lake Trout where anadromy has been confirmed, migrations to and from marine environments are relatively
short (i.e., in the order of a few kilometres). In the Halokvik River on Victoria Island, we captured two anadromous Lake
Trout in a weir used jointly for commercial fishing and stock assessment research. Both fish were captured during the fall up-
stream migration, some 50 km from their presumed fresh water spawning or overwintering locations. This observation extends
the current knowledge regarding the distribution of anadromous populations in this species and suggests that migration to marine

habitats can be much longer than previously expected.

Key Words: Lake Trout; Salvelinus namaycush; anadromy; migration; Victoria Island

Introduction

Migration is a coordinated movement that is evolu-
tionarily advantageous because of the fitness benefits
associated with movement to an alternate habitat (Hen-
dry et al. 2004; Binder et al. 2011). Fishes in particular
are well known for extraordinary migrations that can be
classified based on the aquatic biomes through which
they move. Diadromy is a type of migration in which
the marine—fresh water boundary is crossed (Binder ef
al. 2011), and anadromy is a unique type of diadromy,
in which fishes hatched in fresh water subsequently
migrate to marine habitats for feeding and, eventually,
undertake a return migration to fresh water for spawn-
ing, overwintering, or both (McDowall 1997, 2007).
This life history strategy is exemplified in the salmonid
fishes and is typically more prevalent at higher latitudes
(Gross et al. 1988; McDowall 2008). Anadromous mi-
grations within this group can be quite remarkable,
ranging from thousands of kilometres (Stephenson et al.
2005) to just hundreds of metres (Harris et al. in press).

Among salmonids, Lake Trout (Salvelinus namay-
cush) are generally thought to be the least tolerant of
saltwater conditions (Quinn 2005). Lake Trout were
typically believed to be invariably non-anadromous
(Hendry et al. 2004), but were recently shown to be able
to survive salinities equal to that of full-strength sea
water (i.e., 30%, Hiroi and McCormick 2007). Lake

Trout have been observed sporadically, although rarely,
in marine waters (dating back to the 1950s and reviewed
in Martin and Olver 1980), and recent data from otolith
microchemistry and stable isotope analyses confirm that
some coastal Canadian Arctic populations of Lake Trout
do make annual migrations to marine environments
(Swanson et al. 2010). Among four study populations,
27% of Lake Trout made annual migrations to the sea.
and anadromous Lake Trout were in better condition
and had lower concentrations of mercury than fresh
water resident Lake Trout (Swanson and Kidd 2009:
Swanson et al. 2010). Further study using isotope mix-
ing models found that marine prey items constituted a
large proportion (66%) of the diet of anadromous Lake
Trout (Swanson ef al. 2011).

Despite this recent increase in our knowledge of
anadromy in the Lake Trout, much remains unknown
regarding the distribution of its life history across the
northern range of the species and the magnitude of the
migrations it undertakes. Where anadromy has been
confirmed in Lake Trout, migration distances have been
relatively short (< 3 km, Swanson et al. 2010), suggest-
ing that Lake Trout employing this life history tactic
are associated with systems in which the distance to
marine waters is short. Here, we describe a case of long-
distance migration in anadromous Lake Trout based on
observational evidence and capture data accumulated

260

2014

Halokvik Lake

NOTES

261

Wellington
Bay

__ Halokvik (30 Mile) River

~~,

Ficure |. Map of the study area showing the location of the Halokvik River (30 Mile) on Victoria Island, Nunavut. Also shown
is the location of the weir constructed in 2013 (red rectangle), the position of the acoustic receiver deployed in the river
(black circle), and Halokvik Lake where anadromous Lake Trout (Sa/velinus namaycush) are presumed to overwinter,

some 50 km from the weir.

from a fisheries weir assessment on the Halokvik River
(locally known as 30 Mile), Victoria Island. This river
is located in the Canadian Arctic (Figure 1), approxi-
mately 80 km west of the community of Cambridge
Bay in Nunavut.

Methods and Observations

In 2013, Fisheries and Oceans Canada operated a
conduit weir on the Halokvik River with the intention
of enumerating anadromous Arctic Char (Sa/velinus
alpinus) migrating upstream. The Halokvik River
drains a large system of lakes that flow into the west
side of Wellington Bay. The last lake draining into
this river, locally known as Halokvik Lake, is located
some 50 km upstream from the ocean and is presumed
to be the furthest downstream lake capable of sup-
porting Arctic Char and Lake Trout (J. Hiniliak, Cam-
bridge Bay elder, personal communication, 2013).

The weir, which spanned the entire river (Figure 2)
to ensure that all upstream-migrating fish were cap-
tured, was designed following the methods outlined
by McGowan (1990). It was located near the river
mouth, 50 m from tidal-influenced brackish waters.
The weir was operated continually from August 8 to
September 10, 2013. In mid-August 2013 (August 16
and 18), two Lake Trout (Figure 3) were captured in

the weir as they were most certainly migrating back
into fresh water to spawn and/or overwinter, presum-
ably in habitats located 50 km upstream. Lake Trout
1 was 576 mm in fork length and weighed 2100 g
and Lake Trout 2 was 531 mm in length and weighed
1700 g. Both fish were photographed for morphome-
tric analyses, tissue samples were obtained, and the
opportunity was taken to implant both with acoustic
transmitters (V16 Coded Acoustic Transmitters,
VEMCO, Bedford, Nova Scotia).

Discussion

Although the Lake Trout is often believed to be res-
tricted to fresh water for its entire life cycle, evidence
that anadromy may be common in high latitude popu-
lations is accumulating. Populations in which anadro-
my has previously been documented, however, have
been found at very short distances from the marine
environment (within 3 km, Swanson e/ a/. 2010). Here,
we report the capture of two Lake Trout returning from
marine waters some 50 km downstream from the clos-
est lake where overwintering and/or spawning would
be possible. It is highly unlikely that these two fish
were riverine Lake Trout, given that the Halokvik Riv-
er freezes solid during winter over most of its course.
Furthermore, they were captured in a weir approxi-

: FR
262 THE CANADIAN FIELD-NATURALIST Vol. 128

FIGURE 2. Aerial view of the weir across the Halokvik River, Victoria Island, Nunavut, used in 2013 as part of an Arctic Char
(Salvelinus alpinus) population assessment. The red arrow indicates the upstream direction of migration. Photo courtesy
of Denise LeBleu Images.

FIGURE 3. The two Lake Trout (Sa/velinus namaycush) captured in the Halokvik (30 Mile) River weir.
while migrating from marine waters to spawning and overwinterin

(B) Lake Trout 2.

( Victoria Island, Nunavut,
g locations located 50 km upstream. (A) Lake Trout 1:

2014

mately 50 m from tidal influenced waters and, given
the energy costs of migration, it is highly unlikely that
Lake Trout would migrate 50 km to exploit fresh water
river habitat, which is much less productive than the
adjacent brackish and marine habitats. Thus, to our
knowledge, this represents the longest documented
anadromous migration of Lake Trout, highlighting an
unappreciated ability for long-distance migration to
and from marine environments in a species that was
once thought to be restricted to fresh water.

Local knowledge in the Cambridge Bay region in-
dicates that Lake Trout are sometimes caught in marine
waters (B. Nakashook, Cambridge Bay resident, per-
sonal communication, 2013). Recently, quantitative
evidence of anadromy in Arctic populations of Lake
Trout has become available to corroborate this local
knowledge (Swanson ef a/. 2010) and, combined,
these pieces of information indicate that anadromy in
this species may not be as rare as once assumed. For
instance, in Nauyuk Lake, a system located on the
mainland of Canada approximately 90 km southwest
of the Halokvik River, 40% of assessed Lake Trout
were classified as anadromous (Swanson ef al. 2010).
However, marked variation occurred in the proportion
of resident versus anadromous fish in other proximate
systems (Swanson et a/. 2010) and, currently, no ful-
ly anadromous populations are known. Further work
is thus required to truly understand intra- and inter-
population variation in anadromy in this species and to
resolve the specific mechanisms responsible for driv-
ing such differences.

Information on the population and biological param-
eters of anadromous Lake Trout are virtually absent
(but see Swanson et al. 2010, 2011). Little is known
regarding sizes and ages of anadromous individuals,
however, among the four systems located across Dease
Strait that were studied by Swanson et al. (2010), age
of first migration in Lake Trout varied widely: from 3
to 29 years of age (mean, 13 years). Lake Trout started
migrating to sea at a significantly older age on average
than sympatric anadromous Arctic Char (5 years of
age), and at 13 years of age, first-year migrant Lake
Trout were larger (mean 400 mm fork length) than first-
year migrant Arctic Char (mean 275 mm, Swanson ef
al. 2010). The anadromous Lake Trout observed in this
assessment are within the length ranges reported by
Swanson et al. (2010).

Little information is available on the distribution of
anadromy in Lake Trout, and further work is required
to quantify the prevalence of this life history tactic
throughout the northern range of the species. There are,
however, reports of anadromous Lake Trout on other
Canadian Arctic islands (Manning 1953), and work is
ongoing to provide the first assessment on the distri-
bution of anadromy in this species across the coastal
Canadian Arctic. Finally, marine habitats used by an-
adromous Lake Trout are not well characterized. As
with other salmonids, they likely prefer brackish coastal

NOTES

263

waters for feeding on marine invertebrates and fishes
(Swanson ef a/. 2011). Acoustic receivers are currently
positioned in the brackish and marine habitats near
the mouth of the Halokvik River, and future acoustic
telemetry readings may provide the first data on marine
habitat use by this species.

Given the considerable distance presumably covered
by Lake Trout in our study, marine migrations are not
likely opportunistic forays, but rather concerted efforts
to access marine habitats. High energetic costs of
migration and the potential increased risk of predation
(Hendry ef al. 2004) mean that the fitness benefits of
migrating to marine habitats must be substantial and
must outweigh the fitness benefits of remaining in fresh
water (Gross 1987). Indeed, oceans are much more
productive than fresh water in north-temperate and
Arctic environments, and the prevalence of anadromy
appears to increase at higher latitudes (McDowall 1987;
Gross et al. 1988). Thus, it is likely that anadromous
Lake Trout observed in the Halokvik River were ac-
cessing marine habitats to take advantage of rich food
sources and that this results in higher growth rates,
larger size-at age, and greater energy stores (Hendry
et al. 2004). This in turn, may result in higher fecun-
dity and overall fitness compared with their fresh water
counterparts (Roff 1988). Swanson ef a/. (2010) found
that the diet of anadromous Lake Trout was composed
of about 66% marine-derived prey (based on stable
isotope mixing models) and that anadromous Lake
Trout were typically in better condition than their fresh
water counterparts. This is consistent with the hypoth-
esis that anadromous Lake Trout are migrating to ex-
ploit relatively rich marine food sources.

In conclusion, our observations provide novel insight
into the distribution of anadromy in Lake Trout pop-
ulations in the Arctic, now confirming it on Victoria
Island, and highlight the potential for long-distance
migration in this species as a tactic in a life history that
has remained relatively enigmatic.

Acknowledgements

We thank the Ekalluktutiak Hunters’ and Trappers’
Organization for help with logistic planning over the
course of the field season. Jack (Meyok) Omilgoetok
worked on all aspects of the field work and was instru-
mental in the success of our field season. Jimmy Hanil-
iak helped with tagging at the Halokvik River and
Johnny Pedersen assisted with weir construction. We
also thank Jimmy Haniliak and Brent Nakashook for
providing information on the Halokvik River system.
Funding for this project was provided by Fisheries and
Oceans Canada (through Nunavut Implementation
Funds) and the Nunavut Wildlife Management Board.

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265

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New Records of the Ogilvie Mountains Collared Lemming (Dicrostonyx
nunatakensis) in central Yukon

, > 11,4 a “Ty2 3
THOMAS S, JUNG!:4, BRIAN G. SLOUGH?, Davip W. NAGORSEN®, AND PtlA M. KUKKA!

‘Yukon Department of Environment, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada
“35 Cronkhite Road, Whitehorse, Yukon Y1A SS9 Canada

“Mammalia Biological Consulting, Victoria, British Columbia V9C 324 Canada
*Corresponding author: thomas.jung@gov.yk.ca

Jung, Thomas S., Brian G. Slough, David W. Nagorsen, and Piia M. Kukka. 2014. New records of the Ogilvie Mountains Col-
lared Lemming (Dicrostonyx nunatakensis) in central Yukon. Canadian Field-Naturalist 128(3): 265-268.

The Ogilvie Mountain Collared Lemming (Dicrostonyx nunatakensis Youngman, 1967), reported only from the Ogilvie
Mountains of central Yukon, is among the least known mammals in Canada. It was first discovered in 1961 and, since then,
only 13 specimens had been collected, all from one mountain, in central Yukon. We conducted a targeted survey to determine
the distribution of the species by trapping areas of apparently suitable habitat on 12 mountains within 40 km of the known
location, Many of our traps were disabled by other mammals; however, we captured three Ogilvie Mountain Collared Lem-
mings on two mountains 25.9 km and 29.6 km from the original location. Our findings suggest that this lemming may be
more widely distributed than indicated by earlier specimens. We suggest further surveys to delineate the range of the Ogilvie
Mountain Collared Lemming.

Key Words: Ogilvie Mountain Collared Lemming; Dicrostonyx nunatakensis; distribution; Tombstone Territorial Park; Yukon

Introduction

The Ogilvie Mountain Collared Lemming (Dicro-
stonyx nunatakensis Youngman, 1967) is known from
only one mountain in central Yukon, Canada. Collared
Lemmings in the Ogilvie Mountains are geographi-
cally separated from populations of Nearctic Collared
Lemming (Dicrostonyx groenlandicus) to the northeast
by about 250 km of apparently unoccupied habitat.
The nearest recorded population of Nearctic Collared
Lemming is in the Richardson Mountains, northeast of
the Ogilvie Mountains. Youngman (1967) speculated
that Collared Lemmings in the Ogilvie Mountains rep-
resent a relic population of Dicrostonyx that became
isolated on nunataks (unglaciated mountain tops) dur-
ing the Last Glacial Maximum and that they may
remain reproductively isolated. Although the Ogilvie
Mountain Collared Lemming is afforded species status
by Wilson and Reeder (2005), the taxonomic status of
Collared Lemmings in the Ogilvie Mountains is unre-
solved (J. Eger, Royal Ontario Museum, personal com-
munication).

The Ogilvie Mountain Collared Lemming ts one of
the least known terrestrial mammals in North America
(Nagorsen 1998). Only 13 museum specimens exist, all
collected from a single mountain, Trapper Mountain
(64.583°, =138.217°), located in Tombstone Territorial
Park, Yukon (Youngman 1964, 1967, 1975; Slough and
Jung 2007). The specimens were collected at 1625—
1676 m above sea level in dry rocky alpine tundra
(heath) at the base of glacial cirques.

Limited survey effort has hampered our ability to
assess accurately the distribution of this apparently
range-restricted small mammal. Because surveys have
been constrained by the remote high-elevation habi-
tats in the Ogilvie Mountains, it is unknown whether
this species is restricted to a single mountain. We con-

ducted a targeted survey for the Ogilvie Mountain Col-
lared Lemming to determine whether it inhabits other
nearby mountains. This information is essential for bet-
ter understanding the distribution and conservation sta-
tus of this population.

Study Area

During 19-31 August 2011, we surveyed 12 moun-
tains in the southern Ogilvie Mountains, all located
within Tombstone Territorial Park. Tombstone Territo-
rial Park is approximately 2,200 km?, much of which
lies above the tree line. Our sampling sites were distrib-
uted over five distinct mountain ranges (Table 1 and
Figure 1). The mountains we surveyed were distrib-
uted throughout the park to provide spatial coverage
of the alpine habitat within the park. We did not sam-
ple at Trapper Mountain because the species was al-
ready known from there (Youngman 1964, 1967, 1975;
Slough and Jung 2007). Survey sites were accessed
by helicopter.

Methods

On each mountain surveyed, we selected a sampling
site in alpine tundra that was similar in elevation and
habitat characteristics (dry heath, gentle slope) to cap-
ture sites on Trapper Mountain (Youngman 1975; Jung
et al., unpublished data). At each sampling site, we set
snap-traps in variable-length trap lines that contained
60 trapping stations (49 in | case) set 10-15 m apart.
At each trapping station, we set both a Museum Spe-
cial and a Victor snap-trap (Woodstream Corp., Lititz,
Pennsylvania) |—2 m apart. Traps were baited with
peanut butter and rolled oats and left unattended for
9-11 days (x = 10.3 + 0.3 standard deviation [SD]:
Table 1). We recorded the number of specimens cap-
tured per trap line and the number of traps that were

266 THE CANADIAN FIELD-NATURALIST

a
© Vintenone aM
y

ae

\ Tombstone erritorial Park

_

FiGureE |. Boundaries of Tombstone Territorial Park, Yukon,
Canada, and the 12 sites sampled for the Ogilvie
Mountains Collared Lemming (Dicrostonyx nunatak-
ensis) in 2011. Filled circles indicate locations where
lemmings were captured, open circles where they
were not detected. Note: Trapper Mountain is the only
mountain where specimens were previously collected.

sprung or not functioning at the end of the sampling
period. Our sampling protocol represented a maximum
sampling effort of 14 638 trap nights (1 trap night = 1
trap set for | night).

Captured animals were identified by morphology
and pelage. Small mammals were identified using the
key provided by MacDonald (2003). Identification of
the Ogilvie Mountains Collared Lemming was based
on morphologic and pelagic characteristics, particu-
larly very short tail and ears, densely furred feet, and
buff-gray fur with a black dorsal stripe (MacDonald
2003). We deposited specimens at the Royal Ontario
Museum (Toronto, Ontario).

Results and Discussion

Overall the number of animals captured was small:
26 captures of small mammals of four species, includ-
ing three specimens of the Ogilvie Mountains Collared
Lemming (Table |). Ogilvie Mountains Collared Lem-
mings were collected from two of the 12 mountains
surveyed, both in the Seela Range (Table 1). The two
sites were 5.7 km apart and 25.9 km and 29.6 km west
of previous collections on Trapper Mountain. Capture
locations were among the most distant sampling sites
from Trapper Mountain, and they were separated from

TABLE 1. Summary of site information, sampling effort, and trap captures during a targeted survey of the Ogilvie Mountains Collared Lemming (Dicrostonyx nunatakensis) in Tomb-

stone Territorial Park, Yukon, Canada, August 2011.

Site information

Latitude

Trap captures*

Sampling effort

Montane

Tundra

Northern

Collared Lemming Red-backed Vole

Ogilvie Mountains

% sprung

No. trapping

Elevation

Longitude

Mountain

Shrew

Vole

5

nights trapst

traps set

(°N) (°W) (m)

range

Site

76.7

84.7

+

86.7

0

0

100.0
60.8

len |

64.2

0

86.7

0

120

0

oS

0
0

0

75.0

Vol.

AN

co

ON

WES

Tombstone

64.6414

Blackstone

5

Tombstone

Tombstone

Prospector

Prospector

64.8221

McFarland

10

64.6703

Blackstone

Seela

64.6686

*Note: Northern Red-backed Vole, Myodes rutilus; Tundra Vole, Microtus oeconomus; Montane Shrew, Sorex monticolus.

+Traps set off without apparently capturing an animal.

{No data.

128

2014 NOTES

Trapper Mountain by the Blackstone River, a potential
barrier to dispersal (Figure 1).

Two Ogilvie Mountains Collared Lemmings were
juvenile males and the other was a juvenile female.
They were collected in dry heath type habitat at ele-
vations (1631-1709 m above sea level; Figure 2) typi-
cal of previous captures at Trapper Mountain (Young-
man 1967; Slough and Jung 2002). Vegetation at the

267

capture sites was dominated by mountain avens (Dryas
spp.), dwarf willow (Salix spp.), mosses, lichens
(Thamnolia vermicularis, Flavocetraria nivalis), and
grasses (e.g., Anthoxanthum monticola), along with
other ericaceous species (e.g., Arctous alpinus, Betula

glandulosa, Empetrum nigrum, Cassiope tetragona,

Vaccinium vitis-idaea; Figure 2). Small mammal
species caught in association with the Ogilvie Moun-

i r | ine hez ahit: re > > Mo ai ‘ol-
2. Landscape (top) and micro-site (bottom) views of the high alpine heath habitat where the ven ie Mc = He Co
JURE 2. Landscape Op? ¢ ; central Vi ‘anada. / Scale shown in
or | i L tae (Dicrostonyx nunatakensis) was collected in central Yukon, ¢ anada, August 2011. Scale s
ared Le gU SEN

the bottom panel is

approximate. Photos: Brian Slough (top) and Thomas Jung (bottom).

268

tains Collared Lemming were the Northern Red-backed
Vole (Myodes rutilus), the Tundra Vole (Microtus oeco-
nomus), and the Montane Shrew (Sorex monticolus;
Table 1).

Many traps (x = 81.2% + 3.7% SD) were prema-
turely sprung or missing when we returned to check
them (Table |), which substantially reduced the sam-
pling effort. Based on earlier survey work using the
same trapping protocol in the study area (Jung e7 al.,
unpublished data), we suspect that most traps were
disabled early within the sampling period, resulting in
the low number of small mammals captured. However,
the time from deployment during which our traps were
functional is unknown. Arctic Ground Squirrels (Uro-
citellus parryii) were abundant at all sampling sites,
and we suspect that they were responsible for the vast
majority of traps that were prematurely sprung or miss-
ing. For further surveys, means should be sought to
reduce disturbance of traps by Arctic Ground Squir-
rels, such as placing a wire enclosure over the trap.

We confirm that Ogilvie Mountains Collared Lem-
mings are more widely distributed than solely on Trap-
per Mountain. We suspect that they were present on
some of the 10 mountains we surveyed without detect-
ing them. The habitat at all sites surveyed was similar.
The functional loss of many traps likely resulted in a
low sampling effort, hindering our ability to capture
collared lemmings at some survey sites. The occur-
rence of Ogilvie Mountains Collared Lemmings on
three mountains within the Ogilvie Mountains suggests
that they may be somewhat widespread in suitable habi-
tats there. Moreover, Youngman (1975) speculated that
the Ogilvie Mountains Collared Lemming likely also
occurs in the adjacent Wernecke and Selwyn Moun-
tains of central Yukon, but there have been few (if any)
small mammal surveys in these ranges. Further sur-
veys are needed to delineate the range of the Collared
Lemming within the Ogilvie Mountains and adjacent
ranges.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Acknowledgements

We thank D. Amos, C. Eckert, and T. Powell for
facilitating this work and assisting with field logistics.
We thank L. Hughes, S. Hughes, M. Kienzler, and A.
McCulley for field assistance. We thank J. Eger for
discussions about collared lemmings and archiving our
specimens at the Royal Ontario Museum. Two anony-
mous reviewers provided thoughtful comments on an
earlier draft of this manuscript.

Literature Cited

MacDonald, S. O. 2003. The small mammals of Alaska: a
field handbook of the shrews and small rodents. Unpub-
lished manuscript. Accessed 20 September 2013. http://
science.nature.nps.gov/im/units/swan/assets/docs/reports
/inventories/MacDonaldS 2003 AKRO_ SmallMammal
HandbookCondensed_572288.pdf

Nagorsen, D. W. 1998. Dicrostonyx nunatakensis. Page 88 in
North American Rodents: Status Survey and Conservation
Action Plan. Edited by D. J. Hafner, E. Yensen, and G. L.
J. Kirkland. IUCN Publications, Gland, Switzerland.

Slough, B. G., and T. S. Jung. 2007. Diversity and distribu-
tion of the mammals of the Yukon Territory: a review.
Canadian Field-Naturalist 121: 119-127.

Wilson, D. E., and D. A. Reeder (editors). 2005. Mammal
Species of the World. A Taxonomic and Geographic Ref-
erence. Third edition. The Johns Hopkins University Press,
Baltimore, Maryland, USA. 2142 pages.

Youngman, P. M. 1964. Range extensions of some mammals
from Northwestern Canada. Natural History Papers 23.
National Museum of Canada, Ottawa, Ontario, Canada.

Youngman, P. M. 1967. A new subspecies of varying lem-
ming, Dicrostonyx torquatus (Pallas), from Yukon Territo-
ry (Mammalia, Rodentia). Proceedings of the Biological
Society of Washington 80: 31-34.

Youngman, P. M. 1975. Mammals of the Yukon Territory.
Publications in Zoology 10. National Museum of Canada,
Ottawa, Ontario. 192 pages.

Received 2 October 2013
Accepted 3 February 2014

2014

NOTES 269
A Note on Bird Song: Samuel Hearne’s Observations on the Snow
Bunting (Plectrophenax nivalis)
Davip L. G. Noakes!: and JEFFREY D. NOAKES?
'Fisheries and Wildlife Department and Oregon Hatchery Research Center, 104 Nash Hall, Oregon State University, Corvallis,

Oregon 97331-3803 USA
“Canadian War Museum, | Vimy Place, Ottawa, Ontario K1S 4V5 Canada
*Corresponding author: david.noakes@oregonstate.edu

Noakes, David L. G., and Jeffrey D. Noakes. 2014. A note on bird song: Samuel Hearne’s observations on the Snow Bunting
(Plectrophenax nivalis). Canadian Field-Naturalist 128(3): 269-271.

In 1795, the Arctic explorer Samuel Hearne recorded detailed observations on the distribution, ecology, molt, and behaviour
of the Snow Bunting (Plectrophenax nivalis). The most significant of his observations was that Snow Buntings imitated the
vocalizations of Atlantic Canaries (Serinus canarius) when housed with that species. His account has apparently not been

widely recognized by ornithologists, but it is one of the first such observations on bird’s acquisition of vocalizations.

Key Words: Arctic; bird song; Atlantic Canary; Snow Bunting;

Introduction

The study of avian vocalizations, especially their
acquisition and development, is a central part of what
is often recognized as “classical” ethology (Kroods-
ma and Miller 1996). Niko Tinbergen was one of the
founders of ethology (Tinbergen 1951), and details of
his field studies of Snow Buntings (Plectrophenax
nivalis) in Greenland remain a classic (Tinbergen 1939;
Dawkins et al. 1991). This note brings attention to an
apparently unrecognized early description of the vocal-
izations of Snow Buntings in the written accounts of a
European Arctic explorer, Samuel Hearne.

The species-typical nature of avian vocalizations has
long been recognized and is the basis for field identi-
fication by expert field naturalists (Baptista 1975). Per-
nau is now recognized for attributing birds’ acquisition
of species-typical song to learning in the 16th century,
but his observations are not widely known or generally
appreciated (Birkhead 2003). Some of the earliest ex-
perimental observations on the acquisition of avian
vocalizations were reported in the scientific literature
by Barrington (1773). He hand reared young Linnet
(Carduelis cannabina) and concluded that they ac-
quired their vocalizations as a result of early exposure
to those of adult birds. His experimental approach
showed the way for many who followed (Kroodsma
and Baylis 1982); however, it was not until recording
and playback equipment became available that much
progress could be made in the study of avian vocal-
izations (Baker 2001). The considerable advances in
the study of avian vocalizations from the 1950s were
a result of the use of equipment, especially the sono-
graph, that had been developed during World War II
for detecting and recording sounds (Thorpe 1954).

Our understanding of the development of avian
vocalizations is well past the classical studies of Mar-
ler (1952) and Thorpe (1954). Complexities of song
acquisition include both auditory and visual cues, vary-
ing degrees of plasticity, dialects, vocal mimicry, and
restricted learning (Brooke and Birkhead 1991). De-

Samuel Hearne; Serinus canarius; Plectrophenax nivalis

tailed models have been proposed to account for song
learning in birds, including instruction, selection, and
instruction followed by selection in production (Mar-
ler 1997). However, what might appear to be simple
descriptions of natural history observations, such as
those by Hearne, are always valuable in any area of sci-
ence. In this case, they document an important histor-
ical accomplishment that deserves credit in itself. They
are also important as they may be useful to test the gen-
erality of our current hypothesis or suggest yet another
interesting prediction to be tested in another species.

Historical Record

Samuel Hearne was one of the first European explor-
ers in the region that is now part of the Canadian Arc-
tic (Cavell and Noakes 2010). His writings (Hearne
1911) provide a wealth of details on numerous mun-
dane aspects of daily routines as well as more unique
events. He gave detailed accounts of the distribution
and abundance of a number of bird species, mostly in
relation to hunting them as food. However, he also
remarked on the behaviour of Snow Buntings that he
held as part of an extensive menagerie (Newman 1998).
He described how they could be captured, or shot, at
different times of the year, and he recorded that, when
held in the company of exotic Atlantic Canaries (Ser-
inus canarius), Snow Buntings learned to imitate the
song of the canaries.

With reference to the vocalizations of Snow Bunt-
ings, Hearne wrote: “They live long in confinement,
have a naturally pleasing note, and when in company
with Canary birds soon imitate their song. I have kept
many of them in cages in the same room with Canary
birds, and always found they sung in Winter as well as
in Summer; but even in confinement they change their
plumage according to the season, the same as in a wild
state” (Hearne 1911, page 386).

Unfortunately, his description lacks some valuable
details. For example, the sex and age of the Snow Bunt-
ings, particularly as they might relate to song acquisi-

tion and development, are not given in his descrip-
tion. We now know from a wealth of detailed experi-
mental studies a good deal about song development in
Oscines, the group that includes the Snow Bunting
(Marler 1997). We know that these birds develop their
songs by learning early in ontogeny. It seems unlikely
that Hearne would have reared Snow Buntings in cap-
tivity from the early age (about 20 days after hatching)
that is required to demonstrate initial song learning.
Thus, it would seem that he observed Snow Buntings
changing their songs later in life, after they had ac-
quired their species-typical songs in nature. There is
evidence that some oscine species can change their
songs later in life, as a result of exposure to vocaliza-
tions from other species (Marler 1990; Beecher and
Brenowitz 2005; Laiolo et al. 2011).

Hearne’s descriptions do not indicate whether his
observations refer only to males or to both sexes of
Snow Buntings. Females of some oscine birds do vocal-
ize (Brenowitz and Arnold 1986). Hearne did not report
whether the Snow Buntings replaced their song with
the canary songs. He also does not provide any further
details as to how he managed to hold the birds alive
for such a long time under what were terribly demand-
ing conditions for humans, let alone their domestic
companion species (Newman 1998). Incredibly, he
does not mention how or why he happened to have
Atlantic Canaries available to him during his explo-
rations of the North American Arctic!

Discussion

Hearne’s Arctic accounts are widely and well known,
but his observations of Snow Bunting vocalizations
have not been noted before, despite reviews in the
ornithological literature (Houston 1989). There is ex-
tensive literature on the domestication of the Atlantic
Canary that dates from at least the 16th century. The
songs of the canaries and their colours attracted a great
deal of attention, and the birds were widely distrib-
uted by amateur bird keepers throughout Europe and
Britain (Birkhead 2003). The 18th century European
explorers who came to North America would certainly
have been familiar with canaries, but it is remarkable
that they brought them on their voyages (Houston ef
al. 2003).

Recent descriptions of the behaviour and ecology
of Snow Buntings have included descriptions of their
vocalizations (Thorpe and Lade 1961; Lyon and Mont-
gomerie 1985, 2011;Espmark 1995, 1999; Alsop 2001;
Hofstad et al. 2002; Meltofte 2007). Espmark (1995)
concluded that the song of the Snow Bunting is stereo-
typic, as it is for most Emberizidae species. Alsop
(2001) describes the song as “a series of bold repeti-
tive high trilling notes,” produced only on the breed-
ing grounds. Snow Buntings are reported to have indi-
vidual song patterns and Espmark (1995) concluded
that there is no strong evidence of local song dialects.
There has been no detailed study of the development

THE CANADIAN FIELD-NATURALIST

Vol. 128

of vocalizations in this species; thus, we do not know
whether they learn their song by instruction or selec-
tion, nor do we know the period during which they can
acquire their song. However, Hearne clearly deserves
recognition for his insightful early observations on the
development of behaviour, in addition to his broader
contributions to our understanding of Arctic biology.

Acknowledgements

This note is dedicated to the memory of a friend and
graduate classmate, Luis Baptista, who knew more
about the languages of people and birds than anyone.
We thank Dr. Doug Robinson, Oregon State Univer-
sity, and Dr. David Sherry, University of Western
Ontario, for their reviews of an earlier draft of this
manuscript. The detailed comments of anonymous re-
viewers and the editor significantly improved this man-
uscript and are very much appreciated.

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Received 31 October 2013
Accepted 10 February 2014

Vol. 128

THE CANADIAN FIELD-NATURALIST

Late-winter Habitat Use by the Fisher, Pekania pennanti (Erxleben,
1777), in the Boreal Plains Ecozone of Northwestern Saskatchewan,
Canada

GILBERT PROULX

Alpha Wildlife Research & Management Ltd., 229 Lilac Terrace, Sherwood Park, Alberta T3H 1W3 Canada; email: gproulx
@alphawildlife.ca

Proulx, Gilbert. 2014. Late-winter habitat use by the Fisher, Pekania pennanti (Erxleben, 1777), in the Boreal Plains Ecozone
of northwestern Saskatchewan, Canada. Canadian Field-Naturalist 128(3); 272-275.

Late-winter habitat use by the Fisher, Pekania pennanti (Erxleben, 1777) in northwestern Saskatchewan was assessed in
February 2009, 2011, and 2012. A total of 78 Fisher tracks were recorded over 60 300 m of snowshoe surveys. Fisher tracks
were significantly less frequent than expected in Tamarack (Larix laricina [Du Roi] K. Koch) stands with > 40% crown clo-
sure and mainly 0-10 m trees (P < 0.05) and in open areas. Fishers used other habitat types equal to availability, including
muskeg and coniferous, mixed, and deciduous forest stands. Maintaining mosaics of forest stands of different seral stages
interspersed with muskeg would meet the late-winter habitat needs of Fishers in the Boreal Plains Ecozone of northwestern

Saskatchewan.

Key Words: Fisher; Martes pennanti; Pekania pennanti; muskeg; Saskatchewan; winter habitat; Boreal Plains

Introduction

In western Canada, Fisher (Pekania pennanti |Erx-
leben, 1777]; formerly Martes pennanti, Sato et al.
2012) habitat studies have been conducted in conifer-
ous, mixed, and deciduous forests (Badry et al. 1997;
Weir and Harestad 1997; Proulx 2006). However, none
has been carried out in the Boreal Plains Ecozone of
northern Saskatchewan, where treed fens and bogs (re-
ferred to as muskeg) are abundant and the Fisher is a
species of economic importance because of its value as
a furbearer and its role in supporting the subsistence of
Aboriginal groups (Koback 2014*). The purpose of this
study was to assess late-winter habitat use by the Fisher
in northwestern Saskatchewan.

The study was conducted in Mistik’s Forest Manage-
ment Agreement (FMA) area, north of Meadow Lake
(54°07'N, 108°25'W), adjacent to the Alberta border.
The area encompasses approximately 1.8 million ha of
forest, water, and non-forested land. It is a mosaic of
upland deciduous and coniferous boreal forest, muskeg,
and water (McLaughlan ef al. 2010) resulting from fre-
quent fires of various sizes and intensities (Parisien ef
al. 2004). The area lies within the Boreal Plains Eco-
zone where White Spruce (Picea glauca [Moench]
Voss), Black Spruce (Picea mariana [Miller] Britton,
Sterns & Poggenburgh), Jack Pine (Pinus banksiana
Lambert), and Tamarack (Larix laricina [Du Roi] K.
Koch) are the dominant conifers (Environment Canada
2008). Deciduous stands of Trembling Aspen (Populus
tremuloides Michaux), Balsam Poplar (Populus bal-
samifera Linnaeus), and Paper Birch (Betula papyrifera
Marshall) occur throughout the study area (Rowe 1972).

Field inventories were carried out in February 2009,
2011, and 2012. The range of temperatures was similar
among years (0 to —30°C), but, based on monthly
average temperatures for the region, the 2012 winter
(—10.8°C) was slightly warmer than 2009 (—16.3°C)

and 2011 (—16.5°C) (Munroe 2012*; Government of
Canada 2014*). During inventories, snow depths ranged
from 30 to 65 cm in 2009 and 2011, and 30 to 45 cm
in 2012. The survey method followed Proulx (2006,
2011). A random-stratified approach (Krebs 1978) was
used to locate linear transects averaging > | km long
and = | km apart that crossed all habitat types. Differ-
ent transects were surveyed from year to year. Tran-
sects were plotted on 1:15 000 scale maps, and start-
ing points were located using compass bearings and
distance to distinct photographic features. Transects
were followed on snowshoes, using a compass and a
hip chain (a device used to record linear distances).
Transect lengths varied according to accessibility, safe-
ty, and environmental conditions (e.g., open water, un-
expected snowstorm, etc.). I recorded only well-defined
tracks: those not melted or deformed, not filled with
crusty snow, and judged to be less than 48 h old (a sub-
jective assessment based on my experience). Because
Fisher and American Marten (Martes americana [Tur-
ton, 1806]) footprints are similar (Halfpenny er ai.
1995), when mustelid tracks were encountered, I inves-
tigated both sides of transects and within forest stands
to find the best tracks available. The combination of
footprint (size, presence/absence of toe prints) and trail
(gait, distance between jumps, and dragging of the feet)
characteristics were used to identify all tracks (Murie
1975; Rezendes 1992; Halfpenny et al. 1995). I record-
ed the location of tracks as both linear distance along
the transect and universal transverse mercator.
Autocorrelation is often present in ecological data
and cannot be completely avoided (Proulx and O’Do-
herty 2006). It may occur during analyses of track
survey data because of uncertainty whether one or
more animals has made the tracks being counted. It is
difficult to confirm that a series of tracks along a tran-
sect belongs to a single animal (de Vos 195 1) as home

2014

ranges overlap (Badry et al. 1997; Weir 2003), and win-
ter dispersal movements are known to occur (Arthur
et al. 1993). Because of rugged environmental condi-
tions, I did not follow tracks that crossed close together
to learn whether the same animal made them. On the
basis of track characteristics, Proulx (2006, 2011) de-
duced that two different animals could be as close as
100 m along the same transect. To minimize spatial
autocorrelation, a minimum spacing of tracks and a
minimum spacing of transects were used (Proulx and
O*Doherty 2006). Only tracks > 100 m apart within
the same forest stand were recorded as two inde-
pendent tracks. Tracks < 100 m apart but in two dif-
ferent stand types were also recorded as two inde-
pendent tracks (Bowman and Robitaille 1997; Proulx
2006).

The Silvacom Group (Edmonton, Alberta) produced
1:15 000 scale vector maps for data analyses, using
Saskatchewan Forest Inventory Vegetation datasets
(Saskatchewan Environment 2004*). Nearly half the
total length of transects surveyed in 2009, 2011, and
2012 was in treed bogs and fens. Independent of their
age, these habitats varied considerably in structure ac-
cording to many factors, such as landscape position,
origin, slope gradient, water table location, drainage,
etc. (Smith ef a/. 2007). Habitat classes were, there-
fore, based on the vegetation composition (= 60%
Black Spruce stands, > 60% Tamarack stands, pure
and mixed Jack Pine stands with less than 60% Black
Spruce or Tamarack, and deciduous or mixed-conif-
erous-deciduous stands); crown closure (percentage of
ground area covered by the vertical projection of the
crown to the ground, 0-20%, 21-40%, or > 40%); and
tree height (0-10 m, 11—20 m, or > 20 m) in the forest
stands (Saskatchewan Environment 2014%*). Crown
closure values for deciduous stands refers to summer
datasets: I considered crown closure to be 0% in win-
ter. Crown closure values for coniferous stands were
representative of stand conditions throughout the year.
Winter crown closure data for mixed wood stands was
corrected according to the proportion of deciduous
species.

The proportion of habitat types traversed by survey
transects were used to determine the expected frequen-
cy of tracks per habitat type if tracks were distributed
randomly with respect to habitat types (Proulx and
O’ Doherty 2006; Proulx 2011). Habitat use (i.e., ob-
served versus expected frequency of track intercepts)
was tested using x? statistics with Yates correction
(Siegel 1956; Zar 1999). When 7? analyses suggested
an overall significant difference between observed and
expected frequencies, further comparisons were con-
ducted for each habitat class using the G test for cor-
related proportions (Sokal and Rohlf 1981). Probability
values < 0.05 were considered statistically significant.

Transects crossed major habitat types: Black Spruce-
dominated stands, 23.2%; Tamarack-dominated stands,
22.6%: other coniferous stands, 6.6%; deciduous

NOTES

273

stands, 18.4%; mixed wood stands, 22.9%; and open
areas, 6.1%. A total of 78 Fisher tracks were recorded
over 60 300 m of transects. There was a significant dif-
ference between observed and expected frequencies
of tracks per habitat class (y? = 18.7, df 9, P < 0.05)
(Figure 1). No Fisher tracks were encountered in open
areas, 1.e., cut blocks, burns, roads, and pipelines.
Tracks were significantly less frequent than expected
in Tamarack stands with > 40% crown closure (G =
3.9, P< 0.05), which consisted mainly of 0-10 m tall
trees. Fishers used other muskeg and coniferous, mixed
wood and deciduous forest stands as available.

Fishers avoided open areas and Tamarack stands
with > 40% crown cover and small trees because
these stands do not have a well-developed understory
(McLaughlan er a/. 2010; G.P., unpublished). They
offer relatively little ground structural complexity for
prey and provide Fishers with little protection against
weather and predators (G. P., unpublished). This find-
ing is in agreement with previous studies in various
ecozones (Proulx 2006; Lancaster et a/. 2008; Weir
and Corbould 2010).

In the Boreal Plains Ecozone, late-winter Fisher
habitat may correspond to any forested area that pro-
vides suitable prey, as suggested for other ecozones by
Strickland et a/. (1982) and Arthur et a/. (1989). In
Mistik’s FMA area, large forested areas dominated by
Trembling Aspen were being used by Fishers, as was

25
0 Observed
2 @ Expected
S 20
£
be
o
a
a 645
-
—
°
=
Ss 10
£
=
Zz.
5 |
0 q
@eaAxz G 4 t& 2
% Ge, %, % be, 4, 44,
a A ~ %, > 3°
% 2 Y yyy % & &@
% Ge, ) t,t ip S %, %, &
a he %,* Ya % % . 2
!
% % % A
2 t 2%,% 4%%
© 2, % % Ge &
>% > %” % > ?
»
Habitat Class
Figure 1. Observed and expected distribution of Fisher

tracks (n = 78) by habitat class (including percent-
age crown closure [cc] and range of tree heights) in
late-winter 2009, 2011, and 2012, Mistik’s Forest
Management Agreement area, Saskatchewan.

274

found in the Prairie Ecozone (Badry ef al. 1997). In
these ecozones, the presence of dense coniferous cov-
er during winter may not be necessary when there is
a well-developed understory. Fishers were found in
deciduous stands and in muskeg where a lack of crown
cover was compensated for by well-developed under-
growth (McLaughlan e¢ a/. 2010; G.P., unpublished),
which may provide Fishers with homoeothermic ad-
vantages (Buskirk ef al. 1989; Badry et al. 1997) and
access to prey (Powell and Zielinski 1994). In treed
bogs, Fishers investigated shrub thickets and the bases
of trees, and their tracks were often associated with
Snowshoe Hare (Lepus americanus Erxleben, 1777)
trails (G.P., unpublished). Thus, this study suggests
that conserving mosaics of muskeg interspersed with
forest stands of diverse composition and structure
would meet the late-winter habitat needs of Fishers.

Acknowledgements

This study was funded by Mistik Management Ltd.
I thank Al Balisky, Kevin Gillis, and Roger Nesdoly of
Mistik Management for their logistic support through-
out the study and Neil MacKenzie and Kim Stang of
Alpha Wildlife for field assistance. I am grateful to John
Peters and Ryan Spooner from the Silvacom Group
for the development of maps. I thank associate editor
David Nagorsen and two anonymous reviewers for
their comments on an earlier version of the manu-
script.

Documents Cited (marked * in text)

Government of Canada. 2014. Historical climate data. Gov-
ernment of Canada, Ottawa, Ontario, Canada. Accessed
May 7, 2012 http://climate.weather.gc.ca/

Koback, L. 2014. Saskatchewan wild fur harvest and cash
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port, Government of Saskatchewan, Regina, Saskatchewan,
Canada.

Munro, M. 2012. Last winter was third-warmest in decades:
Environment Canada. Postmedia News, May 7, 2012.
Accessed May 7, 2012. www.canada.com/technology/Last
+winter+third+warmest+decades+Environment+Cana-
da/6581378/story.html

Saskatchewan Environment 2004. Saskatchewan forest
vegetation inventory. Version 4.0. Forest Service, Saskat-
chewan Environment, Prince Albert, Saskatchewan, Cana-
da.

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NOTES py

Nn

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Received 12 February 2014
Accepted 9 April 2014

A Tribute to Warren Baxter Ballard, 1947—2012

FRANCIS R. COOK

Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada; email: freook(@ripnet.com

Cook, Francis R. 2014. A tribute to Warren Baxter Ballard, 1947-2012. Canadian Field-Naturalist 128(3): 276-288.

Warren Baxter Ballard, professor and Bricker Chair
in Wildlife Management, Department of Natural Re-
sources Management, Texas Tech University, Lubbock,
Texas, lost a battle with pancreatic cancer on 12 January
2012,

He had gained broad research experience in wildlife
biology in the boreal forest of Alaska in the northwest
and New Brunswick in the east and very different habi-
tats in the southwestern United States. In addition, he
had been to New Zealand, Poland, Sweden, and Russia.
His extensive publication record spanning 1971 to 2013
included 27 contributions to The Canadian Field-Nat-
uralist covering a variety of vertebrate taxa, including
ducks, turkeys, deer, caribou, foxes, wolves, and bears
and usually coauthored with students and colleagues.
His editorial contributions to journals and other pub-
lications were also extensive. As associate editor of The
Canadian Field-Naturalist for mammals 1994-2001,
2007-2011, and before and between these periods as an
occasional reviewer, he contributed over 280 individual
reviews himself or assigned them to graduate students
as part of their training.

Warren was born on 28 April 1947, in Boston, Mas-
sachusetts. His family moved to Albuquerque, New
Mexico, in the early 1950s and Warren attended St.
Pius X High School there and, subsequently, enrolled
at New Mexico State University. He held numerous
summer positions with the New Mexico Department of
Game & Fish, the United States Army Corps of Engi-
neers, the Bureau of Land Management, and the De-
partment of Animal, Range and Wildlife Sciences, New
Mexico State University. He obtained a bachelor of sci-
ence degree in fish and wildlife management in 1969
and a master’s in environmental biology from Kansas
State University, Manhattan, Kansas, in 1971. His the-
sis for the latter. was entitled, “Importance of social ag-
gression in booming ground hierarchy of greater prairie
chickens (Zympanuchus cupido pinnatus)”; his super-
visor was Dr. Robert J. Robel.

From | August 1971 to 31 July 1973, Warren was a
wildlife biologist with the U.S. Fish and Wildlife Serv-
ice, Tulsa, Oklahoma. From | August 1973 to | Novem-
ber 1990, he was a research biologist with the Alaska
Department of Fish and Game, in Juneau, Homer, Glen-
nallen, Palmer, Anchorage, and Nome, Alaska, but re-
signed when his application for educational leave was
rejected. From 11 January 1990 to 1 January 1992, he

?

\

FiGure |: Warren Baxter Ballard, about 2004, at Texas Tech
University. Photographer Artie Limmer.

was a research associate at the School of Renewable
Natural Resources, University of Arizona, Tucson He
received a Ph.D. in Wildlife Science, University of
Arizona, in 1993; his dissertation was “Demographics,
movements, and predation rates of wolves in northwest
Alaska” (supervisor: Dr. Paul R. Krausman),

Briefly, 1-10 January 1992, he was a wildlife re-
search biologist with the U.S. National Park Service.
Anchorage, Alaska. From 10 January 1992 to | July
1993, he was a senior wildlife ecologist with LGL
Alaska Research Associates, Inc., Anchorage, Alaska.
This was followed by a position as director and asso-
ciate professor in the Cooperative Fish and Wildlife Re-
search Unit, University of New Brunswick, Fredericton.
New Brunswick, where he stayed until August 1996.

VALS

2014

£2

Moving southwest, he spent | September 1996 to |
February 1998 working as research supervisor, Ari-
zona Game and Fish Department, Phoenix, Arizona. In
a final move, from 7 January 1998 to 31 August 2003,
he was an associate professor in the Department of
Natural Resources Management at Texas Tech Univer-
sity, where he became associate chair on 2 July 2002
and was elevated to professor on | September 2003.
On 13 April 2006, he was appointed to the prestigious
Bricker Chair in Wildlife Management, a position he
held until his death. In addition, he held appointments
as adjunct professor, Faculty of Biology, University of
New Brunswick (9 January 1993) adjunct associate
professor, School of Renewable Natural Resources,
University of Arizona, Tucson (1 October 1996), an
adjunct to faculty, Division of Biology, Kansas State
University, Manhattan (1 September 1998).

A highly regarded teacher, Warren taught seven
courses over 39 terms achieving a 4.5 out of 5 rating in
student response. He supervised 44 graduate students
(30 at the M.Sc. level and 14 Ph.D.s) as well as two
postdoctoral students.

A prolific writer and supervisor, Warren wrote or co-
wrote an ever-increasing multitude of journal papers,
monographs, contributions to books and conference
proceedings, and miscellaneous publications, as well as
invited presentations and progress reports which were
not published.

Cook: A TRIBUTE TO WARREN BAXTER BALLARD, 1947-2012

je Tok x > 2 : 7 _
ea hee 7 BD > aE :

eT

FIGURE 2. Warren Baxter Ballard and Heather Whitlaw on a visit to editor’s residence in 2002. Photograph by Francis Cook.

His editorial contributions were numerous: in addi-
tion to his long service as associate editor of The Cana-
dian Field-Naturalist mentioned above, he served as
editor in-chief of The Wildlife Society Bulletin (2002—
2005). He was co-editor of A/ces (1998-2001); The
Moose Call, the newsletter of the North American
Moose Conference and Workshop, Canada (1994—
1996); and the Third International Moose Sympo-
sium, Soviet Union (1991—1992). He was consulting
editor for the University of Alberta Press (1992) and
associate editor for Wildlife Monographs (2007, 2004,
1995-1996, and 1989), The Wildlife Society Bulletin
(1998-2001), and Alces (1996-1997). He carried out
peer reviews for the Canadian Journal of Zoology, the
Journal of Mammalogy, the Journal of Wildlife Dis-
eases, The Journal of Wildlife Management, Alces, the
Proceedings of Desert Bighorn Sheep Council, the Pro-
ceedings of the Second International Wolf Symposium,
International Bear Conference, Ecoscience, the Journal
of Range Management, Texas Tech University Press,
Biological Conservation, Science of the Total Environ-
ment, the Texas Journal of Science, and the European
Journal of Wildlife Research.

Diverse awards have recognized Warren’s many
contributions: Distinguished Moose Biologist Award
from the North American Moose Conference at St.
John’s, Newfoundland (1989); an Award for Excellence
from the School of Renewable Natural Resources, Uni-

278

THE CANADIAN FIELD-NATURALIST

Vol. 128

FIGURE 3. ne en Baxter Ballard on a ae trip in Tees October 2003. Eroierapl courtesy Heather Whitelaw.

versity of Arizona, for an outstanding dissertation
(1994); the Order of Alces at the 31st North Ameri-
can Moose Conference for service to Alces (1995); a
Special Service Award from the University of New
Brunswick for contributions to the New Brunswick
Cooperative Fish and Wildlife Research Unit from
1993 to 1996 (1996); admission to Sigma Xi honorary
research fraternity (1999); the Big 12 Faculty Fellow-
ship from Kansas State University (1999); a Teacher of
the Year award from the Range, Wildlife, and Fisheries
Club, Texas Tech University (1999); an Outstanding
Researcher Award from the College of Agricultural
Sciences and Natural Resources, Texas Tech Univer-
sity (2002); Best Article for 2002 from The Wildlife
Society in Bismarck, North Dakota, for his paper enti-
tled ““Deer—predator relationships: a review of recent
North American studies with emphasis on mule and
black-tailed deer” (2002); a Special Service Recogni-
tion Award from The Wildlife Society, Bismarck, North
Dakota (2002); a Chancellors Council Distinguished
Research Award from Texas Tech University (2002):
the life-time title of TWS Fellow from The Wildlife
Society, Madison, Wisconsin (2005); an Outstanding
Achievement Award from the Texas Chapter of the
Wildlife Society, Beaumont, Texas (2006); and a Best
Monograph award from The Wildlife Society for a
manuscript entitled “Pathogens, nutritional deficien-

cy, and climate influences on a declining moose pop-
ulation” authored by Dennis L. Murray, Eric W. Cox,
Warren B. Ballard, Heather A. Whitlaw, Mark S.
Lenarz, Thomas W. Custer, Terri Barnett, and Todd K.
Fuller, Tucson, Arizona (2007).

Warren was exceptionally active in the science com-
munity and professional societies. He held profes-
sional membership in The Wildlife Society, the Texas
chapter of The Wildlife Society, the American Society
of Mammalogists, the Canadian Field-Naturalist, Sig-
ma Xi, and the Southwestern Association of Natural-
ists. He held numerous positions on committees and
served as a consultant to the University of Minneso-
ta, the British Columbia provincial government, LGL
Alaska Research Associates, Inc., Entrix Inc.. and
Revett Silver Mine concerning grizzly bear issues.

Warren had definite views on wildlife research and
education, which he put into practice in his teaching,
editing, and society involvement, as well as his own re-
search. In his curriculum vitae, he stated,

The purpose of wildlife research is to test hypotheses,
develop better techniques, and build a sound database for
the management and conservation of wildlife through-
out the world. Wildlife research should alw ays use and
follow the scientific method. Research should be orient-
ed both towards testing theory and solvi ing technical
problems. Dissemination of research results through

2014

publication is essential to a successful and productive
research program.

Education in wildlife and biological sciences
should provide students with basic concepts and
theories, and equip them with the necessary tools
of the trade to be productive biologists. Knowl-
edge of ecological concepts, effective communi-
cation skills, statistical expertise, and critical
thinking should be stressed at all levels of college
education. In addition, recent and current research,
as well as management findings and problems
should be incorporated into the education process.

On Warren’s death, Texas Tech president, Guy Bai-
ley, wrote, “Dr. Ballard was a Texas Tech Horn Pro-
fessor and the Bricker Chair in Wildlife Management.
A Horn professorship is the highest honor a faculty
member can receive from the university. Horn profes-
sors are a testament to the quality of our academics
because they represent the very best of our faculty.”

“His legacy lives on in the students, faculty and
research projects he touched,” said Michael Galyean,
interim dean of the College of Agricultural Sciences
and Natural Resources.

“Warren was my friend since graduate school, an
internationally-recognized research scientist, a major
figure in this wildlife program, and an irreplaceable
part of our department,” said Mark Wallace, chairman
of Tech’s Department of Natural Resources Manage-
ment.

On 18 January 2012, Canadian colleague, Graham
Forbes, posted a note on a “tribute wall” (Lake Ridge
Chapel and Memorial Designers postings): “Warren
was very well-liked here at the University of New
Brunswick during his time as director of the Wildlife
Coop Unit. He initiated some great projects and was
valued by all who worked with him. We missed him
when he left UNB and it’s hard to acknowledge he has
passed. I presently occupy his old office and just had
a large shot of whisky, in his memory, and his honour.
Our thoughts are with Heather.”

Dwayne Sabine also posted a message in January:
“T was Warren’s first grad student. I led one of two deer
studies he initiated here in New Brunswick, Canada,
and he hired Heather to run the other.... Warren was
certainly a positive influence on my career and life. I
have fond memories of time spent in the field and at
camp with Warren and Heather. He always had great
stories at hand, and (unfortunately, perhaps) I may have
given him one or two more during his time here (the
blindfolded deer that ‘got away’ comes to mind...).”

Graham Forbes recalled an additional anecdote from
Warren’s time at the University of New Brunswick
(personal communication 18 February 2013): “Warren
became ‘known’ to campus security one autumn when
he took his Wildlife Investigation Techniques class on
to the rugby field on the edge of campus for some drug
;mmobilization training. The class was target practising

Cook: A TRIBUTE TO WARREN BAXTER BALLARD, 1947-2012

2719

with some Cap-Chur rifles (CO,-powered for tran-
quilizing projectiles); in hindsight, it would have been
prudent to inform security first!”

Personally, | will miss our many exchanges on man-
uscript evaluation and on life over 30 years. Our last
conversation was shortly before he died — a compar-
ison of our success with walkers. Warren, who had
worked efficiently and productively with little inter-
ruption over several health setbacks, was ever upbeat
and proud of his dexterity with his walker and encour-
aged me to improve with mine.

Warren was predeceased by his father, Warren Bax-
ter Ballard. He was survived by his mother LaVerne
Rosemary Ballard (née Bernat) and his wife, Heather
Whitlaw, as well as children Cynthia Bergamo (hus-
band Greg Bergamo), Warren B. Ballard, III (mother
Ms. Sheila Sturtz), Laurina Wittig (husband Thomas
Wittig), and Raymond Ballard (mother Ms. Tina Cun-
ning); grandchildren Ezra Bergamo, and Blair and
Brandon Ballard; mother-in-law, Nan McGhee, father-
in-law, David Whitlaw, sister-in-law, Patricia Whitlaw;
and nieces, Elizabeth and Paige Jones. He is fondly
remembered by graduate students from the several uni-
versities where he had been a staff member as well as
by research colleagues and friends from around the
world.

Acknowledgements

Warren’s wife and frequent coauthor, Heather Whit-
law, provided Warren’s 2007 comprehensive curricu-
lum vite, an updated 2012 list of Warren’s publications,
and a photograph. Roger Applegate searched for the
full citation for many incomplete references to publi-
cations that appeared after Warren’s death. At Texas
Tech University, Kristina Woods Butler, Research and
Academic Communications, Office of the Vice Presi-
dent for Research, and Norman Martin, College of
Agricultural Sciences and Natural Resources, traced
the source of Figure | and approved its credit and use,
respectively. The notice in The Lubbock Avalanche-
Journal, 13 January 2012, and online comments post-
ed on the Internet guest book of Lake Ridge Chapel
and Memorial Designers, 6025 82nd Street, Lubbock,
Texas were drawn upon for additional information.
Also accessed was the article by Roger D. Applegate
and Paul R. Krausman. 2012. Warren B. Ballard, Jr.
1947-2012: A personal tribute. The Wildlife Society
36(1): 4-5. Roger Applegate, Graham J. Forbes, and
Heather Whitlaw read earlier drafts of this article and
made many appreciated corrections and additions.

Bibliography

Publications (in chronological order)

Ballard, W. B. 1971. Importance of social aggression in
booming ground hierarchy of greater prairie chickens (7\m-
panuchus cupido pinnatus). M.Sc. Thesis, Kansas State
University, Manhattan, Kansas, USA. 90 pages.

Ballard, W. B., and D. Bowen. 1971. Capture methods for
upland plovers. Bird Banding.

Davis, C. A., D. Babb, and W. B. Ballard. 1973. Food habits
of mourning doves in the Mesilla Valley of south-central
New Mexico, New Mexico State University Agricultural
Experiment Station. 8 pages.

Robel, R. J., and W. B. Ballard. 1974. Lek social organiza-
tion and reproductive success in the greater prairie chick-
en. American Zoologist 14: 121—128.

Ballard, W. B., and R. J. Robel. 1974. Reproductive impor-
tance of dominant male greater prairie chickens. Auk 91:
75-85.

Ballard, W. B. 1978. Status and management of the moun-
tain goat in Alaska. Pages 15—23 in Proceedings of Ist Inter-
national Mountain Goat Symposium, Kalispell, Montana,
February 19, 1977. Edited by W. Samuel and W. G. Mac-
Gregor. Fish and Wildlife Branch.

Taylor, K. P., and W. B. Ballard. 1979. Moose movements
and habitat use along the Susitna River near Devil’s Can-
yon, Alaska. Proceedings of North American Moose Con-
ference and Workshop 15: 169-186.

Ballard, W. B., H. Merriam, and P. Coppock. 1979. Evalu-
ation of methods utilized to estimate deer harvest in Alas-
ka. Pages 196-206 in Sitka Black-tailed Deer: Proceedings
of Conference in Juneau, Alaska. Edited by J. O. C. Wallmo
and W. Schoen, Department of Agriculture, Forest Service,
Alaska Region.

Ballard, W. B., A. W. Franzmann, K. P. Taylor, T. Spraker,
C. C. Schwartz, and R. O. Peterson. 1979. Comparison
of techniques utilized to determine moose calf mortality in
Alaska. Proceedings of North American Moose Conference
and Workshop 15: 362-387.

Ballard, W. B. 1980. Brown bear kills gray wolf. Canadian
Field-Naturalist 94: 91.

Ballard, W. B., C. L. Gardner, and S. D. Miller. 1980.
Influence of predators on summer movements of moose
in south-central Alaska. Proceedings of North American
Moose Conference and Workshop 16: 333-359.

Franzmann, A. W., W. B. Ballard, C. C. Schwartz, and T.
H. Spraker. 1980. Physiologic and morphometric meas-
urements in neonatal moose and their cows in Alaska.
Proceedings of North American Moose Conference and
Workshop 16: 362-387.

Ballard, W. B., and R. W. Tobey. 1981. Decreased calf pro-
duction of moose immobilized with anectine administered
from helicopter. Wildlife Society Bulletin 9: 207-209.

Ballard, W. B., T. H. Spraker, and K. P. Taylor. 1981.
Causes of neonatal moose calf mortality in southcentral
Alaska. Journal of Wildlife Management 45: 335-342,

Ballard, W. B., C. L. Gardner, J. H. Westlund, and S. M.
Miller. 1981. Use of mandible versus longbone to evalu-
ate percent marrow fat in moose and caribou. Alces 17:
147-164.

Miller, S. D., and W. B. Ballard. 1982. Homing of transplant-
ed Alaskan brown bears. Journal of Wildlife Management
46: 869-876.

Ballard, W. B., S. D. Miller, and T. H. Spraker. 1982. Home
range, daily movements, and reproductive biology of brown
bear in southcentral Alaska. Canadian Field-Naturalist 96:
1-5.

Peterson, R. O., C. C. Schwartz, and W. B. Ballard. 1982.
Tooth eruption as an index to population quality in moose.
Journal of Wildlife Management 47: 884-888.

Eide, S., and W. B. Ballard. 1982. Apparent case of surplus
killing of caribou by gray wolves. Canadian Field-Natu-
ralist 96: 87-88.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Miller, S. D., and W. B. Ballard. 1982. Density and biomass
estimates for an interior Alaskan brown bear, Ursus arctos,
population. Canadian Field-Naturalist 96: 448-454.

Ballard, W. B., and R. O. Stephenson. 1982. Wolf contro]
— take some and leave some. Alces 18: 726-300.

Ballard, W. B., A. W. Franzmann, and C. L. Gardner. 1982.
Comparison and assessment of drugs used to immobilize
Alaskan gray wolves (Canis lupus) and wolverine (Gulo
gulo) from a helicopter. Journal of Wildlife Diseases 18:
339-342.

Ballard, W. B. 1982. Gray wolf-brown bear relationships in
the Nelchina Basin of southcentral Alaska. Pages 71—80 in
Wolves of the World. Edited by F. H. Harrington and P. C.
Paquet. Noyes Publications, Park Ridge, New Jersey, USA.
474 pages.

Ballard, W. B., and J. Dau. 1983. Characteristics of gray
wolf, Canis lupus, den and rendezvous sites in southcentral
Alaska. Canadian Field-Naturalist 97: 299-302.

Ballard, W. B., R. Farnell, and R. O. Stephenson. 1983.
Long distance movement by gray wolves. Canadian Field-
Naturalist 97: 333.

Franzmann, A. W., C. C. Schwartz, D. C. Johnson, J. B.
Faro, and W. B. Ballard. 1984. Immobilization of moose
with carfentanil. Alces 20: 259-281.

Tobey, R., and W. B. Ballard. 1985. Increased mortality in
gray wolves captured with acepromazine and etorphine
hydrochloride in combination. Journal of Wildlife Diseases
21: 188-190.

Goodwin, E. A., and W. B. Ballard. 1985. Use of tooth
cementum for age determination of gray wolves. Journal
of Wildlife Management 49: 313-316.

Foster, J. W., and W. B. Ballard. 1985. Radio-telemetry as
an aid in the study of wolf den site behavior. Annual Pro-
ceedings of the American Association of Zoo Veterinarians.
2 pages.

Whitman, J. S., W. B. Ballard, and C. L. Gardner. 1986.
Home range and habitat use by wolverines in southcentral
Alaska. Journal of Wildlife Management 50: 460-463.

Gardner, C. L., W. B. Ballard, and J. Jessup. 1986. Long
distance movement of a wolverine. Journal of Mammalogy
67: 603.

Ballard, W. B., S. M. Miller, and J. S. Whitman. 1986.
Modelling a southcentral Alaskan moose population. Alces
22: 201-244.

Aumiller, L., and W. B. Ballard. 1986. Documented range
extension of the mountain goat, Oreamnos americanus, in
Alaska. Canadian Field-Naturalist 100: 560.

Zarnke, R. L., and W. B. Ballard. 1987. Serologic survey for
selected microbial pathogens of w ay es in Alaska, 1975-
1982. Journal of Wildlife Diseases 23: 77-85.

Miller, S. D., E. F. Becker, and W. B. Ballard. 1987. Densi-
ty estimates using modified capture-recapture techniques
for black and brown bear populations in Alaska. Interna-
tional Conference on Bear Research and Management 7:
23-35.

Ballard, W. B., and J. S. Whitman. 1987. Marrow fat dynam-
ics in moose calves. Journal of Wildlife Management 51:
66-69.

Ballard, W. B., and D. G. Larsen. 1987. Implications of pred-
ator—prey relationships to moose management. Swedish
Wildlife, Research Supplement 1: 581—602.

Ballard, W. B., J. S. Whitman, and N. G. Tankersley. 1987.
Impact mechanisms of hydroelectric dev elopment projects
on moose in North America. Swedish Wildlife, Research
Supplement 1: 737—740.

2014

Ballard, W. B., J. S. Whitman, and C. L. Gardner. 1987,
Ecology of an exploited wolf population in south-central
Alaska. Wildlife Monograph 98. 54 pages.

Bergerud, A. T., and W. B. Ballard. 1988. Wolf predation on
caribou: the Nelchina herd case history, a different inter-
pretation. Journal of Wildlife Management 52: 344-357.

Ballard, W. B., A. F. Cunning, and J. S. Whitman. 1988.
Hypotheses of impacts on moose due to hydroelectric proj-
ects. Alces 24: 34-47.

Ballard, W. B., and A. T. Bergerud. 1988. Wolf population
fluctuations in the Nelchina Basin of south-central Alaska:
are revisions of previous estimates justified? 3rd North
American Caribou Workshop, Fairbanks, Alaska. 2 pages.

Bergerud, A. T., and W. B. Ballard. 1989. Wolf predation on
Nelchina caribou: a reply. Journal of Wildlife Management
53: 251-259.

Taylor, Jr., W. P., H. G. Reynolds IIL, and W. B. Ballard.
1989. Immobilization of grizzly bears with tiletamine

hydrochloride and zolazepan hydrochloride. Journal of

Wildlife Management 53: 978-981.
Roney, K. E., L. A. Ayres, and W. B. Ballard. 1989. Count-

ing grizzly bears in northwest Alaska. U.S. Department of

the Interior Park Science 9: 22—24.

Ballard, W. B., K. E. Roney, L. A. Ayres, and D. N. Larsen.
1990. Estimating grizzly bear density in relation to devel-
opment and exploitation in northwest Alaska. Internation-
al Conference on Bear Research and Management 8: 405—
413.

Ballard, W. B., S. D. Miller, and J. S. Whitman. 1990.
Brown and black bear predation on moose in southcentral
Alaska. Alces 26: 1-8. Also published in Global Trends in
Wildlife Management, | 8th International Union of Game
Biologists Congress, Jagiellonian University, Krakow,
Poland. August, 1987, pages 177—184.

Ballard, W. B., and S. D. Miller. 1990. Effects of reducing
brown bear density on moose calf survival in southcentral
Alaska. Alces 26: 9-13. Also published in Global Trends
in Wildlife Management, 18th International Union of Game
Biologists Congress, Jagiellonian University, Krakow,
Poland, August 1987. Pages 171-176.

Ballard, W. B., L. A. Ayres, K. E. Roney, and T. H. Spraker.
1991. Immobilization of free-ranging gray wolves with a
mixture of tiletamine hydrochloride and zolazepan hydro-
chloride. Journal of Wildlife Management 55: 71-74.

Ballard, W. B., J. S. Whitman, and D. J. Reed. 1991. Popu-
lation dynamics of moose in south-central Alaska. Wildlife
Monograph 114. 49 pages.

Ballard, W. B., L. A. Ayres, C. L. Gardner, and J. A. Foster.
1991. Den site activity patterns of gray wolves, Canis lupus,
in southcentral Alaska. Canadian Field-Naturalist 106: 497-
504.

Ballard, W. B. 1992. Bear predation on moose: a review of

recent North American studies and their implications. Alces
28, Supplement 1: 162-176.

Miller, S. M., and W. B. Ballard. 1992. Analysis of an effort
to increase moose calf survivorship by increased hunting
of brown bears in south-central Alaska. Wildlife Society
Bulletin 20: 445-454.

Ballard, W. B. 1993. Modeled impacts of bear and wolf

predation on moose. Alces 28: 79-88.

Ballard, W. B. 1993. Demographics, movements, and pre-
dation rates of wolves in northwest Alaska. Ph.D. thesis,
University of Arizona, Tucson. 374 pages.

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281]

Franzmann, A. W., and W. B. Ballard. 1993. Use of phys-
ical and physiological indices for monitoring moose pop-
ulation status — a review. Alces 29; 125—134.

Ballard, W. B., L. A. Ayres, D. J. Reed, S. G. Fancy, and
K. E. Roney. 1993. Demography of Noatak grizzly bears
in relation to hunting and mining development in north-
western Alaska. U. S. National Park Service Monograph
23. 112 pages.

Ballard, W. B. 1994. Effects of black bear predation on cari-
bou — a review. Alces 30: 25-36.

Van Ballenberghe, V., and W. B. Ballard. 1994. Limita-
tion and regulation of moose populations: the role of pre-
dation. Canadian Journal of Zoology 72: 2071-2077.

Forbes, G., P. Chamberland, E. Daigle, and W. B. Ballard.
1994. The lack of problem bear issues in Fundy National
Park. Proceedings of 12th Eastern Black Bear Workshop,
Knoxville, Tennessee, IBA Publications, Knoxville, Ten-
nessee, USA.

Stephenson, R. O., W. B. Ballard, C. A. Smith, and K.
Richardson. 1995. Wolf biology and management in Alas-
ka, 1981-1991. Pages 43-54, in Ecology and Conservation
of Wolves in a Changing World. Edited by L. N. Carbyn,
S. H. Fritts, and D. R. Seip, Canadian Circumpolar Insti-
tute, University of Alberta, Edmonton, Alberta, Canada.

Fancy, S. G., and W. B. Ballard. 1995. Monitoring wolf ac-
tivity by satellite. Pages 329-334, in Ecology and Conser-
vation of Wolves in a Changing World. Edited by L. N.
Carbyn, S. H. Fritts, and D. R. Seip, Canadian Circumpolar
Institute, University of Alberta, Edmonton, Alberta, Cana-
da.

Brand, C. J., M. J. Pybus, W. B. Ballard, and R. O. Peter-
son. 1995. The role of infectious and parasitic diseases in
population dynamics of the gray wolf (Canis lupus). Pages
419-430, in Ecology and Conservation of Wolves in a
Changing World. Edited by L. N. Carbyn, S. H. Fritts, and
D. R. Seip, Canadian Circumpolar Institute, University of
Alberta, Edmonton, Alberta, Canada.

Ballard, W. B. 1995. Public attitudes and wildlife science.
Northeast Wildlife 51: 63-70.

Ballard, W. B. 1995. Bone marrow fat as an indicator of un-
gulate condition — how good is it? Alces 31: 105-109.
Ballard, W. B., G. M. Matson, and P. R. Krausman. 1995.
Comparison of two methods to age gray wolf (Canis lupus)
teeth. Pages 455-460, in Ecology and Conservation of
Wolves in a Changing World. Edited by L. N. Carbyn, S.
H. Fritts, and D. R. Seip, Canadian Circumpolar Institute,

University of Alberta, Edmonton, Alberta, Canada.

Ballard, W. B., D. J. Reed, S. G. Fancy, L. A. Ayres, and
P. R. Krausman. 1995. Accuracy, precision, and per-
formance of satellite radio collars for monitoring wolf
movements. Pages 461-468, in Ecology and Conservation
of Wolves in a Changing World. Edited by L. N. Carbyn,
S. H. Fritts, and D. R. Seip, Canadian Circumpolar Insti-
tute, University of Alberta, Edmonton, Alberta, Canada.

Ballard, W. B., C. L. Gardner, and D. J. Reed. 1995. Use
of line-intercept transects for estimating wolf densities.
Pages 469-480, in Ecology and Conservation of Wolves in
a Changing World. Edited by L. N. Carbyn, S. H. Fritts,
and D. R. Seip, Canadian Circumpolar Institute, University
of Alberta, Edmonton, Alberta, Canada.

Ballard, W. B., S. G. Fancy, P. R. Krausman, and W. P.
Burger. 1995. Potential applications of satellite telemetry
to wildlife studies in remote locations. Pages 566-569 in
Integrating People and Wildlife for a Sustainable Future:
Proceedings of the first International Wildlife Manage-

ment Congress. Edited by J. A. Bissonette and P. R. Kraus-
man, The Wildlife Society, Bethesda, Maryland.

Ballard, W. B., and M. A. Cronin. 1995. Arctic Alaska cari-
bou herds in relation to oil field development. Proceedings
of 6th Symposium on Impacted Wildlife, Thorne Ecolog-
ical Institute, Glenwood Springs, Colorado, USA. Pages
35-46.

Lohr, C., W. B. Ballard, and A. Bath. 1996. Attitudes toward
gray wolf (Canis /upus) reintroduction to New Brunswick.
Wildlife Society Bulletin 24: 414-420.

Lohr, C., and W. B. Ballard. 1996. Historical occurrence of
wolves, Canis lupus, in the Maritime Provinces. Canadian
Field-Naturalist 110: 607-610.

Pollard, R., W. B. Ballard, M. Cronin, and J. Bryan. 1996.
Parasitic insect abundance and microclimate of gravel pads
and tundra within the Prudhoe Bay Oil Field, Alaska, in
relation to use by caribou, Rangifer tarandus granti. Cana-
dian Field-Naturalist 110: 649-658.

Pollard, R. H., W. B. Ballard, L. E. Noel, and M. A. Cronin.
1996. Summer distribution of caribou, Rangifer tarandus
granti, in the area of the Prudhoe Bay Oil Field, 1990—
1994. Canadian Field-Naturalist 110: 659-674.

Balsom, S. E., W. B. Ballard, and H. A. Whitlaw. 1996.
Mature coniferous forest as critical moose habitat. Alces
32: 131-140.

Whitlaw, H. A., M. W. Lankester, and W. B. Ballard.
1996. Parelaphostrongylus tenuis in terrestrial gastropods
from white-tailed deer winter and summer ranges in north-
ern New Brunswick. Alces 32: 75-83.

Sabine, D. L., A. H. Boer, and W. B. Ballard. 1996. Impacts
of habitat fragmentation on pairing success of male Oven-
birds, Seiurus aurocapillus, in southern New Brunswick.
Canadian Field-Naturalist 110: 688-693.

Ballard, W. B., P. J. MacQuarrie, A. W. Franzmann, and
P. R. Krausman. 1996. Effects of winters on physical con-
dition of moose in Alaska. Alces 32: 51—59.

Ballard, W. B., and P. R. Krausman. 1997. Occurrence of
rabies in Alaskan wolves. Journal of Wildlife Diseases 33:
242-245.

Cronin, M. A., B. J. Pierson, S. R. Johnson, L. E. Noel,
and W. B. Ballard. 1997. Caribou population density in
the Prudhoe Bay Region of Alaska. Journal of Wildlife Re-
search 2: 59-68.

Banfield, E., and W. B. Ballard. 1997. Are corridors justi-
fied as a wildlife management tool? Journal of Wildlife
Research 2: 182-185.

Cunningham, S. C., R. Engel-Wilson, P. Smith, and W.
B. Ballard. 1997. Food habits and nesting characteristics
of sympatric mourning and white-winged doves in south-
central Arizona. Journal of Wildlife Research 2: 242-253,

Ballard, W. B., L. A. Ayres, P. R. Krausman, D. J. Reed,
and S. G. Fancy. 1997. Ecology of wolves in relation to
a migratory caribou herd in northwest Alaska. Wildlife
Monograph 135. 47 pages.

Miller, S. D., G. C. White, R. A. Sellers, H. V. Reynolds,
J. W. Schoen, K. Titus, V. G. Barnes Jr., R. B. Smith,
R .R. Nelson, W. B. Ballard, and C. C. Schwartz. 1997.
Brown and black bear density estimation in Alaska using
radio-telemetry and replicated mark-resight techniques.
Wildlife Monograph 133. 55 pages.

Ballard, W. B., H. A. Whitlaw, D. L. Sabine, R. A. Jenkins,
S. J. Young, and G. F. Forbes. 1998. White-tailed deer.
Odocoileus virginianus, capture techniques in yarding and
non-yarding populations in New Brunswick. Canadian
Field-Naturalist 112: 254-261.

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Vol. 128

Spaulding, R. L., P. R. Krausman and W. B. Ballard. 1998.
Summer diet of gray wolves, Canis /upus, in northwest-
ern Alaska. Canadian Field-Naturalist 112: 262-266.

Cronin, M. A., S. C. Amstrup, G. M. Durner, L. E. Noel,
and W. B. Ballard. 1998. Caribou distribution during the
post-calving period in relation to infrastructure in the Prud-
hoe Bay oil field, Alaska. Arctic 51: 85—93.

Cronin, M. A., W. B. Ballard, J. D. Bryan, B. J. Pierson,
and J. D. McKendrick. 1998. Northern Alaska oil fields
and caribou: a commentary. Biological Conservation 83:
195-208.

Ballard, W. B., and V. Van Ballenberghe. 1998. Moose
predator relationships: research and management needs.
Alces 34: 91-105.

Whitlaw, H. A., W. B. Ballard, D. L. Sabine, S. J. Young,
R. A. Jenkins, and G. F. Forbes. 1998. Survival and cause-
specific mortality rates of adult white-tailed deer in New
Brunswick. Journal of Wildlife Management 62: 1335-
1341.

Noel, L. E., R. H. Pollard, W. B. Ballard, and M. A.
Cronin. 1998. Activity and use of active gravel pads and
tundra by caribou, Rangifer tarandus granti, within the
Prudhoe Bay oil field, Alaska. Canadian Field-Naturalist
112: 400-409.

Gipson, P. S., and W. B. Ballard. 1998. Accounts of famous
North American wolves, Canis /upus. Canadian Field-
Naturalist 112: 724-739.

Gipson, P. S., W. B. Ballard, and R. M. Nowak. 1998. Fa-
mous damaging wolves and the credibility of early wildlife
literature. Wildlife Society Bulletin 26: 806-816.

Van Ballenberghe, V., and W. B. Ballard. 1998. Popula-
tion dynamics. Pages 223-245 in Ecology and Manage-
ment of the North American Moose. Edited hy A. W.
Franzmann and C. C. Schwartz, Wildlife Management
Institute, Washington, D. C., USA (Awarded best book
by The Wildlife Society)

Ballard, W. B., and V. Van Ballenberghe. 1998. Predator—
prey relationships. Pages 247-273 in Ecology and Man-
agement of the North American Moose. Edited by A. W.
Franzmann and C. C. Schwartz, Wildlife Management
Institute, Washington, D. C. (Awarded best book by The
Wildlife Society)

Ballard, W. B., S. S. Rosenstock, and J. C. deVos, Jr. 1998.
The effects of artificial water developments on ungulates
and large carnivores in the Southwest. Pages 64-105 in
Proceedings of Symposium on Environment, Economics.
and Legal Issues Related to Rangeland Water Develop-
ments, Arizona State University, College of Law, Tempe,
Arizona, USA.

deVos, Jr., J. C., W. B. Ballard, and S. S. Rosenstock. 1998.
Research design considerations to evaluate efficacy of
wildlife water developments. Pages 606-612 in Proceed-
ings of Symposium on Environment, Economics, and Legal
Issues Related to Rangeland Water Developments, Arizona
State University, College of Law, Tempe, Arizona, USA.

Cunningham, S. C., C. R. Gutavson, L. A. Haynes, and W.
B. Ballard. 1999. Diet selection of mountain lions in south-
east Arizona. Journal of Range Management 52: 202-207.

Spaulding, R. L., P. R. Krausman, and W. B. Ballard. 1999.
Observer bias in scat analyses. Journal of Wildlife Re-
search 2: 128-132.

Rosenstock, S. S., W. B. Ballard, and J. C. deVos, Jr. 1999.
Viewpoint: benefits and impacts of wildlife water devel-
opments. Journal of Range Management 52: 302-311.

2014

Ballard, W. B., H. A. Whitlaw, S. J. Young, R. A. Jenkins,
and G. F. Forbes. 1999. Predation and survival of white-
tailed deer fawns in northcentral New Brunswick. Journal
of Wildlife Management 63: 574-579.

Ballard, W. B., M. Edwards, S. G. Fancy, S. Boe, D. J.
Reed, and P. R. Krausman. 1999, comparison of VHF
and satellite telemetry for estimating wolf territory sizes in
northwest Alaska. Wildlife Society Bulletin 26; 823-829,

Forbes, G., D. Johnson, E. Daigle, P. Chamberland, and
W. Ballard. 1999. Ecological magnets and black bears in
the Greater Fundy Ecosystem. Pages 57-68 in Black Bear
Ecology and Management Issues for Atlantic Canadian
National Parks: Proceedings of Parks Canada Atlantic Re-
gion Black Bear Workshop. Edited by G. Forbes and E.
Daigles, Parks Canada, Ottawa, Ontario, Canada.

Forbes, G., P. Chamberland, E. Daigle, and W. Ballard.
1999. The lack of problem bear issues in Fundy National
Park. Pages 89-94 in Black Bear Ecology and Manage-
ment Issues for Atlantic Canadian National Parks: Pro-
ceedings of Parks Canada Atlantic Region Black Bear
Workshop. Edited by G. Forbes and E. Daigles, Parks Cana-
da, Ottawa, Ontario, Canada.

Daigle, E., G. Forbes, W. Emrich, and W. Ballard. 1999.
Black bear den characteristics at forest stand and site scales,
New Brunswick. Pages 95-103 in Black Bear Ecology and
Management Issues for Atlantic Canadian National Parks:
Proceedings of Parks Canada Atlantic Region Black Bear
Workshop. Edited by G. Forbes and E. Daigles, Parks
Canada, Ottawa, Ontario, Canada.

Rupp, S. P., W. B. Ballard, and M. C. Wallace. 2000. A
nationwide evaluation of deer hunter harvest survey tech-
niques. Wildlife Society Bulletin 28: 570-578.

Connor, K. J., W. B. Ballard, T. Dilworth, S. Mahoney,
and D. Anions. 2000. Changes in structure of a boreal
forest community following intense herbivory by moose.
Alces 36: 111—132.

Gipson, P. S., W. B. Ballard, R. M. Nowak, and L. D.
Mech. 2000. Accuracy and precision of estimating age of
gray wolves by tooth wear. Journal of Wildlife Management
64: 752-758.

Kamler, J. F., K. D. Seyffert, and W. B. Ballard. 2000.
Range expansion of the Eurasian Collared-dove in the
Texas Panhandle. Texas Ornithological Society Bulletin
33: 23-24.

Ballard, W. B., H. A. Whitlaw, B. F. Wakeling, R. L.
Brown, J. C. devos Jr., and M. C. Wallace. 2000. Sur-
vival of female elk in northern Arizona. Journal of
Wildlife Management 64: 500-504.

Ballard, W. B., P. R. Krausman, S. Boe, S. Cunningham,
and H. A.Whitlaw. 2000. Short-term response of gray
wolves, Canis lupis, to wildfire in northwestern Alaska.
Canadian Field-Naturalist 114: 241-247.

Kamler, J. F., W. B. Ballard, and K. Mote. 2000. Aggres-
sive behavior exhibited by a swift fox, Vulpes velox.
Canadian Field-Naturalist 114: 506.

Spaulding, R. L., P. R. Krausman, and W. B. Ballard.
2000. Methodology for calculating biomass consumed
by gray wolves. Wildlife Society Bulletin 28: 947-950.

Cronin, M. A., H. A. Whitlaw, and W. B. Ballard. 2000.
Northern Alaska oil fields and caribou. Wildlife Society
Bulletin 28: 919-922.

Ballard, W. B., M. A. Cronin, R. Rodrigues, RAO
Skoog, and R. H. Pollard. 2000. Arctic f0x, Alopex
lagopus, den densities in the Prudhoe Bay oil field, Alas-
ka. Canadian Field-Naturalist 114: 453-456.

Cook: A TRIBUTE TO WARREN BAXTER BALLARD, 1947-2012

Ballard, W. B., M. A. Cronin, M. D. Robards, and E. H.
Follman, 2000. Body sizes, ages, reproductive status,
and sex ratios of arctic foxes, Alopex lagopus, in the
Prudhoe Bay oil field, Alaska. Canadian Field-Naturalist
114: 493-494.

Ballard, W. B., M. A. Cronin, and H. A. Whitlaw. 2000.
Interactions between caribou and oil fields: the Central
Arctic Herd Caribou Case History. Pages 85-104 in The
Natural History of an Arctic Oil Field Development and
the Biota. Edited by J. C. Truett and S. R. Johnson, Aca-
demic Press, San Diego, California, USA.

Cronin, M. A., H. A. Whitlaw, and W. B. Ballard. 2001.
Addendum to Northern Alaska oil fields and caribou.
Wildlife Society Bulletin 29: 764.

Ballard, W. B., E. H. Follman, D. G. Ritter, M. D.
Robards, and M. A. Cronin. 2001. Rabies and canine
distemper in an arctic fox population in Alaska. Journal
of Wildlife Diseases 37: 133-137.

Cunningham, S. C., W. B. Ballard, and H. A. Whitlaw.
2001. Survival and cause-specific mortality rates of a
heavily exploited mountain lion population in southeast-
ern Arizona. Southwestern Naturalist 46: 76-80.

Sabine, D., W. B. Ballard, J. Bowman, G. Forbes, and H.
A. Whitlaw. 2001. Use of mixedwood stands by winter-
ing white-tailed deer in eastern Canada. Forestry Chroni-
cle 77: 97-103.

Ballard, W. B., D. Lutz, T. W. Keegan, L. H. Carpenter,
and J. C. deVos Jr. 2001. Deer-predator relationships: a
review of recent North American studies with emphasis
on mule and black-tailed deer. Wildlife Society Bulletin
29: 99-115.

Kamler, J. F., W. B. Ballard, and D. A. Swepston. 2001.
Range expansion of mule deer in the Texas Panhandle.
Southwestern Naturalist 46: 378-379.

Ballard, W. B. 2001. Deer-predator relationships. Pages
52-69 in The role of predator control as a tool in game
management: Proceedings of a symposium, 18-19 April
2001, Kerrville, Texas. Edited by T. F. Ginnett and S. E.
Henke, Texas Agricultural Research & Extension Center,
San Angelo, Texas, USA.

Kamler, J. F., and W. B. Ballard. 2002. Distribution and
taxonomy of red foxes in the western United States. The
Wildlife Society Bulletin 30: 370-379.

Ballard, W. B., J. C. deVos Jr., J. F. Kamler, and H. A.
Whitlaw. 2002. A need for an integrated radiotemetry
database. The Wildlife Society Bulletin 30: 263-264.

Sabine, D., S. F. Morrison, H. A. Whitlaw, W. B. Ballard,
G. Forbes, and J. Bowman. 2002. Migration behavior of
white-tailed deer under varying winter climate regimes in
New Brunswick. Journal of Wildlife Management 66: 718—
728.

Bowman, J., M. C. Wallace, W. B. Ballard, J. H. Brunjes,
M. S. Miller, and J. M. Hellman. 2002. Evaluation of
two techniques for attaching radio transmitters to turkey
poults. Journal of Field Ornithology 73: 276-280.

Spears, B. L., W. B. Ballard, M. C. Wallace, R. S. Phillips,
D. H. Holdstock, J. H. Brunjes, M. Miller, R. D.
Applegate, P. S. Gipson, and T. Barnett. 2002. Retention
times of miniature radiotransmitters glued to wild turkey
poults. The Wildlife Society Bulletin 30: 861-867.

Kamler, J. F., R. M. Lee, J. C. deVos, Jr., W. B. Ballard,
and H. A. Whitlaw. 2002. Survival and cougar predation
of translocated bighorn sheep in Arizona. Journal of Wild-
life Management 66: 1267-1272.

Marshal, J. P., P. R. Krausman, V. C. Bleich, W. B. Ballard,
and J. S. McKeever. 2002. Rainfall, El Nino, and dynam-
ics of mule deer in the Sonoran Desert, California. Journal
of Wildlife Management 66: 1283-1289.

Kamler, J. F., W. B. Ballard, R. L. Gilliland, and K. Mote.
2002. Improved trapping methods for swift foxes and sym-
patric coyotes. The Wildlife Society Bulletin 30: 1262
1266.

Kamler, J. F., W. B. Ballard, B. Bullock, D. Butler, and R.
Ward. 2003. General remains of great horned owl pellets

from Union County, New Mexico. New Mexico Journal of

Science 43: 68—70.

Kamler, J. F., W. B. Ballard, R. L. Gilliland, P. R. Lemons
II, and K. Mote. 2003. Impacts of coyotes on swift foxes
in northwestern Texas. Journal of Wildlife Management
67: 317-323.

Kamler, J. F., W. B. Ballard, R. L. Gilliland, and K. Mote.
2003. Spatial relationships between swift foxes and coy-
otes in northwestern Texas. Canadian Journal of Zoology
81: 168-172.

Kamler, J. F., R. M. Lee, J. C. deVos, Jr., W. B. Ballard,
and H. A. Whitlaw. 2003. Mortalities from climbing acci-
dents of translocated bighorn sheep in Arizona. The South-
western Naturalist: 48: 145-147.

Kamler, J. F., L. A. Green, and W. B. Ballard. 2003. Recent
occurrence of black bears in the southwestern Great Plains.
The Southwestern Naturalist 48: 303-306.

Ballard, W. B., M. A. Cronin, M. D. Robards, and W. A.
Stubblefield. 2003. Heavy metal concentrations of arctic
foxes, Alopex lagopus, in the Prudhoe Bay oil field, Alas-
ka. Canadian Field-Naturalist 117: 119-121.

Kamler, J. F., W. B. Ballard, B. R. Helliker, and S. Stiver.
2003. Range expansion of raccoons in the western Utah
and eastern Nevada. Western North American Naturalist 63:
406-408.

Kamler, J. F., and W. B. Ballard. 2003. Range expansion of
red foxes in eastern Nevada and western Utah. Journal of
Arizona-Nevada Academy of Science 36: 18-20.

Kamler, J. F., W. B. Ballard, E. B. Fish, P. R. Lemons, K.
Mote, and C. C. Perchellet. 2003. Habitat use, home
ranges, and survival of swift foxes in a fragmented land-
scape: conservation implications. Journal of Mammalogy
84: 989-995.

Avey, J. T., W. B. Ballard, M. C. Wallace, M. H. Humphrey,
P. R. Krausman, F. Harwell, and E. B. Fish. 2003. Habi-
tat relationships between sympatric mule and white-tailed
deer in south-central Texas. The Southwestern Naturalist
48: 644-653.

Cunningham, S. C., W. B. Ballard, L. M. Monroe, M. J.
Rabe, and K. D. Bristow. 2003. Black ear habitat use in
burned and unburned areas, central Arizona. Wildlife Soci-
ety Bulletin 31: 786-792.

Carrera, R., and W. B. Ballard. 2003. Elk distribution in
Mexico: a critical review. Wildlife Society Bulletin 31:
1272-1276.

Rodriguez, M. C., P. R. Krausman, W. B. Ballard, C.
Villalobos, and W. W. Shaw. 2003. Attitudes of Mexican
citizens about wolf translocation in Mexico. Wildlife Soci-
ety Bulletin 31: 971-979.

Spears, B. L., W. B. Ballard, M. C. Wallace, R. G. Apple-
gate, and P. S. Gipson. 2003. Coyote, Canis latrans — Rio
Grande Turkey, Meleagris gallopavo intermedia, interac-
tions. Canadian Field-Naturalist 117: 645-647.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Cronin, M. A., S. P. Haskell, and W. B. Ballard. 2003. The
frequency of antlerless female caribou and reindeer in
Alaska. Rangifer 23: 67-70.

Spears, B. L., W. J. Peterson, and W. B. Ballard. 2003. Bone
marrow fat content of moose from northeastern Minneso-
ta, 1972-2000. Alces 39: 273-285.

Ballard, W. B., D. Lutz, T. W. Keegan, L. H. Carpenter,
and J. C. deVos, Jr. 2003. Deer—predator relationships.
Pages 177-218 in Mule Deer Conservation: Issues and
Management Strategies. Edited by J. C. deVos, M. R.
Conover, and N. E. Headrick, Jack H. Berryman Institute
Press, Utah State University, Logan, Utah, USA.

Ballard, W. B., L. N. Carbyn, and D. W. Smith. 2003. Wolf
interactions with non-prey. Pages 259-271 in Ecology and
Conservation of the Wolf. Edited by L. D. Mech and L.
Boitani, University of Chicago Press, Chicago, Illinois.
USA. (Awarded best book by The Wildlife Society).

Avey, J. T., W. B. Ballard, M. C. Wallace, M. H Humphrey,
P. R. Krausman, E. B. Fish, and P. J. Zwank. 2003.
Determining habitat variability between sympatric deer
species in west-central Texas. Proceedings Western States
and Provinces Deer and Elk Workshop.

Lemons, P. R., W. B. Ballard, R. M. Sullivan, and M. A.
Sovada. 2003. Den site activity patterns of adult male and
female swift foxes, Vulpes velox, in northwestern Texas.
Canadian Field-Naturalist 117: 424429.

Kamler, J. F., and W. B. Ballard. 2003. White color phase
of the swift fox, Vulpes velox. Canadian Field-Naturalist
117: 468-469.

McGee, B. K., and W. B. Ballard. 2004. Observations of a
bald eagle (Haliaeetus leucocephalus) feeding on prairie
dogs (Cynomys ludovicianus) in the northwest Texas pan-
handle. Bulletin Texas Ornithological Society 37: 14.

Kamler, J. F., and W. B. Ballard. 2004. First record of a
long-tailed weasel from the Texas Panhandle. The Prairie
Naturalist 32: 141-142.

Kamler, J. F., W. B. Ballard, K. Mote, and R. L. Gilliland.
2004. Coyote (Canis latrans) movements relative to cattle
Bos taurus) carcass areas. Western North American Nat-
uralist 64: 53-58.

Kamler, J. F., W. B. Ballard, P. R. Lemons, and K. Mote.
2004. Variation in mating system and group structure in two
populations of swift foxes. Animal Behavior 68: 83-88.

Kamler, J. F., W. B. Ballard, E. M. Gese, R. L. Harrison, S.
Karkis, and K. Mote. 2004. Adult male emigration and
a female-based social organization in swift foxes. Vulpes
velox. Animal Behavior 67: 699-702.

Rominger, E. M., H. A. Whitlaw, D. L. Weybright, W. C.
Dunn, and W. B. Ballard. 2004. The influence of mountain
lion predation on bighorn sheep translocations. Journal of
Wildlife Management 68: 993-999.

Lawrence, R. K., S. Demarais, R. A. Relyea, S. P. Haskell,
W. B. Ballard, and T. L. Clark. 2004. Desert mule deer
survival in southwest Texas. Journal of Wildlife Manage-
ment 68: 561—569.

Reed, J. E., R . J. Baker, W. B. Ballard, and B. T. Kelly.
2004. Differentiating Mexican gray wolf and coyote scats
using DNA analysis. Wildlife Society Bulletin 31: 685—
692.

Cunningham, S. C., and W. B. Ballard. 2004. Effects of a
castrophic wildfire on black bear demographics in central
Arizona. Wildlife Society Bulletin 31: 928-937.

Pence, D. B., J. Kamler, and W. Ballard. 2004. Ectoparasites
of the swift fox in northwestern Texas. Journal of Wildlife
Diseases 40: 543-547.

2014

Kamler, J. F., W. B. Ballard, and P. S. Gipson. 2004. Occur-
rence of feral dogs in northwest Texas: an observation.
Texas Journal of Agriculture and Natural Resources 15:
75-78.

Haskell, S. P., and W. B. Ballard. 2004. Factors limiting
productivity of the Central Arctic Caribou Herd of Alaska.
Rangifer 24: 71-78.

Spears, B. L., W. B. Ballard, M. C. Wallace, R. S. Phillips,
D. H. Holdstock, J. H. Brunjes, M. Miller, R. D. Apple-
gate, and P. S. Gipson. 2005. Survival of Rio Grande wild
turkey chicks. Journal of Field Ornithology 76: 12-20.

Kamler J. F., W. B. Ballard, P. L. Lemons, R. L. Gilliland,
and K. Mote. 2005. Home range and habitat use of coy-
otes in an area of native prairie, farmland, and CRP fields.
American Midland Naturalist 153: 396-404.

Kamler, J. F., W. B. Ballard, R. L. Harrison, and C. G.
Schmitt. 2005. Range expansion of red foxes in northwest-
ern Texas and northeastern New Mexico. Southwestern
Naturalist 501: 100-101.

Kamler, J. F., W. B. Ballard, E. M. Gese, R. L. Harrison,
and S. M. Karki. 2005. Dispersal characteristics of swift
foxes. Canadian Journal of Zoology 82: 1-6.

Butler, Matthew J., Andrew P. Teaschner, Warren B. Bal-
lard, and Brady K. McGee. 2005. Commentary: Wildlife
Ranching in North America — arguments, issues, and per-
spectives. Wildlife Society Bulletin 33: 381-389.

Livingston, T. R., P. S. Gipson, W. B. Ballard, D. M.
Sanchez, and P. R. Krausmann. 2005. Scat removal: a
source of bias in feces related studies. Wildlife Society
Bulletin 33: 172-178.

Butler, M. J., M. C. Wallace, W. B. Ballard, and S. DeMaso.
2005. From the field: the relationship of Rio Grande wild
turkey distributions to roads. Wildlife Society Bulletin 33:
745-748.

O’Brien, C. S., H. M. Boyd, P. R. Krausman, W. B. Bal-
lard, S. C. Cunningham, and J. C. deVos, Jr. 2005. Influ-
ence of wildfire and coyote presence on habitat use by col-
lared peccaries. Wildlife Society Bulletin 33: 865-875.

McGee, B. K., M. J. Butler, M. C. Wallace, W. B. Ballard,
and K. L. Nicholson. 2005. From the Field: a comparison
of survey techniques for swift fox pups. Wildlife Society
Bulletin 33: 1169-1173.

Nicholson, K. L., B. K. McGee, and W. B. Ballard. 2005.
Swift fox, Vulpes velox, den located next to a railroad track
in northwestern Texas. Canadian Field-Naturalist 119: 584—
585.

Kamler, J. F., and W. B. Ballard. 2006. Ear flashing behav-
ior of black-tailed jackrabbits (Lepus californicus). Amer-
ican Midland Naturalist 155: 390-391.

Cunningham, S. C., L. Kirkendall, and W. B. Ballard.
2006. Gray fox and coyote abundance and diet responses
after a wildfire in central Arizona. Western North American
Naturalist 66: 169-180.

McGee, B. K., K. L. Nicholson, W. B. Ballard, and M. J.
Butler. 2006. Characteristics of swift fox dens in northwest
Texas. Western North American Naturalist 66: 239-245.

Brown, A. D., J. H. Brunjes, R. S. Phillips, W. B. Ballard,
M. C. Wallace, and R. J. Baker. 2006. Eggshell remains
as a noninvasive source of genetic material in wild turkeys
(Meleagris gallopavo). Texas Tech University, National
Science Research Laboratory, Lubbock, Texas, USA. Occa-
sional paper 257. round ee.

Kamler, J. K., and W. B. Ballard. 2006. Canid diversity in
the Texas Panhandle. The Southwestern Naturalist 51:

569-571.

Cook: A TRIBUTE TO WARREN BAXTER BALLARD, 1947-2012

Haskell, S. P., R. M. Nielson, W. B. Ballard, M. C. Cronin,
and T. L. MeDonald. 2006. Dynamic responses of calving
caribou to oilfields in northern Alaska. Arctic 59: 179-190.

McGee, B. K., M. J. Butler, D. B. Pence, J. L. Alexander,
J. B. Nissen, W. B. Ballard, and K. L. Nicholson. 2006.
Possible vector dissemination by swift foxes following a
plague epizootic in black-tailed prairie dogs in northwest
Texas. Journal of Wildlife Diseases 42: 415-420.

Marshal, J. P., P. R. Krausman, V. C. Bleich, S. S. Rosen-
stock, and W. B. Ballard. 2006. Gradients of forage bio-
mass and ungulate use near wildlife water developments.
Wildlife Society Bulletin 34: 620-626.

McGee, B. K., W. B. Ballard, K. L. Nicholson, B. L.
Cyper, P. R. Lemons I, and J. F. Kamler. 2006. Effects
of artificial escape dens on swift fox populations in north-
west Texas. Wildlife Society Bulletin 34: 821-827.

Reed, J. E., W. B. Ballard, P. S. Gipson, B. T. Kelly, P. R.
Krausman, M. C. Wallace, and D. B. Wester. 2006. Diets
of free-ranging Mexican gray wolves in Arizona and New
Mexico. Wildlife Society Bulletin 34: 1127-1133.

Brunjes, K. J., W. B. Ballard, M. H. Humphrey, F. Harwell,
N. E. McIntyre, P. R. Krausman, and M. C. Wallace.
2006. Habitat use by sympatric mule and white-tailed deer
in Texas. Journal of Wildlife Management 70: 1351-1359.

Holdstock, D. P., M. C. Wallace, W. B. Ballard, J. H. Brun-
jes, R. S. Phillips, B. L. Spears, S. J. DeMaso, J. D.
Jernigan, R. D. Applegate, and P. S. Gipson. 2006. Male
Rio Grande turkey survival and movements in the Texas
Panhandle and southwestern Kansas. Journal of Wildlife
Management 70: 1028-1036.

Nicholson, K. L., W. B. Ballard, B. K. McGee, J. Surles, P.
Lemons, and J. F. Kamler. 2006. Swift fox use of black-
tailed prairie dog towns in northwest Texas. Journal of
Wildlife Management 70: 1659-1666.

Butler, D. A., W. B. Ballard, S. P. Haskell, and M. C. Wal-
lace. 2006. From the field: limitations of thermal infrared
imaging for locating neonatal deer in semi-arid shrub com-
munities. Wildlife Society Bulletin: 34: 1458-1462.

McKinney, T., J. C. deVos Jr., W. B. Ballard, and S. R. Boe.
2006. Mountain lion predation of translocated desert big-
horn sheep in Arizona. Wildlife Society Bulletin: 34: 1255—
1263.

McGee, B. K., K. L. Nicholson, W. B. Ballard, and M. J.
Butler. 2006. Characteristics of swift fox dens in northwest
Texas. Western North American Naturalist 66: 239-245.

Butler, M. J., W. B. Ballard, and H. A. Whitlaw. 2006.
Physical characteristics, hematology, and serum chemistry
of free-ranging gray wolves, Canis /upus, in southcentral
Alaska. Canadian Field-Naturalist 120: 205-212.

Murray, D. L., E. W. Cox, W. B. Ballard, H. A. Whitlaw,
M. S. Lenarz, T. W. Custer, T. K. Fuller, and T. Barnett.
2006. Pathogens, nutritional deficiency, and climate influ-
ences on a declining moose population. Wildlife Mono-
graphs 166. 30 pages.

Huffman, R. T., M. C. Wallace, W. B. Ballard, G. Hall,
R. Houchin, R. D.Applegate, S J. DeMaso, and P. S.
Gipson. 2006. Nesting habitat of Rio Grande wild turkeys.
Pages 103-111 in Managing Wildlife in the Southwest.
Edited by J. W. Cain, Ill, and P. R. Krausman, Southwest
Section of The Wildlife Society, Tucson, Arizona, USA.

Butler, M. J., W. B. Ballard, M. C. Wallace, S. J. Demaso,
and R. D. Applegate. 2006. Comparing techniques for
counting Rio Grande wild turkeys at winter roosts. Pages
112-117 in Managing Wildlife in the Southwest. Edited

286

by J. W. Cain, I, and P. R. Krausman, Southwest Section
of The Wildlife Society, Tucson, Arizona, USA.

Spears, B., K. Nicholson, R. Huffman, W. B. Ballard, M.
C. Wallace, R. Applegate, and P. Gipson. 2006. Ecology
of Rio Grande wild turkeys in southwest Kansas. Kansas
Wildlife Bulletin 5. Kansas Wildlife and Parks Department,
Pratt, Kansas, USA.

O’Brien, C. S., H. M. Boyd, P. R. Krausman, W. B. Bal-
lard, R. M. Kattnig, S. C. Cunningham, and J. C. deVos,
Jr. 2006. Nutritional mule deer forage in burned and un-
burned interior chaparral. Pages 31-48 in Managing Wild-
life in the Southwest. Edited by J. W. Cain III, and P. R.
Krausman, Southwest Section of The Wildlife Society.
Tucson, Arizona, USA.

Spears, B. L., M. C. Wallace, W. B. Ballard, R. S. Phillips,
D. H. Holdstock, J. H. Brunjes, M. Miller, R. D. Apple-
gate, and P. S. Gipson. 2007. Habitat use and survival of
pre-flight wild turkey broods. Journal of Wildlife Manage-
ment 71: 69-81.

Nicholson, K. L., W. B. Ballard, B. K. McGee, and H. A.
Whitlaw. 2007. Dispersal and extra-territorial movements
of swift foxes (Vulpes velox) in northwestern Texas. West-
ern North American Naturalist 67: 102-108.

Haskell, S. P., and W. Ballard. 2007. Accounting for radio-
telemetry signal flux in triangulation point estimation.
European Journal of Wildlife Research 53: 204-211.

Haskell, S. P., and W. B. Ballard. 2007. Modeling the West-
ern Arctic Caribou Herd during a positive growth phase:
effects of radio-collars and wolves. Journal of Wildlife
Management 71:619—627.

Kamler, J. F., W. B. Ballard, M. C. Wallace, R. L. Gilliland,
and P. S. Gipson. 2007. Dietary overlap of swift foxes and
coyotes in northwestern Texas. American Midland Natu-
ralist: 158: 140-147.

Butler, M. J., W. B. Ballard, M. C. Wallace, S. J. DeMaso,
and B. K. McGee. 2007. Aerial surveys for estimating wild
turkey abundance in the Texas Rolling Plains. Journal of
Wildlife Management 71: 1639-1645.

Butler, M. J., W. B. Ballard, M. C. Wallace, and S. J.
DeMaso. 2007. Road-based surveys for estimating wild
turkey density in the Texas Rolling Plains. Journal of Wild-
life Management 71: 1646-1653.

Haskell, S. P., W. B. Ballard, D. A. Butler, N. M. Tatman,
M. C. Wallace, C. Kochanny, and P. Aleumbrac. 2007.
Observations on capturing and aging deer fawns. Journal
of Mammalogy 88: 1482-1487.

Butler, M. J., W. B. Ballard, M. C. Wallace, and S. J. Dema-
so. 2007. Wild turkey (Meleagris gallopavo) detectability
from helicopters and ramifications for estimating abun-
dance. European Journal of Wildlife Research 54: 148-152.

Hall, G. I., M. C. Wallace, W. B. Ballard, D. C. Ruthven
II, M. J. Butler, D. P. Holdstock, R. L. Houchin, R. T.
Huffman, R. S. Phillips, B. L. Spears, and R. Apple-
gate. 2007. Rio Grande wild turkey habitat selection in the
southern Great Plains. Journal of Wildlife Management
71: 2583-2591.

Rice, M. B., W. B. Ballard, E. B. Fish, and D. Holdermann.
2007. Predicting private landowner support toward recol-
onizing Black Bears in the Trans-Pecos Region of Texas.
Human Dimensions of Wildlife 12: 405-415.

Haskell, S. P., W. B. Ballard, D. A. Butler, M. C. Wallace,
T. R. Stephenson, O. Alcumbrac, and M. H. Humphrey.
2007. Factors affecting birth dates of sympatric deer in
west-central Texas. Journal of Mammalogy 89: 448-458.

THE CANADIAN FIELD-NATURALIST

Vol. 128

McGee, B. K., W. B. Ballard, and K. L. Nicholson. 2007.
Swift fox, Vulpes velox, den use patterns in northwestern
Texas. Canadian Field-Naturalist 121: 71—75.

Holdstock, D. P., M. C. Wallace, W. B. Ballard, J. H.
Brunjes, R. S. Phillips, B. L. Spears, S. J. Demaso, J.
D. Jernigan, R. D. Applegate, and P. S. Gipson. 2007.
Male Rio Grande wild turkey habitat characteristics in the
Texas Panhandle and Southwestern Kansas. Ninth Wild
Turkey Symposium: 217-230.

Butler, M. J., G. I. Hall, M. C. Wallace, W. B. Ballard, R.
S. Phillips, J. H. Brunjes, R. Huffman, R. Houchin, J.
Bullock, S. J. DeMasso, R. D. Applegate, and M. Frisbie.
2007. Utility of poult-hen counts to index productivity of
Rio Grande wild turkeys. Ninth Wild Turkey Symposium:
159-168.

Brunjes, J. H., W. B. Ballard, M. C. Wallace, R. S. Phillips,
D. H. Holdstock, B. L. Spears, M. J. Butler, M. S.
Miller, N. E. McIntyre, S. J. DeMasso, R. Applegate,
and P.S. Gibson. 2007. Patterns of capture related mortal-
ity in Rio Grande wild turkeys. Proceedings of the Nation-
al Wild Turkey Symposium 9: 75-81.

Phillips, R. S., W. B. Ballard, M. C. Wallace, D. P. Hold-
stock, B. L. Spears, M. S. Miller, J. H. Brunjes, and S.
J. Demaso. 2007. Movement, fidelity and dispersal of Rio
Grande wild turkeys in the Texas Panhandle. Ninth Wild
Turkey Symposium: 149-158.

Hall, G. I., M. J. Butler, M. C. Wallace, W. B. Ballard, D.
P. Holdstock, R. L. Houchin, R. T. Huffman, R. S.
Phillips, B. Spears, D. C. Ruthven, and R. Applegate.
2007. Rio Grande wild turkey home range characteristics
in the southern Great Plains. Proceedings of Annual Con-
ference of Southeast Association of Fish and Wildlife Agen-
cies: 60: 36-42.

Ranft, R., P. Gipson, W. Ballard, M. Wallace, D. Wester, J.
Bonner, R. Huffman, and G. Hall. 2007. Coyote diet in
the Texas Panhandle and southwestern Kansas. Proceed-
ings of the North American Prairie Conference 20: 255—
268.

Mills, B. H., B. D. Vogt, M. L. Sumner, and W. B. Ballard.
2008. Diets of feral emus in the Cross Timbers and Prairies
Region of Texas. Southwest Naturalist 53: 385-388.

Carrera, R., J. E. Reed, W. B. Ballard, P. Gipson, B. Kelly,
P. Krausman, and M. Wallace. 2008. Comparison of coy-
ote and Mexican wolf diets in the Blue Range Wolf Recov-
ery Area, Arizona and New Mexico. Journal of Wildlife
Management 72: 376-381.

Cariappa, C., W. Ballard, S. Breck, A. Piaggio, and M.
Newbaum. 2008. Estimating population size of Mexican
wolves noninvasively. Ecological Restoration 26: 14-16.

Haskell, S. P., and W. B. Ballard. 2008. Annual re-habituation
of calving caribou to oilfields in northern Alaska: impli-
cations for expanding development. Canadian Journal of
Zoology 86: 627-637.

McRoberts, J. T., M. J. Butler, W. B. Ballard, M. C. Wal-
lace, H. A. Whitlaw and D. A. Haukos. 2008. Aerial sur-
veys for prairie grouse: detectability, disturbance response,
and distance sampling. Proceedings of Annual Conference
of Southeast Association of Fish and Wildlife Agencies
62: 203. [Abstract]. ;

Nicholson, K. L., W. J. Peterson, and W. B. Ballard. 2008.
Comparisons and trends in white-tailed deer. Odocoileus
virginianus, in northeastern Minnesota, 1974-1990. Cana-
dian Field-Naturalist 122: 253-261.

McRoberts, J. T., W. B. Ballard, M. J. Butler, H. A. Whit-
law, K. Boydston, and A. H. Swift. 2009. Lesser prairie

2014

chicken conservation and wind development in the Texas
Panhandle: a case for proactive wildlife management. Pages
74-76 in Proceedings of the National Wind Coordinating
Collaborative Wind Wildlife Research Meeting VII. Octo-
ber 28-29, 2008, Milwaukee, Wisconsin, USA. Edited by
S. Savitt Schwartz, National Wind Coordinating Collab-
orative, Washington, D. C., USA,

Brunjes, K. J., W. B. Ballard, M. H. Humphrey, F. Harwell,
N. E. MeIntyre, P. R. Krausman, and M. C. Wallace.
2009. Home range size and overlap of male sympatric mule
and white-tailed deer in Texas. Western North American
Naturalist 69: 125-130.

Brunjes, Kristina J., Warren B. Ballard, Mary H. Hum-
phrey, Fieldling Harwell, Nancy E. McIntyre, Paul R.
Krausman, and Mark C. Wallace. 2009. Home ranges of
sympatric mule deer and white-tailed deer in Texas. South-
western Naturalist 54: 253-260.

Butler, D. J., S. P. Haskell, W. B. Ballard, M. C. Wallace,
C. J. Britton, and M. H. Humphreys. 2009. Differences
in timing of birth, birthing sites, and bedding sites in sym-
patric populations of deer. Southwestern Naturalist 54:
261-271.

Merkle, J. A., P. R. Krausman, D. W. Stark, J. K. Oakleaf,
and W. B. Ballard. 2009. Summer diet of the Mexican
gray wolf (Canis lupus baileyi). Southwestern Naturalist
54: 480-524.

Rice, M. B., W. B. Ballard, E. R. Fish, N. McIntyre, and
D. Holdermann. 2009. The importance of accurate lan-
duse/landcover maps for assessing habitat utilization by
black bear (Ursus americanus) in the Trans-Pecos region
of Texas. Wildlife Biology in Practice 4: 48-56.

Rice, M. B., W. B. Ballard, E. B. Fish, N. E. McIntyre, and
D. Holdermann. 2009. Habitat-distribution modeling of
a recolonizing black bear, Ursus americanus, population in
the Trans-Pecos region of Texas. Canadian Field-Naturalist
123: 246-254.

Tatman, N. M. J. T. McRoberts, W. A. Smith W. B. Ballard,
F. P. Kehoe, and T. G. Dilworth. 2009. Nest success and
duckling survival of greater scaup, Avthya marila, at Grassy
Island, New Brunswick. Canadian Field-Naturalist 123:
323-328.

McRoberts, J. T., W. B. Ballard, M. J. Butler, H. A. Whit-
law, K. Boydston, and A. H. Swift. 2009. Lesser prairie
chicken conservation and wind development in the Texas
panhandle: a case for proactive e wildlife management.
Pages 71—73 in Proceedings of the Wind Wildlife Research
Meeting VII. Edited by S. S. Schwartz. National Wind
Coordinating Collaborative, 28-29 October 2008, Milwau-
kee, Wisconsin, USA.

Holt, R. D., M. J. Butler, W. B. Ballard, and C. A. Kukal.
2010. Disturbance of lecking lesser prairie-chickens (7)-
panuchus pallidicitus) by ring-necked pheasants in (Pha-
sianus colchicus) Hemphill County, Texas. Western North
American Naturalist 70: 241-244.

O’Brien, C. S., P. R. Krausman, H. M. Boyd, W. B. Bal-
lard, S. C. Cunningham, and J. C. deVos, Jr. 2010. Influ-
ence of coyotes on habitat use by mule deer following a
wildfire. California Fish and Game 96: 7-22.

Haskell, S. P., D. A. Butler, W. B. Ballard, M. J. Butler,
M. C. Wallace, and M. H. Humphrey. 2010. Deer density
estimation in west-central Texas: old versus new ground
techniques with mark-resight as a comparative baseline.
Proceeding of the Western States and Provinces Deer and
Elk Workshop 7: 30-47.

Cook: A TRIBUTE TO WARREN BAXTER BAI LARD, 1947-2012

287

Erxleben, D. R., M. J. Butler, W. B. Ballard, M. C. Wallace,
J. B. Hardin, and S. J. DeMaso. 2010. Encounter rates
from road-based surveys of Rio Grande wild turkeys in
Texas. Journal of Wildlife Management 74: 1134-1140.

Butler, M. J., W. B. Ballard, R. D. Holt, and H. A. Whitlaw.
2010. Sound intensity of booming in lekking lesser prairie-
chickens. Journal of Wildlife Management 74: 1160-1162.

Ballard, W. B. 2010. Predator-prey relationships. /n Biolo-
gy and Management of White-tailed Deer. Edited by D.
Hewitt, CRC Press, Chicago, Illinois.

Mech, L. D., W. Ballard, E. Bangs, and B. Ream. 2010.
Restricting wolves risks escapes. BioScience 60: 485-486.

McBride, R. T., A. J. Giordano, and W. B. Ballard. 2010.
Diets of jaguars in the transitional Chaco of Paraguay. Bell-
bird 4: 1-4.

Haskell, S. P., W. B. Ballard, M. C. Wallace, M. H. Hum-
phrey, and D. A. Butler. 2010. Postpartum group cohe-
sion of sympatric deer species in west Texas. Journal of
Wildlife Management 74: 1686-1692.

Tatman, N. M., W. B. Ballard, M. C. Wallace, S. P. Haskell,
J. deVos, Jr., and O. J. Alcumbrac. 2010. Evaluation of
use of vaginal-implant transmitters in mule deer (Odo-
coileus hemionus). Southwestern Naturalist 56: 247-251.

Cariappa, C. A., J. K. Oakleaf, W. B. Ballard, and S. W.
Breck. 2010. A reappraisal of the evidence for regulation
of wolf populations. Journal of Wildlife Management 75:
726-730.

Giordano, A. J., and W. B. Ballard. 2010. Noteworthy record
of a black howler monkey from the central dry Chaco of
Paraguay. Neotropical Primates 17: 74-75.

Walker, R. N., M. C. Wallace, W. B. Ballard, R. S. Phillips,
S. L. McKenzie,R. M. Swearingin, and M. Janis. 2010.
Lack of cottonwood regeneration in the Southern Great
Plains: Implications for Wild Turkeys. Proceedings of the
National Wild Turkey Symposium 10: 73-78.

Swearingin, R., M. C. Wallace, W. B. Ballard, M. J. Butler,
R. S. Phillips, R. N. Walker, S. L. McKenzie, B. E.
Petersen, and D. Ruthven IIT. 2010. Chronology of Rio
Grande wild turkey flocking behavior in the Texas Rolling
Plains. Proceedings of the National Wild Turkey Sympo-
sium 10: 243-249.

Swearingin, R., M. C. Wallace, W. B. Ballard, M. J. But-
ler, R. S. Phillips, R. N. Walker, S. L. McKenzie, B. E.
Petersen, and D. Ruthven III. 2010. Winter roost char-
acteristics of Rio Grande wild turkeys in the Rolling Plains
of Texas. Proceedings of the National Wild Turkey Sym-
posium 10: 251-262.

Wallace, M. C., W. B. Ballard, D. P. Holdstock, and B.
Petersen. 2010. Rio Grande wild turkey diets in the Texas
Panhandle. Proceedings of the National Wild Turkey Sym-
posium 10: 201-211.

McKenzie, S. L., M. C. Wallace, W. B. Ballard, M. J. But-
ler, C. Ruthven, and O. Aleumbrac. 2010. Survival, be-
havior, and physical effects of surgically implanted radio-
transmitters on Rio Grande turkey poults. Proceedings of
the National Wild Turkey Symposium 10: 111-118.

Phillips, R. S., W. B. Ballard, and M. C. Wallace. 2010.
Rio Grande turkey habitat associations during dispersal.
Proceedings of the National Wild Turkey Symposium 10:
213-225.

McRoberts, J. T., M. J. Butler, W. B. Ballard, H. A.
Whitlaw, and D. A. Haukos. 2011. Detectability of lesser
prairie-chicken leks: a comparison of surveys from air-
craft. Journal of Wildlife Management 75: 771-778.

McRoberts, J. T., M. J. Butler, W. B. Ballard, Mark C.
Wallace, H. A. Whitlaw, and D. A. Haukos. 2011. Res-
ponse of lesser prairie-chickens on leks to aerial surveys.
Wildlife Society Bulletin 35: 27-31.

Erxleben, D. R., M. J. Butler, W. B. Ballard, M. C. Wallace,
M. J. Peterson, N. J. Silvy, W. P. Kuvlesky Jr., D. G.
Hewitt, S. J. DeMaso, J. B. Hardin, and M. K. Dom-
inguez-Brazil. 2011. Wild turkey (Meleagris gallopavo)
association to roads: implications for distance sampling.
European Journal of Wildlife Research 57: 57-65.

Breck, S. W., B. M. Kluever, M. Panasci, J. Oakleaf, T.
Johnson, W. Ballard, and L. Howery, and D. L. Berg-
man. 2011. Domestic calf mortality and producer detec-
tion rates in the Mexican wolf recovery area: Implications
for livestock management and carnivore compensation
scemes. Biological Conservation 144: 930-936.

Panasci, M., W. B. Ballard, S. Breck, D. Rodriguez, L.
D. Densmore III, D. B. Wester, and R. J. Baker. 2011.
An evaluation of fecal DNA preservation techniques and
effects of sample age and diet on genotyping success. Jour-
nal of Wildlife Management 75: 1616-1624.

Giordano, A. J., R. Carrera, and W. B. Ballard. 2011.
Assessing the credibility of jaguarundi observation using
diagnostic criteria and witness qualification. Human-
Dimensions of Wildlife 16: 360-367.

McDonald, L., T. Stanley, D. Otis, D. Biggins, J. Kuprows-
ki, and W. Ballard. 2011. Recommended methods for
range-wide monitoring of the prairie dog in the United
States. United States Geologic Survey Special Scientific
Publication, Fort Collins, Colorado, USA. Jn press.

Butler, M. J., B. A. Collier, R. D. Holt, W. B. Ballard, M. J.
Peterson, N. J. Silvy, and M. C. Wallace. 2011. Reten-
tion of butt-end aluminum bands by wild turkeys. Journal
of Wildlife Management 75: 1807-1811.

Schwalm, D. L, W. B. Ballard, E. B. Fish, and H. B. Whit-
law. 2012. Distribution of the swift fox Vulpes velox in
Texas Southwestern Naturalist 57: 393-398.

McRoberts, J. T., N. T. Quintana, W. A. Smith, W. B.
Ballard, F. P. Kehoe, and T. G. Dilworth. 2012. Greater
scaup, Ayvthya marila, nest site characteristics on Grassy
Island, New Brunswick. Canadian Field-Naturalist 126:
15-19.

Conard, J. M., B. K. Sandercock, P. S. Gipson, and W. B.
Ballard. 2012. Factors influencing survival of female elk
in a harvested population. Journal of Fish and Wildlife
Management 3: 199-208.

Flock, B. E., P. S. Gipson, R. D. Applegate, and. W. B. Bal-
lard. 2012. Distance-based habitat associations of north-

THE CANADIAN FIELD-NATURALIST

Vol. 128

ern bobwhites in a fescue-dominated landscape in Kansas.
Proceedings of the National Quail Symposium 7: 42-51.

Conard, J. M, B. K. Sandercock, P.S. Gipson, and W. B.
Ballard. 2012. Factors influencing survival of female elk
in a harvested population. Journal of Fish and Wildlife
Management 3(2): 199-208

Flock, B. E., P. S. Gipson, R. D. Applegate, and W. B.
Ballard. 2012. Distance-based habitat associations of
northern bobwhite in southeastern Kansas. Quai-VII: 42-
oul.

McDonald, D. T., W. B. Ballard, M. C. Wallace, W. H.
Miller, and J. C. deVos. 2011. Seasonal forage use and
availability by pronghorn antelope in north-central Ari-
zona. 24th Biennial Pronghorn Workshop: Partnering for
Pronghorn. /n press.

Submitted

Haskell, S. P., W. B. Ballard, M. C. Wallace, P. R. Kraus-
man, M. H. Humphrey, O. J. Alcumbrac, D. A. Butler,
and A. M. Haskell. 2011. Population dynamics of sym-
patric white-tailed and mule deer in Texas. Fauna Mono-
graphs.

Tatman, N. M., W. B. Ballard, M. C. Wallace, S. P.
Haskell, J. deVos, Jr., O. J. Alcumbrac, C. O’Brien,
and P. R. Krausman. 2011. Cause-specific mule deer fawn
mortality in central Arizona. Journal of Mammalogy.

Carrera, R., W. Ballard, M. C. Wallace, J. deVos, S. Cun-
ningham, and O. Aleumbrac. 2011. Population dynamics
of desert mule deer in central Arizona, 1960-2006. Wildlife
Monographs.

Nicholson, K., P .R. Krausman, T. Smith, W. B. Ballard,
and T. McKinney. Mountain lion use of urban landscapes
in Arizona. Journal of Urban Ecology.

Nicholson, K., P.R. Krausman, T. Smith, W. B. Ballard,
and T. McKinney. 2011. Mountain lion habitat selection
in Arizona. Journal of Wildlife Management.

Holt, R. D., M. J. Butler, W. B. Ballard, M. C. Wallace,
and R. Perez. 2010. Acute effects of radio transmitters in
Rio Grande turkeys. Journal of Wildlife Management.

Cariappa, C. A., R. W. Tobey, W. B. Ballard, and S. W.
Breck. 2010. Nelchina redux: revisiting the Nelchina
caribou-wolf system. Journal of Wildlife Management.

Phillips, R. S., W. B. Ballard, N. E. McIntyre, M. C. Wal-
lace, E. B. Fish, D. K. Holdstock, B. P. Spears, and S.
J. DeMaso. 2011. Causal mechanisms behind sex-biased
dispersal in Rio Grande wild turkeys. Ecology.

Received 5 February 2013
Accepted 10 March 2014

A Naturalist for All Seasons: Richard Merrill Saunders, 1904—1998

PHILIP COLLINS

41 York Avenue, Crosby, Liverpool L23 SRN United Kingdom; email: collinsp79@hotmail.com

Collins, Philip. 2014. A naturalist for all seasons: Richard Merrill Saunders, 1904-1998. Canadian Field-Naturalist 128(3):

289-294.

The year 2014 marks the 110" anniversary of Richard
Saunders’ birth and this tribute invites the reader to dis-
cover, or rediscover, something about the life of this
popular ornithologist.

The lure of the unexplored, the beckoning finger of nov-

elty — what field naturalist can pass them by? The
chance of finding a new bird, a new flower, the hope of

having a new experience with some old friend of the

wild in a new setting — these are sure enticements to

send the ardent naturalist afield in search of adventure.

Saunders wrote these words in 1945, and they indeed
sum up this ardent naturalist, who spent most of his long
life observing the birds and wildflowers of Canada and
beyond. The natural world, however, was not the voca-
tion for which he had trained. As a professor of history,
he taught at a number of universities for over 40 years,
during which time he published several academic pa-
pers. Had this been his only achievement, he could still
be considered to have made a worthy contribution to
society. But there was another side to the man: his love
of nature and his joy in communicating and sharing it
with many people. It is for this that he most deservedly
should be remembered.

Richard Merrill Saunders was born on 16 November
1904 in the bustling fishing port of Gloucester, Massa-
chusetts, where his father, Lee Saunders, was in the
catering business. Two years later his parents’ marital
problems reached crisis point and his father left the fam-
ily home. His widowed maternal grandmother lived in
Worcester, 60 miles to the east and it was there that his
mother, Grace Merrill Saunders, together with Richard
and his younger brother, Carroll, moved after the break-
down of the marriage. To provide for her family, Grace
lived in Boston, where she worked full-time as a sec-
retary in what was then predominately a male-domi-
nated world of business, while her son was looked after
by his grandmother, Sarah Jane, and a bachelor uncle
who was the boy’s masculine influence. To break the
family up still further, his brother was looked after by
a family in Quincy. It says much for Saunders’ charac-
ter as a youth and a teenager that, under those unusual
family arrangements, he successfully completed his
schooling in Worcester and, having decided on a teach-
ing career, he entered Clark University as an under-
craduate in 1920, obtaining his bachelor’s degree four
years later at the age of 20 and his master’s the follow-
ing year.

FIGURE |. Richard Saunders ¢ 1935.

He began his long academic career, not in the United
States, but in distant Beirut, Lebanon, at the American
University, where he taught history. It was an appoint-
ment that was to change his life in a most unexpected
way. Unbeknown to him, Dr. West, one of the profes-
sors at the University had died leaving a family of six
children in the care of his widow. She had remained
at the campus and had eventually become the house-
keeper for the entire University. She must have been a
resourceful woman for she put all six children through
Ivy League universities or Seven Sisters institutions.

One of her daughters, Anne, had just completed her
studies at Vasser College in New York when Saunders
met her on board ship as she returned to Beirut. It
proved to be the happiest of meetings, and it soon blos-
somed into courtship. While in Beirut (despite a bad

289

290

bout of malaria), Saunders made a number of visits to
Europe, including France, where he purchased a beau-
tiful Sévres vase for Anne, a gift which must surely
have made a large hole in his pay packet but which
must equally have impressed her hugely. They married
in June 1929 at South Kent, Connecticut, having left
Beirut and settled back in the United States in Ithaca.
Shortly afterwards, Saunders took up a post teach-
ing history as a Boldt Fellow at Cornell University in
New York. During this time, he studied for and in 1931
achieved his Ph.D. What followed next is recalled by
his daughter:
It might be wondered how an American ended up at a
Canadian university. When father completed his Ph.D.
it was deep in the Depression. He sent resumes to all the
universities in North America and got no replies. One
day Mother was house cleaning and picked up the rug
in front of the door that had the mail slot. Under the
rug was a letter from the University of Toronto saying
they would accept Father, but they needed to know the
next day! Needless to say, an expensive phone call was
made immediately.

The couple moved to Toronto in 1931, where Saun-

ders accepted the position of associate professor of

history at the University of Toronto, specializing in
the Age of Enlightenment European history. There he
remained until his retirement from academia in 1970.

Such are the bare facts of the career of this ener-
getic and much-loved teacher. But what of the man
himself, and what sparked off a parallel career in nat-
ural history? He provides a slight clue in a book re-
counting his visit to South Carolina. Describing the
experience of being mobbed by Least Terns (Sterna
antillarum), he likens it to a similar encounter with
Common Terns (S. hirundo) in Massachusetts. From
this, we might deduce that his interest dated from his
childhood or teens there. By 1938, however, he had be-
come sufficiently well known and respected in Toron-
to ornithological circles to be invited to become the
editor of the Toronto Field Naturalists’ newsletter, a
position he held for 27 years.

During the war, he continued to teach at the Univer-

sity, publishing several academic papers on aspects of

French Canada, (although he always considered him-
self a teacher first and a writer second) and of course
continuing his birdwatching. This latter activity was not
without its risks; having a pair of binoculars around
your neck and being in the vicinity of what are now
termed “sensitive installations” was a recipe for trouble.
On more than one occasion he was apprehended and
interrogated in the local police station. Experiences like
this were common among the birding fraternity in the
early war years, such was the fear of spies. However,
as time went on the birdwatchers became known to the
police, who treated them at best with interest and at
worst as harmless eccentrics.

Flashing Wings appeared in 1947, the year Saunders
became a committee member of the American Ornithol-

THE CANADIAN FIELD-NATURALIST

Vol. 128

Gees: “ane a. Bi

FiGURE 2. Saunders with Sally, his caciies straw Seay pick-
ing at Orangeville in 1944/5.

ogists’ Union, speaking at a dinner they held in Toron-
to. Far from being the only book he published (there
were at least a dozen more titles covering historical and
natural history subjects), it is perhaps the most unin-
tentionally autobiographical. Tucked away in its pages
are various clues and insights into some of his other
talents. For example, speaking of the value of learning
bird calls, he says: “Probably my own musical train-
ing gave me an advantage.” He was a good pianist, but
he suddenly gave it up in the late 1940s because the
pressure of his work (and no doubt his constant bird-
watching) did not allow him to keep practising to his
satisfaction. However, he was interested in encourag-
ing his students to know and appreciate music from the
period he taught, and they would gather in his house
to listen to records of Mozart and Haydn.

His book covers the period from 1934 to 1946, 12
years that saw him combining a busy program of teach-
ing at the University with his passion for the outdoors
and the birds of his adopted city of Toronto. One of the
great virtues of Saunders’ writing is his power to evoke
landscape. There is a passage at the beginning of the
book in which he details a snow-covered wood of hem-
lock trees near Maple where he watched Red Crossbills
(Loxia curvirostra) and White- winged Crossbills (LZ.
leucoptera) mov ing from branch to branch feeding. It
is a magical description. He was a man clearly sensitive

2014 COoLLins: A NATURALIST FOR ALL SEASONS:

to the beauty of nature and more than once observed
that no artist could possibly do justice to the scene in
which he was birding. For three of those 12 years he
was president of the Toronto Field Naturalists, a further
indication of the esteem in which he was held. During
this time he also experienced first extreme happiness
with the birth of Sally his daughter and then extreme
despair when his four-year-old son Alan died from
leukemia. This affected Saunders in an unimaginable
way, creating within him a sort of denial (henceforth,
he rarely mentioned his son’s death) driving him to find
solace in his field trips with the Toronto Field Natu-
ralists and his work at the University.

He recounts in Flashing Wings many of those field
trips he led, prompting George Bryant to recall recent-
ly:

On Wednesday mornings, when I was a child my friend

and | would cycle to Cedarvale Ravine before school

where Saunders led weekly early morning spring mi-

grant walks. He was a professor of history and had a

most commanding presence. His voice was resonant,

mellifluous and instantly recognisable.

Another participant on these May morning walks
was Bruce Falls who recalls: “[Saunders’] deep reso-
nant voice could be heard calling out the names of
the birds he heard and observed for the benefit of his
flock of devoted followers,” later adding, “he was an
enthusiastic, indeed exuberant birdwatcher,” and, “‘one
of the most active and best known birders of his day.”

His appearance at the field trip meetings was im-
posing — leather jacket, breeches, high laced-up boots,
and fur hat in the winter or, on other occasions, French
beret, shirt with special plaid tie, and wool jacket. He
was a naturalist for all seasons and an ornithologist for
all weathers. In November, he writes with gentle cen-
sure, “The woods are empty of watchers. They are at
home, no doubt, sleeping away the winter like the bear
— hibernating naturalists!”

His daughter, who frequently accompanied her
father on his birding trips, comments that her father
spoke as he wrote, whether it was trying to get people
to see a bird or to conduct a lecture, or both, as on one
occasion while he was taking a tutorial he suddenly
asked his students to put on their coats and go out into
Queen’s Park to see a Great Grey Owl. Falls, who was
professor of zoology at Toronto recalls:

I was surprised to learn that some of his colleagues in

the Department of History regarded Professor Saunders

as rather retiring. Apparently they had not attended his

lectures that were animated and forceful. He sometimes

appeared in period costume to heighten the drama.

Being in his presence must certainly have been an
exciting experience. Sally recalls that during a lecture
on Martin Luther, her father became so enthusiastic
when recounting how Luther had nailed his ‘“Ninety-
five theses” to the door of the chapel in Wittenberg,
Germany, that he stamped his foot so hard it went clean
through the floor boards of the lecture room. Other

RICHARD MERRILL SAUNDERS, 1904—1998

291

FIGURE 3. Saunders with his wife Anne in 1947.

examples of his humour surface from time to time in
his writing, such as when he mentions his birdwatching
— although unrelated namesake — W. E. Saunders:

How old Mr. Saunders gets around. He is seventy-four
and he goes everywhere and all the time to see his birds
— down to Kingsville, up to Toronto and over to Hamil-
ton all in one week. Quite a record for an old man. But
he is not an old man! His birds and flowers keep him
young; a retired business man but what a one. Most of
them die of boredom and golf. He lives.

Flashing Wings chronicles a quite astonishing coy-
erage of bird life in and around Toronto. In those days,
the city was much smaller than it is now although there
were signs of what was to come with his frequent ref-
erences to building concessions and the loss of habitat.
However, a// has not been lost, and some of the areas
that survive include Saunders’ beloved Cedarvale Ra-
vine (although parts have been lost to a subway station),
Scarborough Bluffs, and Grenadier Pond. To read this
book is to walk through a vanished landscape, but one
that is described so accurately that one might be watch-
ing a documentary film about it. The book includes 37
fine drawings by his friend Terence Shortt (1911
1986). Shortt was then artist-ornithologist with the
Division of Birds at the Royal Ontario Museum, be-
coming, in the year after F/ashing Wings was published,
chief artist of the Department of Art and Exhibits. In an
appendix, Saunders also includes what is almost cer-

tainly the first table of arrival and departure dates of
birds in the Toronto region, which is based on his and
Jim Baillie’s observations, making it a significant con-
tribution to the ornithology of Ontario.

Saunders seems to have integrated his academic and
the ornithological activities well, particularly after pur-
chasing a house on McMaster Avenue, in a leafy Toron-
to suburb; it became the family home until he left it the
year before his death. Many are the observations he
made of birds seen during the mile or so walk to and
from the university, and it was there that he wrote his
next book Carolina Quest, published in 1951. It des-
cribes a birdwatching trip to South Carolina with
Thomas Murray, a student in modern history at the
University of Toronto and an active birdwatcher in the
young group that birded with the Toronto Field Natu-
ralists.

Although it was written four years after Flashing
Wings, Carolina Quest in fact chronicles a visit to South
Carolina made in June 1936 when Saunders was 32
years old. The earlier book had been well received, a
fact which no doubt prompted him to build on its suc-
cess. The narrative in the first section of Carolina Quest
is a fine evocation of life and landscape in the Deep
South, culminating in a brief but unforgettable sight-
ing of a pair of Ivory-billed Woodpeckers (Campephilus
principalis), thus numbering him among the small
band of ornithologists who have seen this birdwatch-
ers’ Holy Grail, a bird now almost certainly extinct.
The 12 beautiful woodcuts in the book are by Sylvia
Hahn (1911-2001) who was the daughter of the artist
and teacher, Gustav Hahn, and the sister of Saunders’
secretary at the University. In 1934, she had joined the
staff of the Royal Ontario Museum, to which she dedi-
cated almost her entire artistic and amazingly versatile
career. The final chapters deal with some of the birds
that, in the past, had seemed so characteristic of the
South and which were now increasingly to be found
within the borders of Canada, thus providing an avian
bond between the “Dominion of the North” and the
“Deep South.” A hint of things to come occurs when
Saunders cites global warming as a possible factor for
this expansion. He also includes what must be one of
the earliest (and most humorous) descriptions of a full-
blown “twitch” (numbering 80 birders) when a King
Rail (Rallus elegans) appeared at Catfish Pond, High
Park in Toronto.

The 1960s saw a quite radical decision made in the
Saunders household. Having lived in Canada since they
were married, Richard and Anne had grown used to,
and now preferred, the Canadian way of life and so
chose to take the nationality of their adopted country
— although in those days it was a requirement that in
so doing they had to relinquish their American citizen-
ship. They felt that having Canadian passports made
travel easier, particularly as America was then heavily
involved in the Vietnam War, a fact that may well have
contributed to their decision.

THE CANADIAN FIELD-NATURALIST

Vol. 128

They were soon to make good use of these new
passports, as Saunders, accompanied by Anne, took a
sabbatical from the University, first visiting Europe,
then the United Kingdom, travelling by steamship. For
several weeks, they journeyed around Europe, mainly
in France, a country whose language he spoke fluently,
before crossing the channel to England. Here they stayed
for four months while Saunders undertook historical
research. Nevertheless, he still found time to make
many field trips and to meet and become acquainted
with Britain’s ornithologists and discover its birds. The
second part of the sabbatical was spent back in North
America, driving down the east coast to Florida then
continuing west to California, before turning north to
Vancouver, taking in Yellowstone Park and the Cana-
dian Rockies and returning home to Toronto. As her
husband had never learned to drive, this challenging
itinerary was undertaken by Anne.

Resuming his University duties, Saunders recalled
how on one occasion his authority was challenged:

In a fourth-year seminar in Sidney Smith Hall a girl ar-
rived one afternoon, sat down and took out a nice black
pipe which she proceeded to light up and to puff bold-
ly, cocking an eye at me every so often. The rest of the
group also watched to see what would happen. As it was
obvious that, in the spirit of that day, she was trying to
push me into some sharp reaction I decided to pay no
attention at all and carried on the group discussion as
usual. As it worked out she never tried anything like it
again and proved to be a very good and attentive student.
Nonetheless, she defied the professor and had done her
own thing in good 1960s style.

Saunders’ calm and experienced approach to han-
dling this situation says much about his character.

In 1965, he relinquished his position as editor of the
Toronto Field Naturalists’ newsletter, bowing out with
a résumé of the activities of that lively institution, which
he published in the 26-page booklet Joronto Field
Naturalists’ Club — Its History and Constitution. He
retired from academic life a few years later, having
become increasingly afflicted by deafness which also
dramatically reduced his birding activity. Bird identi-
fication through birdsong had always been his
strength, and he once observed that when springtime
arrived with its heavy foliage, 75% of bird observi ing

was done by the ears.

To compensate for the onset of deafness, there were
the wildflowers — an area of natural history that had
interested him as a photographer since his meeting with
Mary Ferguson in the late 1960s at the Toronto Photo-
graphic Club. She had a deep knowledge of flowers,
and they had an immediate rapport. Mary’s husband
was a professor in the Department of Pharmacy at the
University and the Saunders and Ferguson families
soon became close friends. They began to go on field
trips together and were frequent guests at the Fergu-
sons’ cottage on the shores of Lake Huron at Georgian
Bay.

9) . Re F
2014 CoLLins: A NATURALIST FOR ALL SEASONS:

In 1973, the two families’ most extensive journey
together took them, this time by aeroplane, on an ex-
tended tour of Europe across France, Germany, Austria,
and Switzerland. It is regrettable that Saunders never
turned this trip (or indeed that made on his sabbatical)
into a book. While in Europe, it is likely that plans were
discussed with Mary, which resulted in the joint produc-
tion of two books, Canadian Wildflowers (1971) and
Canadian Wildflowers through the Seasons (1982). In
the preface to the latter, Saunders wrote:

The discovery of a rare or unusual flower, a new spe-

cies... Or an expanse of flowers in fields, woods or

swamps exhilarates and spurs us on. We forget the
swarming black flies and mosquitoes, the difficult pro-
gress into a bog or the tiring climb up a mountain trail...

When you go out with the same purpose, take along

patience and perseverance as part of your equipment,

and take pictures for the joy of it.

Note how his words, “the discovery of a rare plant
or unusual flower,” echo those quoted in the opening
paragraph of this tribute: “the chance finding of a new
bird.” Saunders was very much an active, rather than a
passive naturalist.

His contribution to ornithology and conservation can
be summed up in those four last words from the pref-
ace quoted above: “for the joy of it.” He communicat-
ed that joy in an infectious yet authoritative way, and
in so doing introduced a great many people, both young
and old, to a lasting enjoyment in studying birds.

The final decade of Richard Saunders’ life was total-
ly overshadowed by his wife’s suffering a severe stroke
which confined her to hospital until her death in 1993.
They had been happily married for 64 years. Now pro-
foundly deaf, and at his request, Saunders spent his last
year in a senior retirement home close to his daughter
where, sitting on his veranda, yet unable to hear his be-
loved birds, he could at least enjoy the sight of them at
the feeders. After a short illness, he died in a hospital in
Huntsville on 25 July 1998 aged 93 and is buried in the
family grave in his beloved adopted city of Toronto.

That he was an interesting man is without question,
and interesting people must never be allowed to sink
into oblivion. If this tribute has opened a small window
onto his life and stimulated others to research it still
further, then it will have achieved its purpose. Let the
final words be from the man himself:

Often people ask me why I continue to look at the same

birds week in and week out, year after year. Somehow

they don’t seem to realise that a sort of friendship springs
up between observer and birds. And whoever comes to
the end of wanting to see friends and to learn more about

them?

Acknowledgements

The author gratefully acknowledges the invaluable
advice, support, and recollections of Richard Saunders’
daughter, Sally Saunders, who also made available the
photographs. Thanks are due also to Margaret McRae,

RICHARD MERRILL SAUNDERS, 1904-1998 293

FiGuRE 4. Anne Saunders favourite photo of her husband. Late
1960s.

president of the Toronto Field Naturalists (TFN) for
her helpful information and for enabling the initial
contact to be made with Sally; Marilynn Murphy,
formerly on the TFN newsletter committee; George
Bryant and Bruce Falls for their recollections; and
Sandi Kane of Clarke University Massachusetts and
Michel Duquet of the Canadian Historical Associa-
tion.

Bibliography of Richard Merrill Saunders

Saunders, R. M. 1925. The Historical Geography of Cape
Ann. Clark University Press, Worcester, Massachusetts,
USA.

Saunders, R. M. 1931. The Provincial Academies of France
during the 17th and 18th Centuries. Ithaca. New York,
USA.

Saunders, R. M. 1939. The Emergence of the Coureur de
Bois as a Social Type. Canadian Historical Association
Annual Report 18(1): 22-33.

Maheux, A., and R. M. Saunders. (translator) 1942. French
Canada and Britain: A New Interpretation. Ryerson Press,
Toronto, Ontario, Canada.

Saunders, R. M. 1943. History and French-Canadian Sur-
vival. Canadian Historical Association Annual Report
22(1): 25-34.

Saunders, R. M. 1944. Whither the Near East? Canadian
Institute of International Affairs, Canadian Association for
Adult Education, Toronto, Ontario. Behind the Headlines
4(4).

Wilkinson, B., and R. M. Saunders. 1944. Stories from Cana-
dian History: No. 30. Imperial Optical Company. Toronto,
Ontario, Canada.

Saunders, R. M. 1945. Flashing Wings. McClelland and
Stewart, Toronto, Ontario, Canada. 388 pages.

Saunders, R. M. 1946. The French-Canadian Outlook. G.
A. Klinck, 16 pages.

Saunders, R. M. 1946. Introduction. Pages ix—xili in Educa-
tion for Tomorrow: a series of lectures... organised by the
Committee Representing the Teaching Staff of the Univer-
sity of Toronto. Edited by Richard M. Saunders, University
of Toronto Press, Toronto, Ontario, Canada.

Saunders, R. M. 1951. Carolina Quest. University of Toronto
Press, Toronto, Ontario, Canada and University of South
Carolina Press, Columbia, South Carolina, USA. 119 pages.

Saunders, R. M. 1955. The Function of History: A Philo-
sophe View. Canadian Historical Association, Report of the
Annual Meeting 34(1): 33-41.

Associate Editor’s Footnote

R. Saunders should not be confused with two un-
related W. Saunders, who are familiar to many Cana-
dian naturalists.

“Known as Will or W.E. (short for William Edwin),
W. E. Saunders was born in 1861, second in William
Saunders’ family of one daughter and five accom-
plished sons. The senior Saunders later became found-
ing director of the Dominion Experimental Farm in
Ottawa.

“Of the brothers, only W.E. remained in London,
where he operated a wholesale pharmaceutical compa-
ny until his death in 1943. Saunders’ broad-ranging
interests found outlets in civic, historical, horticultural,
educational and musical circles.” From Wake, Win-
nifred. 2011. W. E. Saunders, London’s beloved natu-
ralist. Londoner 18 October. Available: www.thelon
doner.ca/2011/10/18/w-e-saunders-londons-beloved-
naturalist (accessed 19 May 2014).

Will Saunders was a major part of the early natural-
ists group active at Point Pelee as partly documented
by a later University of Toronto historian who includ-

THE CANADIAN FIELD-NATURALIST

Vol. 128

Saunders, R. M. 1965. Toronto Field Naturalists’ Club — Its
History and Constitution. Toronto Field Naturalists’ Club,
Toronto, Ontario, Canada. 26 pages.

Saunders, R. M., and M. Ferguson. 1971. Canadian Wild-
flowers. Van Nostrand Reinhold, New York, New York,
USA. 192 pages.

Saunders, R. M. 1979. Birds in the Wild. Van Nostrand Rein-
hold, New York, New York, USA.

Saunders, R. M. 1979. Flower photography. Journal of the
Canadian Rhododendron Society 8(41).

Saunders, R. M., and M. Ferguson. 1982. Canadian Wild-
flowers through the Seasons. Van Nostrand Reinhold, New
York, New York, USA. 160 pages.

Saunders, R. M. 1985. Flower photography. Journal of the
American Rhododendron Society 39(4).

Received 2 April 2014
Accepted 25 April 2014

ed many references in A Life with Birds: Percy A. Tav-
erner, Canadian Ornithologist, 1875-1947 by John L.
Cranmer-Byng. Canadian Field-Naturalist 110(1): 1—
254 (1996).

For information about W. Saunders, senior, see Cody,
W. J., D. B. O. Saville, and M. J. Sarazin. 1986. Sys-
tematics in Agriculture Canada at Ottawa 1886-1986.
Agriculture Canada Historical Series number 28. 81

pages.

About the Author

Philip Collins lives in the United Kingdom and has
been bird watching since childhood. He was a member
of the Southwest Lancashire Ringing Group where he
organized a Barn Owl nest box scheme and study
group. He is currently secretary of the Liverpool
Ornithologists Club. Back in 1964, he was given a
copy of A Birdwatcher s Anthology edited by Roger
Tory Peterson, which included an extract from Saun-
ders’ book Flashing Wings. The passage so captured
and remained in his imagination that he finally deter-
mined to discover all he could about the life and work
of its author.

Book Reviews

Book Review Editor’s Note: We

are conunuing to use the current currency codes. Thus Canadian dollars are CAD, U.S.

dollars are USD, Euros are EUR, China Yuan Remimbi are CNY, Australian dollars are AUD and so on.

ZOOLOGY

Beetles of Eastern North America

By Arthur V. Evans. 2014. Princeton University Press, 41 William Street, Princeton, NJ, USA, 08540-5237. 560 pages,

23.16 USD, Paper.

For those with an inordinate fondness for beetles,
finding suitable field guides and reference material has
traditionally been a bit of a challenge compared to more
charismatic taxa such as butterflies, moths, and (more
recently) odonates. My dog-eared 1983 copy of the
Peterson Field Guide to the Beetles of North America
covers I11 families, and despite the 600+ line drawings,
there are only 65 colour paintings (this holds true for
the 1998 edition as well). More recent field guides to
the tiger beetles (Cicindelidae) and jewel beetles (Bu-
prestidae) are welcome and more colourful additions,
but leave the majority of beetles treated in a fairly su-
perficial way by more general insect books. On the
more technical end of the spectrum, the comprehen-
sive two-volume American Beetles by CRC Press (Ar-
nett and Thomas 2000; Arnett et a/. 2002) are an in-
valuable reference, but the pair will set you back $280,
and the combined 1300+ pages of text may be a bit
daunting for the average field naturalist.

Enter Arthur V. Evans’ Beetles of Eastern North
America. According to its preface, the primary goal
of the book is to present the beetles of eastern North
America in an engaging format that is accessible to
the amateur naturalist interested in beetles, while
authoritative enough to serve the needs of the profes-
sional biologist. Mission accomplished. No longer
will the vast majority of Coleoptera languish under-
appreciated! The 1500+ colour photographs by some
of the best insect photographers in North America
truly capture the beauty and diversity of our eastern
beetle fauna like no other text before it.

Arthur’s fantastic new book covers 1409 species
and all of the 115 beetle families known from North
America east of the Mississippi River. With an 8” x
10” format and 4 Ibs of heft, it is not intended for the
field and, despite its size, covers only about 10% of
the known species for this region. It should however,
allow readers to identify to the species level many of
the conspicuous beetles they are likely to encounter,
and to assign most others to the appropriate genus or
family. Beetles of Eastern North America reflects the
latest taxonomic works, which may confuse some
readers where a once-familiar species is reassigned

to another group. For example, Ghost Tiger Beetle,
long known as Cicindela lepida (including in recent
field guides) is now referred to as Ellisoptera lepida.
Fortunately, Beetles of Eastern North America
includes a taxonomic classification from order
through suborder, series, superfamily, etc. down to
genus/species. While this appendix may be overkill
for some, it can help readers more easily find a reas-
signed species, and also helps to demonstrate the
relationships amongst the diverse beetle groups.

The book opens with about 50 pages of introduc-
tory text covering beetle anatomy, behaviour, natural
history, and where and when to find beetles. Accom-
panying photos helpfully demonstrate antennal types,
body shapes, tarsal claws, and other features. Arthur
has an engaging writing style and it is a much more
interesting read than one might expect. Guidance on
how to explore the world of beetles is provided, from
passive observation and photography to actively col-
lecting and preserving beetles, and even how to rear
live beetles and their larvae. The introduction is fol-
lowed by an illustrated dichotomous key to 68 of the
most commonly encountered families in eastern North
America. While not as comprehensive as the keys in
the two-volume American Beetles, they should point
the reader in the right direction, especially when used
in conjunction with family descriptions from the main
body of the book. No one ever said beetle identifica-
tion was easy!

The main part of the book is organized by family,
with a half- to full-page summary of family-level diag-
nostic characters such as length (mm), shape, color,
and distinctive morphological features. Tips are given
on how to distinguish each family from other, similar-
looking families. A brief overview of the natural his-
tory of the group is also provided, as well as notes on
suitable collection methods. Finally, the number of spe-
cies and genera of each family found in the Nearctic
and eastern North America are given (if known). This
information is very helpful in understanding how di-
verse the group is, and how many potentially co-
occurring species are not illustrated in the book. Sim-
ilarly helpful is the total number of species in each

phe

296

genus known from east of the Mississippi River, which
is typically provided at the end of the species’ accounts.

The species accounts are what set this book apart.
There are generally only four species treated per page;
this generous layout allows each species’ photo to be
sufficiently large that the diagnostic features described
in the accompany text are often visible. And as a bonus,
most of the crisp, vivid photos are of live beetles rather
than pinned specimens. One should really use the keys
to determine the family, but I would not be surprised
if many readers “picture-key” first, and use the text to
confirm a beetle’s identification. The written descrip-
tions of diagnostic features provided for each species
are very useful (if somewhat dry), particularly for
small or subtle characters that may not be evident in
the photo.

The remainder of each species account consists of
concise notes on seasonality, habitat, food preference
(for adults and occasionally larvae) and distribution.

Birds of the Kenya’s Rift Valley

THE CANADIAN FIELD-NATURALIST

Vol. 128

No range maps are provided (which would be daunt-
ing and space-consuming) but the range descriptions
help determine whether a species potentially occurs 1n
your general area. Non-native species are also flagged.
Surprisingly, Emerald Ash Borer and Asian Long-
horned Beetle are not covered in the species accounts
but are presented in the introductory section on /nsects
as Pests. More information on the behaviour and nat-
ural history would have been welcome for many of the
species, but compromises always have to be made
when trying to cover as many species as possible in a
reasonably-sized tome.

In short, Beetles of Eastern North America is an
excellent book that will be much loved by field natu-
ralists and entomologists alike, especially given its
very modest price.

Dr. RoBERT F. FOSTER

Northern Bioscience, 363 Van Horne Street, Thunder Bay,
ON, Canada, P7A 3G3

By Adam S. Kennedy. 2014. Princeton University Press, 41 William Street, Princeton, NJ, USA, 08540-5237. 256 pages,

19.75 USD, Paper.

Birds of Kenya’ Rift Valley is the fourth in a series
of Princeton University Press WILDGuides by Adam
Scott Kennedy, a former safari camp manager and now
full-time professional guide. This new field guide fo-
cuses on the Kenya’s portion of the Rift Valley, which
runs through East Africa from the Gulf of Aden south
to Mozambique. Kenya’s Rift is rich in bird life, with
a variety of habitats from soda Lake Magadi and the
lush wetlands of Lake Navaisha to the cliffs of Hell’s
Gate. In particular, Lake Nakuru and Lake Bogoria are
famed for their vast flocks of flamingos, which can
number in excess of one million birds. Kenya is a re-
nowned location for birding, and this field guide is well
suited for visitors of the many national parks and Im-
portant Bird Areas of the Rift Valley.

Not surprisingly, there is considerable overlap with
Kennedy’s two previous bird guides; half of the 320
species covered in the Rift Valley guide are also includ-
ed in the Birds of the Masai Mara. Nonetheless, Birds
of Kenya Rift Valley covers a fair bit of new ground
since there are approximately 1100 avian species
known from Kenya, and the Rift Valley encompasses
habitats and areas not found in his Serengeti and Masai
Mara guides. For example, Kennedy devotes a full page
to Sharpe’s Longclaw (Hemimacronyx sharpei), an
imperilled grassland species found in the Rift Valley
highlands. Unfortunately, the guide does not include
the similarly endangered Abedare Cisticola (Cisticola
aberdare), which is also found in the highland grass-
lands of Mau Narok and the Abedares. Oddly, it also
doesn’t include such notable species as the Secretary
Bird and Kori Bustard. However, the guide does cover
19 species of weavers and no less than ten species of

starlings (including the aptly named Superb Starling),
which may be surprising to many North Americans
used to just one garden variety starling.

Rather than by standard taxonomic order, the guide
is arranged into six sections: Lakes and Marsh; Up in
the Air; Birds of Prey; Grasslands and Open Areas:
Woodland, Scrub and Garden; and Nightbirds. The
intent is to help novices more rapidly locate the likely
bird species in the guide based on where it is observed,
or by groups of birds (raptors, aerial specialists, noc-
turnal birds) that may range widely over many habitats.
This approach has its limitations due to poorly defined
or overlapping groups (I must admit I prefer the tra-
ditional approach), but may make it easier for begin-
ning birders to identify potential candidates.

What sets this guide apart from other field guides
are the 500+ spectacular photographs and impressive
artistry of the layout, with truly seamless blending of
multiple photos. There are typically 2-3 species per
page, with full page accounts for some of the more
charismatic or spectacular species such as the Long-
tailed Widowbird and Hemprich’s Hornbill. The in-
formative and accessible species’ accounts have tips
on identification and similar species, and touch upon
distinctive songs or calls, behaviour, ecology, and other
points of interest.

The guide is “intended to be an inspirational, port-
able, and easy-to-use introduction to the many hun-
dreds of species that have been recorded in the Rift
Valley”. At 540 g, Birds of Kenya s Rift Valley definite-
ly more compact than the more comprehensive Birds
of Kenya and Northern Tanzania (1000 g, 576 pages,
1114 spp.) by Zimmerman er al., or Stevenson and

2014

Fanshawe’s Birds of East Africa (1125 g, 602 pages,
1388 spp.). Serious birders will likely want to carry
one of these more detailed guides, but Kennedy’s new
book would serve most safari goers well in Kenya’s

The Amazing World of Flyingfish

Book REVIEWS 297

Rift Valley, and its great photos make it a worthy addi-
tion to any birder’s bookshelf.
Dr. ROBERT F. FOSTER

363 Van Horne Street, Thunder Bay, ON, Canada, P7A 3G3

By Steve N. G. Howell. 2014. Princeton University Press, 41 William Street, Princeton, NJ, USA, 08540-5237. 64 pages,

12.95 USD, Cloth.

Many years ago, when I was out on the Gulf Stream
off Carolina, I glimpsed a “bird.” I spun around, but it
had disappeared. Birds just do not disappear, so this
was very puzzling. This repeated several times until |
realised I was looking at flying fish. Puzzlement then
changed to wonder.

I have been fortunate to see these fabulous creatures
many times since that day. Standing on the bow of a big
cruise ship I noticed the other passengers would ignore
birds, but these little fish always caused a stir. There is
something magical about flying fish. They appear out
of “nowhere” and sparkle in the sun over the deep blue
water for a few seconds, before they disappear.

Finding information on these fish has proven diffi-
cult. One book I have (A Field Guide to North Atlantic
Wildlife, by Proctor and Lynch, Yale University Press)
has one page covering five look-alike species. My Field
Guide to Atlantic Coast Fishes (By Robins, Ray and
Douglass, Houghton Mifflin) covers another seven
species. Neither book adds much to my knowledge.
Now Steve Howell has written this delightful book.
Despite its small size, it contains plenty of useful mate-
rial. The author acknowledges that the family Exo-
coetidae is poorly known, but he has assembled all the
data he can find. The more you learn the more fascinat-
ing these fish become. To add to their enchanting ap-
pearance they now have the most intriguing names.

How can you not perk up for a Leopardwing, a Pacific
Necromancer or a Violaceous Rainmaker?

The author admits to taking “a few thousand fuzzy
images, .... blank water or water punctuated by only a
splash.” I too have spent hours trying to get a photo-
graph. Despite there being hundreds of fish actively fly-
ing | have way more than my fair share of fish-less
ocean water. Therefore it is to Howell’s great credit (and
his infinite patience) that this book is so beautifully il-
lustrated. Not only are the fish photos technically accu-
rate, many have an ethereal quality. There are photos
of some other oceanic “flyers” — rays, dolphins, squid,
seals and penguins for comparison. As well there are
some nice sequence shots of boobies catching flying
fish on the wing.

The Amazing World of Flyingfish (yes it is correct-
ly one word) is amazing. Once I opened it I did not
put it down (Perhaps my one complaint is the book is
too short. But then one of the author’s points is the
lack of knowledge about these animals). I now have a
strong urge to be back on the bow, looking at the deep
blue water, waiting for that mysterious explosion of
gossamer and silver. Fish enthusiasts, travelers, natu-
ralists and photographers will love this book, so buy
it for friends and family. Most of all buy a copy for
yourself.

Roy JOHN
2193 Emard Crescent, Ottawa, ON, Canada, K1J 6K5

A Field Guide to the Larger Mammals of Tanzania

By Charles Foley, Lara Foley, Alex Lobora, Daniela De Luca, Maurus Msuha, Tim R. B. Davenport, and Sarah Durant. 2014.
Princeton University Press, 41 William Street, Princeton, NJ, USA, 08540-5237. 320 pages, 29.95 USD, Paper.

When I first saw this book I was disappointed by the
title “Larger Mammals.” I have long been frustrated by
African film makers extreme focus on Lions, Leopards,
Buffalo, Elephant and Rhino to the exclusion of all else.
I have seen three brief videos of the Honey Badger,
totalling less than 15 minutes. This is surely one of the
most inscrutable mammals of Africa. It is strong, fear-
less, tireless, tough and courageous. They will defend
against any animal, regardless of size. Lions usually
lose this fight. Their thick loose skin is impenetrable to
bee stings, porcupine quills, and animal! bites. They eat
anything — insects, frogs, tortoises, rodents, turtles,
lizards, snakes, eggs, birds, honey, fruit and vegetables.

They are immune to the venom of cobras, mambas and
adders. Surely such an astounding beast is worth a few
hours of TV time.

When I opened the book I sighed with relief. The
Honey Badger was included, as well as many other not-
so-large mammals. In fact the authors cover galagos as
small as a 12 cm (5 inches). So mongooses, weasels,
civets, genets, porcupines and hedgehogs are all incor-
porated. And all creatures are treated equally. The Lion
has a two page coverage, but so does the diminutive
White-bellied Hedgehog. The authors cover 135 of the
340 species officially recorded in Tanzania. It does not
cover rodents (except three of the big ones), bats or

298

shrews that make up 60 percent of the mammal list. I
did see and photograph one mouse; a Natal Mastomys,
Mastomys natalensis, that took me two years and
multiple emails to identify (by an expert in Belgium).
The authors provide a URL for The Field Museum in
Chicago (archive. fieldmuseum.org/tanzania/index.html
— try it, it is worth a look) that provides an excellent,
on-line field guide to all the mammals of Tanzania. Only
16 species of confirmed sea mammals are in this book
and the authors state this needs further research, as
more species are listed for neighbouring Kenya. Oddly
the Field Museum lists 24, but the unconfirmed ones
have no detail.

The species accounts are tremendous. The text cov-
ering description biology distribution and population
is sound. The range maps are large and easy to follow.
The authors have added a text box on where to look
for each species. This is very useful to all tourists. I do
not feel so bad for not seeing a (common) Ground Pan-
golin when they say you are “very lucky if you do see
one.”

Each species has at least one and often two or more
photographs. They are consistently admirable. Some
(Bush Hyrax, White-bellied Hedgehog, Clawless Otter,
Cheetah and many antelope) are really cute. The night
photos of animals show the eyes as white or red disks
that look a little odd. I thought that it would look bet-
ter if they were photo-shopped, but it is realistic and
likely what you with a flashlight.

One animal was quite a surprise; the Kipunji. A close
relative of the mangabes, it was the first new monkey
to be discovered in 20 years. Only about a thousand of
these mammals live in a small corner of Tanzania in a
few forest reserves and national parks, making it crit-
ically endangered. Its habitat is degraded and fragment-
ed and it suffers from hunting and illegal habitat des-
truction. No wonder it has a sad look.

THE CANADIAN FIELD-NATURALIST

Vol. 128

There are seven pages of photos that were not la-
belled so they confused me until I back referenced the
pages in the index. They are “Species comparison
spreads” — a set of photos with similar species in close
proximity. So the reader can compare all the spotted
genets and civets, all the mongooses, the antelope and
so on. This is so useful for field comparison of look-
alike species. When the text notes there are similar
species it points you to the correct comparison page.

There is a very important section on the National
Parks and major protected areas. Each site has a large
photograph overlaid by a locator map and some basic
facts. Text boxes list the mammals present with a letter
and coloured code indicating their visibility and the
chance of seeing one on a two or three day trip. Cou-
pled with the information on the individual species this
provides valuable information when planning a trip.

This book is an absolute must for a person going on
a safari in Tanzania. It is better than the other mammal
guides I own or have seen. It will set you up on where
to look. It will help you separate similar species and it
will add wonder to your trip. The only other comment
' have is that I would like to have this information on
Kenya. There is no need to repeat most of the text. A
visitor would only need the range maps and the Na-
tional Park section. Kenya also has a few additional
mammals (Grevy’s Zebra and White Rhinos for exam-
ple) so some type of supplement would be nice.

Roy JOHN
2193 Emard Crescent, Ottawa, ON, Canada, K1J 6K5

Reviewers note: While researching the White-tailed Mongoose
I was taken aback to find a page devoted to the hunting exploits
of a decorated “conservationist.” He had arranged to have bait-
ed traps set by his safari hosts. Not only did he pot his White-
tailed Mongoose, his trusty Remington nailed a Striped Hyena,
two Honey Badgers, a Rock Hyrax, a Crested Porcupine, a
Black tailed Mongoose and an African Wild Cat. These hardly
seem good “trophies” or good to eat.

A Feathered River Across the Sky. The Passenger Pigeon’s Flight to Extinction

By Joel Greenberg. 2014. Bloomsbury Publishing Inc., 1385 Broadway, 5" Floor, New York, NY, USA. 10018. 304 pages,

26.00 USD, Cloth.

The Passenger Pigeon is perhaps the most iconic
species representing modern extinction and its story
has been repeated many times, but only rarely in any
detail. The bird’s plummet from vast numbers to extinc-
tion has often served as a parable illustrating our own
species’ merciless, Darwinian penchant to convert the
rest of nature into ourselves. This year is the 100" since
the last Passenger Pigeon expired in a Cincinnati zoo.
This centennial is being observed in several events
across North America. Joel Greenberg’s book is the
first detailed account in decades of the Passenger
Pigeon’s unbelievably rapid descent from billions to
none. Greenberg provides meticulous descriptions of
our treatment of Ectopistes migratorius; our responses

to its decline and extinction, our overwhelming igno-
rance in trying to explain what happened, our unrelent-
ing avarice in “harvesting” it, and our staggeringly
short-lived and embarrassingly superficial record of
what it was like to have billions of pigeons fly over-
head for days at a time, and to have hundreds of mil-
lions swoop in to occupy a roost or nesting ground.
And what have we learned from this tale of the pigeon
of Biblical abundance? I think an accurate answer
would be, not much so far, but this book gives us
some food for further thought and a basis to consider
critically some current approaches to conservation of
biodiversity and even our place in nature. Such con-
siderations are appropriate on this centennial of the

2014

Passenger Pigeon’s extinction especially given the
recognition that we are facing a new so-called 6" glob-
al extinction for line which, as is usually assumed to be
true for the Passenger Pigeon, we have ourselves to
blame. Perhaps new assessment of the conservation
implications and lessons of the facts of the Passenger
Pigeon’s biology and decimation will help us mitigate
the current biodiversity crisis; or perhaps it won’t.

First, some Passenger Pigeon background, gleaned
from this entertaining book. It was a bona fide pigeon,
closely related to the extant Band-tailed Pigeon (hint
to Passenger Pigeon fans; it would be truly helpful in
thinking about Passenger Pigeon biology to apply
some phylogenetic analysis and direct study of this liv-
ing bird), and similar to, but slightly larger than our
ubiquitous Mourning Dove. It was colourful, a fast,
agile flier reputed to zoom along at 60 mph. It pro-
duced varied vocalizations, though the descriptions
seem somewhat conflicting (screaming, cooing, woo-
ing, bell-like, sleigh bells ...), so it is not clear to me
what their voices sounded like. The Passenger Pigeon
was a typical pigeon in that it built sloppy, rickety nests
and fed its squabs pigeon milk (“a curdy substance
resembling loose rice pudding”) produced by the lining
of the crop of both sexes. They usually laid only | egg
per nest, nested once per year (though this seems un-
certain) and probably lived for 10-20 years or more in
the wild. They ate almost anything although especially
fond of nuts including American Chestnuts, acorns of
most species of oaks, and beechnuts all of which to
Thoreau’s astonishment they swallowed whole. They
also ate plants, fruit, invertebrates, seeds, green veg-
gies, grains and more. Their catholic, omnivorous diet
is one of their oddly human traits. They occupied most
of eastern North America from the Gulf Coast all the
way to Hudson Bay (where it is worth noting, there are
no nuts) moving about in large flocks of 1000’s to mil-
lions to billions. These movements were unpredictable,
but in part appeared tied to availability of food, espe-
cially mast of nut trees. Many times these flocks saved
Native and European villages from starvation and,
hence, to the immigrants at least, proved the existence
of a benign Providence when he showed largesse in
sending in the pigeons to feed deprived colonists. In
gratitude, “we” eradicated this splendid gift of the
Creator.

Greenberg’s book is a wealth of interesting factoids,
here is a small sample to whet your interest. The largest
flock ever “counted” flew over Fort Mississauga, On-
tario, Canada in 1860 and numbered 3,717,120,000
(confidence limits undoubtedly huge) individuals.
Mayor McCallion in her inaugural term was only mild-
ly thrilled, and expressed concern about the effect of
guano on business development and tourism. The main
flock zoomed along at “60 miles per hour”, took 14
hours to pass and of course blotted out the sun. Smaller
flocks continued to fly over for the next several days.
The largest nesting “roost” was reported from Wiscon-

Book REVIEWS

299

sin and “occupied 850 square miles”. The total number
of birds in North America was thought to be about 5 bil-
lion. A large flock (greater than 50,000,000 say), sound-
ed like a “loud rushing roar succeeded by instant dark-
ness”, “a noise like the crackling of a fire among dry
leaves”, “a low pitched hum as they appeared on the
horizon ...that increased to a mighty throbbing... chil-
dren screamed, women sought shelter, horses bolted”.
Audubon recorded a huge flock and mentioned the
blocking of the sun, and how the dung fell “not unlike
melting flakes of snow”, definitely a bravely quaint
way to describe being engulfed in a guano storm. The
volume of these avian missiles never abated for 3 days,
during which of course the sun was eclipsed. Other
claims are made (e.g., it outnumbered every other bird
species on earth or every other terrestrial vertebrate
in North America). All the foregoing is subject to the
error of eyewitness testimony and reporting. But an
unassailable fact is that less than 50 years after the On-
tario super flock, the Passenger Pigeon was extinct in
the wild constituting, if nothing else, a tribute to
human ingenuity. (See http:http://passengerpigeon.org
/index.html (Project Passenger Pigeon), for many more
details)

Greenberg spends much of the book chronicling the
Passenger Pigeon’s abundance, decline and ultimate
extinction. He apologizes for the amount of detail he
incorporates into the latter topic, and perhaps he should,
as it does become a tad tiresome ploughing through
pages describing when, where and by whom the last
one was lost from the wild, particularly when there
seemed to be bigger issues being largely ignored. His
excuse is that he wants to be as scholarly as possible,
but it is a monotonous litany of shooting pigeon—like
birds for the notoriety of bagging the last wild Passen-
ger Pigeon. There is not much of significance to be
learned from this, except perhaps that many people
crave acclaim, especially when it is as easy as pulling
a trigger. The end of wild birds came in about 1902, |
think.

I would like to focus on 4 questions that Greenberg
discusses less intensively than the awe inspiring abun-
dance and slaughter. What was the ecological “role”
of the Passenger Pigeon? How could we know so little
about them given their importance and ubiquity? How
did 19" century attitudes toward their decline and ex-
tinction compare to those we express today about the
nature of species at risk? This question is inextricably
tied to the big question; what caused the extinction of
the Passenger Pigeon, and Greenberg leaves little doubt
about his answer; humans. A fourth question, not real-
ly covered by Greenberg, is whether we can or should
‘de-extinct’ the species, or at least some sort of fac-
simile.

What was the impact of the Passenger Pigeon on
North American ecosystems? Greenberg discusses this
with interesting possible examples in Chapter 1. The
huge numbers of Passenger Pigeon’s were enough to

300

make people wax poetic, express wonder and so on.
Some also quailed and were fearful and described
consequences that are not generally thought of when
we lament the loss of this icon. When the huge flocks
whirred, cooed and jingle-belled into an area to roost,
their sheer numbers and cumulative mass caused their
roosting trees to lose large limbs or to collapse or up-
root. In sound and destruction, big flocks were similar
to a tornado, leaving destruction everywhere, except
tornados don’t routinely deposit vast amounts of gu-
ano. Soon after the birds arrived the faecal output was
over half a metre deep and equivalent to “thousands
of wagon loads”. Therefore, after the masses moved on
not only were all edible fruits, nuts, vegetables, grains,
small invertebrates and valuable timber gone, there
also was a huge mass of downed woody debris, thou-
sands of pigeon carcasses, remaining trees festooned
with excrement, and a choking ground cover of murky
indigestibles smelling like a poorly managed factory
farm for poultry. The remaining trees cried, the under-
story died and then it all dried and a lightning bolt con-
verted the forest to ashes, just like modern logging. In
many cases, people didn’t wait for lightning, but set fire
to the roosting/nesting areas, especially if it was a nest-
ing colony, so as to fry the birds and their flightless
squabs. They also burned down potential roosts to dis-
courage the arriving flocks from staying and siphoning
up local crops. All of this 1s worth thinking about when
one considers how these days a few hundred starlings,
crows, rock doves, or geese elicit bitter complaints from
the citizenry because of noise, droppings, collisions
with aircraft, and occasional direct aggression. What
would we do with a few billion pigeons and their
whimsical falling flakes?

The impact of these birds on eastern North Ameri-
can ecology must have been prodigious as Greenberg
suggests. What that impact was is not at all clear, nor
much discussed in the Passenger Pigeon literature,
which makes this book interesting in that Greenberg
does speculate on some possible effects of the pigeon’s
passing. We hear a great deal these days about the
“functional role” of species X on the “ecosystem” and
how ecosystems will change or even collapse if we pull
out just one species. But the Passenger Pigeon was not
just any species; it was the most abundant terrestrial
vertebrate on earth. Couple that with the American
Chestnut, population 30 billion and both going extinct
within the same century, and it is surprising that any
native land forms of life still exist in North America.
What was the impact? Well, we don’t have native sweet
chestnuts to roast nor those gigantic smelly heaps of
guano. Greenberg gives some specific hypothetical sce-
narios resulting from pigeon extinction, one being the
rapid decline and now near extinction of the American
Burying Beetle (a species worth its own book), and
another being the spread of Lyme disease. Lyme carry-
ing ticks are fed largely by Peromyscus mice which
undergo rapid population increases in high mast years.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Greenberg suggests that billions of Passenger Pigeons
would stem these mouse peaks and in turn keep tick
populations “in check”. But on the other hand, mice cal
gypsy moths, so Passenger Pigeons competing with
mice would lead to moth outbreaks, loss of mast trees
and lower Passenger Pigeon numbers. And so such fun
speculation goes. The only certainty is that most North
Americans have never noticed or thought about these
impacts at all.

Well never mind hypothetical ecological roles; how
is it that we know so little about the biology and be-
haviour of Passenger Pigeons beyond the anecdotal and
mostly dubious anecdotal at that. At times reading de-
scriptions of the flocks sounding like a “1000 threshing
machines...plus 1000 locomotives running full throttle
in a covered bridge” , makes one thinks of those low
budget movies, called ‘invasion of species X°. Let X =
Passenger Pigeon. There were lots of pigeons and many
people recorded that fact, but there, apparently, seri-
ous interest died, except to Hunt them, Kill them, Eat
them, Sell them, Market them, Use them as Stoolies,
and Marvel at their indestructible abundance. In the
1800’s, there were few resources to be wasted studying
birds, especially destructive pests. No one investigated
their ecology, life history, or behaviour, beyond what
was practical knowledge to assist hunting. We can be
absolutely certain that no one was trying to conserve or
protect them until it was much too late. There was no
need, because it was beyond the realm of possibility
that they could be exterminated. There are no known
photos of Passenger Pigeons in the wild, although there
are 2 of wild birds being used as stool pigeons. There
is only one known photo of a squab (young pigeon).
Even the number of eggs in a clutch was a mystery with
the two greatest ornithologists of the time, Alexander
Wilson and John James Audubon, feuding over whether
it was one or two. Apparently, it didn’t occur to them
to look. (Greenberg tells us that it was 1, as depicted on
the book’s cover).

How have attitudes changed since the haleyon days
of Passenger Pigeon hordes? This strikes me as a use-
ful question, and one not really addressed by Green-
berg. Before I read this book, and more correctly, before
I read other sources, I never had much doubt that the
extinction of the Passenger Pigeon was caused by hu-
mans, not just humans, but humans slaughtering the
Passenger Pigeon in market hunting. My experience in
the species at risk game has however taught me that
when there are large numbers of a species, there are
many skeptics who believe that over harvesting cannot
extinguish a species. Think of Northern Cod, Bison.
Eskimo Curlew, and many others. Even after intense
harvest seems to have driven a species to the brink,
there are those, particularly those doing the driving,
who argue that either it is not at the brink, because they
are still abundant, or if they are no longer abundant it
was not the harvesters who caused the problem. The
cod is a good example of denial from fisher persons,

2014

and the blame being shifted to seals, foreigners, mys-
terious changes in the water temperature, or that the
cod have moved elsewhere (a special case of blaming
the victim). Similarly, most 19" 20" or 21" century
accounts of the decline of Passenger Pigeons seem
determined to find something more complex than mere
slaughter by pigeon hunters. What is delicious about
these debates is that the arguments remain so self-
servingly constant. In summary the argument goes like
this: whatever one thinks, the species is not in danger
of extinction, but if it is, then it is not us that caused the
problem. Climate change is similar. It is not occurring,
the measurements are suspect, they are measuring the
wrong things, but if it is true, then it is not man made.
I like Greenberg’s summary for his bird, “Homo sapi-
ens slaughtered the bird methodically and relentlessly.
Most everything else is a matter of speculation”

Denial of Abundance

I think there are at least two significant ways that
people express skepticism of the Passenger Pigeon
story, that are reflected generally today when there is
doubt that an “abundant” species is ‘truly at risk’. The
first and most fundamental is the claim that there never
were that many Passenger Pigeons. This skepticism
is not surprising given the hyperbolic nature of the
descriptions and the paucity of serious effort and abil-
ity to quantify numbers. Dismissal of the estimates of
billions is easy when one can blame the errors on uned-
ucated, credulous bumpkins from the distant past. And
others disparage a few billion, pointing out that there
are a lot more chickens today, so what is the big deal.
True, there are about 20 billion chickens globally, and
7 billion are slaughtered annually in North America.
And come to that, there are over 7 billion of us and
we are a much bigger deal than mere pigeons. Green-
berg reluctantly reports a more politically correct skep-
ticism expressed recently that the species was natu-
rally uncommon historically because Native Peoples
somehow kept their numbers down by hunting and
competition. Then when the Europeans arrived and
decimated Native populations, the pigeons exploded
and for a brief period dominated the skies. There is
no support for this odd argument.

In a recent paper in the Proceedings of the National
Academy of Sciences (Hung ef al. 2014; pages 1-6,
PNAS Early Edition), a research team reported that
the effective population size was much smaller, (on
the order of 130,000-24 million), than the 3-5 billion
usually indicated. The PNAS team based this astound-
ing conclusion on analysis of the partial genome of 5
Passenger Pigeons from museum collections and from
“ecological analysis”. The latter led the team to sug-
gest that Passenger Pigeons underwent inherent, fre-
quent, severe fluctuations in population size, and had
a population biology like that of lemmings, voles, or
even locusts, driven by the periodicity of mast (nuts).
So, in a nutshell (sic) population lows, combined with
hunting and loss of habitat caused the great demise.

Book REVIEWS

301

The utter lack of similarity between Passenger Pigeon
(or Band-tailed Pigeon) and lemming/vole life history,
demography, and food habits was never considered in
the analyses, or at least not mentioned. No one has re-
ported Passenger Pigeons, or any pigeon or perhaps
any bird, with a lemming-like population cycle. It is
interesting that they note that Homo sapiens has an
even smaller effective population size (90 000-170 000)
despite its current robust abundance. The team notes
that humans achieved this difference by recently start-
ing with a small population and a rapid recent rise,
whereas their “ecological analyses” support a Passen-
ger Pigeon history of repeated rapid rises and falls in
abundance over the past millions of years. It appears
that this model assumes that pigeon carrying capacity
depended on annual acorn production. It is true of
course that oaks and beech may fluctuate their mast
crop dramatically, but not so true that the different
species do so neither in synchrony, nor in synchrony
with American Chestnut which fluctuated much less
than oaks, and which dominated eastern forests until
after the demise of the Passenger Pigeon. The notion
that Passenger Pigeons underwent fluctuations might
be true, but that the evidence and basis for such is sadly
lacking or misrepresented in this paper, which has no
reference to Greenberg’s book or to the known biology
of Band-tailed Pigeons, Passenger Pigeons or lem-
mings.

Denial of Decline

Before the arrival of Europeans, natives hunted the
Passenger Pigeon extensively. It is perhaps the most
common species in native middens throughout the
eastern half of the continent and presumably was a
dietary staple. This interaction continued with the early
settlers. When the birds came, everyone turned out to
stock up on meat, eggs, oil (from the fat squabs), and
feathers. They didn’t make a much of a dent in their
numbers, as far as anyone could tell (but who was
counting?) There were not that many people and it was
hard to preserve the birds, so the early impacts may
not have been large, like the impacts of fishing on the
Grand Banks until the factory ships came. In the early
days of Native or Settler harvest, cod, bison, Passenger
Pigeons and their fellow teeming species were not
greatly affected. As the later stages of this tale proceed,
we learn that even when the declines became noticed,
it was more often than not assumed that the birds were
not declining, but merely hiding in remote pigeon
secure zones in the far west of the US with the Dalton
Gang, or in Argentina, or had crossed the oceans to
Europe or Asia. My favourite explanation was present-
ed by Cotton Mather, he of the Salem witch trials, who
argued that the birds “migrated to an undiscovered
satellite, accompanying the earth at a near distance”.
One hears similar arguments today about declining
species, ‘they have moved to where you scientists
can’t find them’, or ‘we haven’t searched thoroughly
enough’.

Denial of Humans as Cause

In the past month, I have watched an interesting video
(http://www. youtube.com/watch?v=FpXkA-BY3YE)
of a PhD talk suggesting that hunting helped to cause
extinction, but loss and fragmentation of habitat was
the real culprit. Of course, this doesn’t absolve humans
from causal agency, but does suggest that we were
only guilty of second degree extinction or maybe neg-
ligent speciescide. We didn’t intend to destroy the spe-
cies and there was no malice or intent, we were only

a little stupid. In this video, the student presents one of

her “models” and suggests that this particular model
indicates that if in the mid 19" century, conservation-
ists had applied today’s IUCN (or COSEWIC) criteria,
Passenger Pigeon would have come out Threatened.
That was interesting. But her model also showed that
hunting was not enough, there had to be a major impact
from habitat loss. One of her assumptions built into
her models was that hunting pressure decreased with
declining Passenger Pigeon numbers, whereas Green-
berg’s painstaking descriptions lead to the opposite
conclusion.

I originally found out about this book through a
thought provoking review in the New Yorker (Rosen
2014). In the review, Rosen also suggests that Green-
berg is biased in his fingering over harvest as the cause
of decline. Rosen makes a confusing, to me at least,
argument that hunters were the true conservationists
and that rural subsistence hunters in regions of eco-
nomic hardship often overlap with areas that harbour
species at risk. I think this means that if people were
to blame, well their sins were justifiable. What Rosen
finds less justifiable are “elitists” who conserve vast
tracts of wilderness for the public good without a broad
consensus. Greenberg doesn’t seem to anticipate these
sorts of arguments, nor Rosen’s expansion on the
book’s mention of racism in the writings of a scien-
tist who dissected Martha the last Passenger Pigeon.
It is of course these digressions by Rosen that made
the review interesting.

The Final Days

In this era of climate change and the inevitable de-
niers, there were Passenger Pigeon extinction/threat
/decline deniers. The deniers rested their position on
the assertion that humans could not possibly extermi-
nate such an abundant, prolific species, just as we have
later argued that we could not possibly decimate North-
ern Cod, or the myriad other species we have decimat-
ed. The story is startling for its monotonous repetitive-
ness. First, no one thinks about risk at all, then there are
feeble concerns expressed that decline is occurring,
then denial combined with often bizarre explanations
to refute the declines (they have moved, there are still
lots of them, there are more than you think, jobs and
subsistence food are at stake), then frantic, sometimes
futile efforts to “save” the species.

I agree with Rosen that Greenberg is quite ready to
blame the less noble properties of humanity for the

THE CANADIAN FIELD-NATURALIST

Vol. 128

demise of this splendid creature. But I think Greenberg
has a case. After providing a picture of the bird’s num-
bers and habits, he embarks on a seemingly endless
account of how they were killed. Nets, bigger nets,
shotguns, rifles, pistols, slingshots, air guns, arrows,
clubs, fire, saws (cutting down nest trees), cannons and
potatoes (this latter only in Orillia, Ontario). In the 19th
century, 3 things changed the playing field for the Pas-
senger Pigeon to a near vertical tilt. The telegraph, the
railway and the refrigerated rail car. The wandering
habits of the Passenger Pigeon now could be tracked
via the telegraph, and a huge number of birds could be
killed in a short time, preserved and shipped to big city
markets for profit. Then ensued relentless and techni-
cally improved hunting, the volume of which Green-
berg relates with gusto. The story of the fur trade, the
feather trade, the timber trade and the fish trade was
repeated here. A market was created with little regu-
lation, then the harvest began to reduce the numbers,
this led to more hunters per bird than before, especially
as more hunters got involved to take advantage of the
growing market. The birds helped by being highly
social allowing large numbers to be killed on the breed-
ing ground after being located by via a telegraph net-
work. The intense hunting pressure further disrupted
breeding and suddenly they were nearly gone, the last
big flocks being decimated in final harvest frenzy, and
then the trickling remnants hunted casually for dinner
or specimens. For the folks who think we couldn’t push
them to zero, think again, rationally. In other areas, con-
servation measures and primitive wildlife management
saved many once abundant widespread species before
it was too late; the goose, the beaver, the fisher, the
white pine, and for a while the northern cod, but for
the Passenger Pigeon, the bison, the Eskimo Curlew
species that were highly clumped at some point in their
life history, conservation enlightenment was too late.
To illustrate this “too little too late’, Greenberg has sev-
eral examples of measures to save the Passenger Pigeon
that were so late, they seem humourous. Ontario is a
typical example. An 1887 Act that protected small
birds excluded Passenger Pigeons. A decade later the
province gave the pigeon protection, 8 years after the
last known specimen was taken in the province. Often
the regulations passed in various jurisdictions pro-
tected the hunters’ rights not the birds’ lives. Nothing
changes; can anyone say Ontario Snapping Turtle.

Raising the Dead

One can’t speculate about the Passenger Pigeon and
its collapse without mention of “de-extinction”, Rapid
advances in genetic tech have beckoned the bio-entre-
preneurs out of their basements smelling the colour of
money and the lure of fame. TED (Technology, Enter-
tainment, Design) italks abound as speakers ask the
rhetorical question, “wouldn’t it be fabulous to see ex-
tinct species return to life?” And of course. Passenger
Pigeon sans guano is one of the most popular candi-
dates. Greenberg spends little space on this, briefly

2014

mentioning cloning in a neutral way. | will be less neu-
tral. De-extinction is in general a foolish idea, and to
quote Stuart Pimm, “a spectacular waste of everyone’s
time”. The idea that we can erase the losses of the
| 000°s of species going extinct right now by resurrect-
ing the lost is ludicrous. Jurassic Park shows what is
really involved, hucksterism. In the case of the poor
Passenger Pigeon, the current hype suggests using the
extant Band-tailed Pigeon as the ‘host’ for Passenger
Pigeon DNA. The irony here is that the Band-tailed
Pigeon is itself a species at risk indicating we can’t
even maintain it much less some chimera cobbled
together by biotechnology. Of sad double irony is that
while people generally get excited to hear of restoring
the glory of abundant Passenger Pigeons, they have
no idea what a Band-tailed Pigeon is, and if it goes ex-
tinct, | am supremely confident there will be no enthu-
siasm to de-extinct it. Metaphorically, this effort is like
ignoring the wounded in a battle and trying to resurrect
the dead. It would be best to wait for the end times
then call on the guy who did it 2 millennia ago.

At the end of my Passenger Pigeon odyssey, I offer
this thought. It struck me one day that the Passenger

BooK REVIEWS

303

Pigeon in its heyday was a lot like us. Think of them
and us as sharing a Darwinian background, being mold-
ed by overproduction, genetic variation and natural
selection. There were 5 billion of them, and there are
7 billion of us. Both are hugely destructive patch dis-
turbers, although we do it on a grossly larger scale
befitting our greater intelligence, bulk and numbers.
We both have limited genetic variation and much small-
er effective population sizes. We are both slow breed-
ers, and long-lived. Both of us are omnivorous and
highly social, and we both use Tweets. I won’t mention
guano again. My point here is not that we are going
extinct, but simply that if we think we are a curse on
the earth, then so were Passenger Pigeons, even if a
more modest one. If we think that their destructiveness
and fabled abundance were wonderful, then we can’t
complain about humans. Nature is such a bitch, and
this book will get you thinking about life and biodi-
versity and us...

RON BROOKS
50 Stone Road East, Guelph, ON, Canada, NIG 2W1

A Sparrowhawk’s Lament — How British Breeding Birds of Prey Are Faring

By David Cobham. 2014. Princeton University Press, 41 William Street, Princeton, NJ, USA, 08540-5237. 256 pages,

35.00 USD, Cloth.

There are 15 species of breeding hawks in the UK
and I have seen them all, but only 13 in Britain. Ex-
ploring the plains of southern England my compan-
ion and I spotted a Montague’s Harrier. As it was fly-
ing away we used the leap-frog technique (one person
watching with other running to keep with the bird.
When the runner stopped and had the bird in sight the
second person became the runner). After a few ex-
changes we were close to the harrier and I had just
caught up with my friend when I saw he was about to
sit on an Adder (a venomous snake). | pushed him side-
ways and in the confusion we lost the bird, but gained
a story. As a teenager I was wandering through Scot-
land with a good friend when we saw a large raptor.
We were very excited when we realised it was an
Osprey. The next day the reserve warden insisted we
took a vow of secrecy to protect the eggs from col-
lectors.

The Osprey epitomises the history of diurnal birds
of prey in the British Isles. The over-rich persecuted
these birds that interfered with their “sport” and added
them to their egg and skin collections, so the Osprey
was extirpated by 1916. It did not recolonise until 1954.
Even then their recovery was hampered by the effects
of organochlorine pesticides and the continual predation
by egg collectors. A huge effort by many concerned
people have resulted in a current population of 250
pairs. Compare this to an estimated 1500 nests and in-
creasing in Ontario (2001).When travelling with some

birders in Assam I was mildly reproached for not show-
ing wild enthusiasm for the daily Osprey sighting. Most
summer days I am in the field I see half a dozen or
more birds as I live in ideal habitat.

David Cobham describes the status of each of the
15 breeding, diurnal raptors in turn (Red Kite, White
tailed Eagle, Western Marsh Harrier, Montagu’s Har-
rier, Hen Harrier, Northern Goshawk

Eurasian Sparrowhawk, Common Buzzard, Euro-
pean Honey Buzzard, Golden Eagle, Osprey, Common
Kestrel, Merlin, Eurasian Hobby and Peregrine). His
descriptions are detailed, accurate and reveal that he
is a careful observer. He often includes large chunks of
someone else’s text to support his own narrative. He
does not need to do this as his writing is most readable.
However these inserts are also well written and fit with
the flow of information. The author is primarily a film
and TV producer and vice-president of the Hawk and
Owl Trust. This film-maker “bias” comes through from
time to time, most noticeably when he details filming
a Goshawks nest in the chapter on Honey Buzzard.

The author’s descriptions and his personal anecdotes
are charming and a pleasure to read, but the real mes-
sage of the book is in the history. While all these birds
go though many up and downs, seven have become
good news stories, while three are still very sad. The
remaining five are satisfactory, but need careful moni-
toring.

304

Of the good tales the Red Kite is representative.
These birds were prevalent in the Middle Ages, but
by the 1900s they were only a few pairs in Wales.
With the efforts of volunteers and the reintroduction
of Swedish birds the author can now report there are
now 1,600 pairs in several locations. The Hobby story
is similar going from a low of 100 pairs to its current
2800 pairs. The Peregrine had a little different fate.
Before World War II there was about 700 pairs, despite
being subject to persecution. In the war several hun-
dred were killed to protect military pigeons carrying
messages. The population did recovered but plunged
again to 400 pairs from effects of DDT. Now the au-
thor reports there are 1,500 pairs. The star story is the
reintroduction of the White-tailed Eagle. This common
bird was extirpated from the UK by 1917. From the
release of the first birds in 1975 the population has
grown to 64 pairs.

The Golden Eagle has 440 pairs, a stable popula-
tion size for many years (for comparison the eastern
Canadian population is only 200-300 pairs). This is
similar to the history of Honey Buzzard and Goshawk.
More distressing 1s the history of the two harriers, Hen
and Montagu’s.

Wildlife of the Caribbean

THE CANADIAN FIELD-NATURALIST

Vol. 128

Although the author reports 630 pairs of Hen Har-
rier for all the UK it is likely they are almost extirpat-
ed in England, although the RSPB reports three nests
this year (up from zero in 2013). Raptors are still per-
secuted despite penalties of a six-month jail sentence
or a £5,000 fine. In a 2007 infamous incident two birds
were shot by someone at Sandringham, the Queen’s
estate. A hunting party composed of Prince Harry, a
family friend and the estate gamekeeper were ques-
tioned, but nobody was prosecuted. In this atmosphere
it is not surprising that in the most bird-oriented country
(they have over ten times more members per capita in
the RSPB than we have in Nature Canada) raptors are
still being slaughtered.

David Cobham has written a very understandable
biology and history of British birds of prey. It was a
pleasure to read the words, but the content was, of
necessity, sometimes disturbing. I would recommend
this book to all who like birds, particularly raptors.
British birders and those who visit (like me) will gain
a lot of valuable information. It would make a great
present to anyone studying hawks.

Roy JOHN
2193 Emard Crescent, Ottawa, ON, Canada, K1J 6K5

By Herbert A. Raffaele, and James W. Wiley. 2014. Princeton University Press, 41 William Street, Princeton, NJ, USA,

08540-5237. 304 pages, 19.95 USD, Paper.

There must be a couple of dozen companies offer-
ing luxurious Caribbean cruises. Each stops for a day
or two on a selection of islands. If you choose not to go
on the cruise line’s shopping trip, you can explore the
some of the wild areas. You can usually find a local
birder to help you out.

This guide focuses on 451 species of plants, birds,
mammals, herptiles, fish and seashells that you are most
likely to see on such a trip. It starts with photographs
and descriptions of about 70 widespread plants. The
authors have included both native and many introduced,
but dramatically beautiful, species. So the reader will
find the (African) Tamarind, used to make the much-
loved candy balls, an island favourite. Another fruit
tree, this time a native, is the Guava famed for its jelly
and juice. The traditional Christmas plant, the Poinset-
tia, has been introduced on many islands, but it is not
the familiar pot plant. In the tropical climate it grows
to a three to four metre-high tree. Perhaps the most
exotic flower is the widespread and dazzling Yellow
Bell or Golden Trumpet. Sadly little of the original
Caribbean vegetation now exists.

The section on birds, the most extensive part of the
guide, follows a conventional field guide style. The
illustrations are expertly painted by artists Kristin
Williams and Nils Navarro. Generally there are only
three to five species per page, giving plenty of room
for well-sized depictions. The authors state that there

aim was “to include conspicuous and widespread spe-
cies” so it is reasonable that only 32 percent of the
regions birds are covered (179 out of 564). (For a more
extensive coverage the ardent birder will need “A
Guide to the Birds of the West Indies, by H. Raffeale,
Wiley, Garrida, Keith, and J. Raffeale) The description
of each species is necessarily brief, but useful. I thought
that Status and Range would be the most helpful in
the field.

I was delighted to see a section on reptiles and am-
phibians. It is always much more difficult to find infor-
mation on these animals. On a typical visit I always
see a few lizards or a turtle or two and have a hard time
identifying them. | am sure I normally see only the
commonest species.

The authors include two drab, freshwater fish and
a shrimp. I do not think I paid any attention to look-
ing underwater in streams. Maybe I should. A similar
section on marine fish is more extensive, covering 89
species. As most of these are either highly coloured or
dramatically large they do claim the tourist’s attention.
Even without snorkelling I managed to identify 10 of
these exotic fish, nine of which are in this book. Oddly
missing is the common Caribbean Silversides.

There are smaller sections on marine and land mam-
mals, reptiles and amphibians, butterflies, land crabs
and snails. A marine segment, as well as fish, covers

2014

corals, shells and marine invertebrates. These give sim-
ilar coverage to the larger divisions f

So how well do they succeed, considering the au-
thor’s aim to provide a general nature book of com-
monly-seen species? I estimate in a couple of trips to
the area I saw 85 percent of the birds covered in the
book, if I exclude the 68 endemics on islands I did not
visit. Because | made a special effort (hiring a local
guide) | saw six rarities not in the book (e.g. Grenada
Dove, Eurasian Little Egret). I believe I saw approxi-
mately 70 percent of the plants depicted. These are
very good percentages and suggest the authors have
succeeded well.

There are a couple of minor points I noticed. There
is no mention of Cayenne Tern and I saw a number of
the yellow-billed birds. They are closely related to the
Sandwich Tern and are probably the same species.
However the taxonomic relationship of these terns is
still unresolved. The authors include the St. Vincent

BOTANY

Book REVIEWS

305

Parrot (population about 700 plus) and the nocturnal
Cuban Solenodon (population low and unknown). Yet
they did not include the iconic Grenada Dove (popu-
lation over 130).

This small (8 * 5 x % inches) book slips easily into
my pants pocket so it is ideal for the travelling natu-
ralist. It covers most of what you will see wandering
the Caribbean. The passionate birder will want to take
Raffaele et al.'s other book mentioned above. While
you are en route in your comfortable cabin you should
read the introductory section which is a good discus-
sion of the Caribbean’s wildlife. The author’s give an
account of the devastation caused by humans and the
efforts being made by some of those humans to retain
what is left of this natural heritage.

Roy JOHN
2193 Emard Crescent, Ottawa, ON, Canada, K1J 6K5

Plants of Southern Ontario, Trees, Shrubs, Wildflowers, Grasses, Ferns and Aquatic Plants

By Richard Dickinson, and France Royer. 2014. Lone Pine Publishing, 87 Pender Street East, Vancouver, BC, Canada, V6A

1S9. 527 pages, 29.95 CDN and USD, Paper.

Lone Pine has produced 33 botanical guides mostly
covering North America. Five of these specifically
cover Ontario: “Trees of Ontario”, “Wetland plants of
Ontario”, “Ontario Wildflowers’, “Forest plants of cen-
tral Ontario”, and “Forest plants of northeastern On-
tario”. There is also the ROM field to wildflowers
(Dickinson et al. 2004), partly authored by Richard
Dickinson. Thus, there are plant groups and regions
that are covered more completely in earlier guides by
the same publisher and by one of the same authors. For
example, Linda Kershaw’s “Trees of Ontario” covers
213 species and the guide considered here covers 90.
There are also many other guides that cover the vari-
ous groups in this new book more completely, but it is
sometimes convenient to start with a book that has a
little of everything. Because it is incomplete however,
it does not provide an accurate identification much of
the time, although the user may get near to the correct
identification and then use more complete books to
go further.

The area covered by this guide includes two large
ecoregions of southern Ontario, the Southern Decid-
uous Forest (also known as the Carolinian Zone) and
the Great Lakes — St. Lawrence Forest. The plants cov-
ered are vascular plants, but although it is implied that
they are wild plants, some plants only found in gardens
are included such as the Maidenhair Tree (Gingko bilo-
ba). When a guide is incomplete, the choice of what to
include becomes a challenge. It seems best to include
what people are most likely to encounter. In this guide
the choices have not always been the best. For exam-

ple, Black Pine, which is not included, is more often
seen in plantings and as an escape than Douglas-fir,
which is included. The less frequent escape Euonymus
europaeus is included while the more common, con-
spicuous, and native Celastrus scandens (in the same
family) is not.

It is also helpful in an incomplete guide to mention
related species that are not covered in depth and there
are many places in this guide where that could have
been done. On p. 245 under the text on Sundew Fam-
ily, there is ample space to say that “four species occur
in Ontario, one with round leaves, one with linear
leaves (both shown here) and two with spoon-shaped
leaves one of which has smooth leaf stalks (D. inter-
media), the other with more or less glandular hairy
leaf stalks (D. anglica). Likewise for Claytonia there
is ample space to refer to the more northern Carolina
Spring Beauty (Claytonia caroliniana) which has
leaves less than eight times as long as broad com-
pared to the Eastern Spring Beauty (Claytonia vir-
ginica) which has leaves more than eight times as
long as broad.

It is not clear what the authority is for the scientif-
ic and common names, but for the scientific names at
least it appears to be ITIS (Integrated Taxonomic Infor-
mation System as noted on page 8). For both kinds of
names, VASCAN (http://data.canadensys.net/vascan
/taxon/6949?lang=en) may be the best choice. Here
current scientific names are available and an attempt
is made to standardize common names. For example
Spiranthes romazoffiana is “Hooded Ladies’ -tresses”

306

rather than just “Ladies’-tresses”. “Zigadenus elegans”
is “Anticlea elegans”. “Carex eburnea” is “Bristle-
leaved Sedge” rather than “Bristle-leaf Sedge”. “Black-
grass Rush” should be “Black-grass Rush”. The “Dwarf
Lake Primrose” should be the “Mistassini Primrose”.
“Pale Coralroot” should be “Early Coralroot’, and etc.

The misidentified photos are disappointing in an
identification guide. Following 1s a list of those I found
(in the pre-distribution copy) including page numbers
with incorrect name of photo followed by the correct
name: 180 right, Coeloglossum viride is Platanthera
hyperborea; 187, Platanthera hyperborea is Platan-
thera dilatata; 421, Elodea canadensis is Najas guada-
lupensis; 455, Carex aurea is Carex garberi; 459,
Carex lasiocarpa 1s Carex aquatilis or C. stricta; 475,
Juncus gerardii is Juncus secundus; 486, Selaginella
rupestris 18 Selaginella eclipes; and 488, Botrychium
multifidum 1s Botrychium lanceolatum.

OTHER

Bird Lady — A Lifelong Love Affair with Birds

THE CANADIAN FIELD-NATURALIST

Vol. 128

There are also mistakes in the descriptions. For
example the flowers of Cypripedium orchids have three
stamens, one of which is sterile and two of which are
reduced but fertile. This guide describes three of them
as having a single stamen.

Although this book cannot be recommended with-
out some reservation, and does not come close to the
remarkably high standard of quality in the Lone Pine
guides for BC produced by Jim Pojar and Andy Mac-
Kinnon, it is still a substantial achievement. It includes
528 pages, 760 species, and many impressive photo-
graphs (although some are dark). It also provides a
useful overview of the wild flora of southern Ontario
which includes approx. 3000 species. It is field guide
size and stitch-bound with good quality paper. It will
stand up well to outdoor use and is good value for the
price.

PAUL CATLING

William Saunders Bldg., Central Experimental Farm, Ottawa,
ON, Canada, K1A 0C6

By Elizabeth Le Geyt. 2014. Friesen Press Inc., Suite 300 — 852, Fort Street, Victoria, BC, Canada, V8W 1H8. 184 pages.

18.57 CAD, Paper.

Elizabeth Le Geyt is an Ottawa icon. Long before
the internet and emails she wrote a column about birds
in the Ottawa Citizen’s which lasted for 39 years.
Using information phoned in by local bird watchers
she reported what birds were where and what they
were doing. This enabled her readers to make informed
decisions on where to go on their own field trips. This
gained her an extensive and loyal local following. Un-
fortunately the column included some bizarre gaffs that
irritated the truly passionate birders. Names would get
mixed up, for example Broad-shouldered Hawk. Did
this mean a Broad-winged Hawk or a Red-shouldered
Hawk? Sometimes it would be the numbers that were
corrupted. Was there one Snow Bunting with a flock
of 250 Snowy Owls or was it really the other way
round? So I was not surprised when I read of her joy
as a teenager at watching a heronry of Great Blue
Herons near her home at Bothwell. The problem is
Bothwell is near Glasgow, Scotland, and there is only
one record of this North American species in the UK
(St Mary’s, Isles of Scilly, 7 December 2007). Clearly
the author means the Grey Heron (13,000 nests and
about 63,000 birds according to the RSPB). The black-
and-white photo of a heron’s nest that follows, is
labelled as Great Blue and I think this is correct. You
cannot see the colour of the thighs, but they look grub-
by, not white.

But this is not a time to pick at the ornithological
details in this book. It is the tale of one woman’s pas-
sion for nature and, in particular, birds. The author

recounts her youth in Great Britain and her joy at dis-
covering its many birds. You can follow her journey to
adulthood and then her marriage and resultant family.
Her husband a Royal Navy officer is posted to Canada;
first Halifax then Ottawa. This gives Le Geyt a new
group of birds to learn and enjoy.

She is asked to take over the bird column in the
Ottawa Citizen. This brings her a whole new coterie of
contacts and friends. Her caring nature leads people
to bring her orphaned and wounded wildlife. Eventu-
ally she forms a liaison with the Wild Bird Care Centre
where injured animals are treated and rehabilitated
and released back into a typical habitat. It is question-
able whether a hand-reared creature placed in the wild
would fare well. If the habitat is ideal it likely has a
resident occupier, who will drive any newcomer away.
Eventually it will be driven to poor habitat where it will
be vulnerable to starvation and predation. However the
process of caring for the distressed and helpless crea-
tures has a powerful message to the untutored public.
Le Geyt and her friends have raised the awareness of
the plight of wildlife and the environment and much
of this has been achieved through their rescue efforts.

The author gives a concise portrayal of the impor-
tant wildlife areas of the Ottawa area. She describes
her adventures with Jacko, an African Grey Parrot, Joey
the Rock Pigeon; and Rattles the kingfisher. She also
reports on trips she took to South Africa, Mexico. Costa
Rica, Trinidad, Arizona and her return to Great Britain.

2014

Elizabeth Le Geyt is a good writer who uses a light,
breezy style to bring the reader into her passion. Her
tales are told with warmth and humour. | really enjoyed
learning of her childhood and comparing it to my own.
The wonder she shows at seeing her first Canadian
birds is something else I can share. When she describes
the exploits of Jacko, Joey and Rattles her writing
reflects her own warm and kindly nature. Her journeys
to other countries again reminds me of my own travels.

The Dismal State of the Great Lakes

Book REVIEWS

307

| think all those readers who followed Le Geyt’s
column will delight in this book. Any one who loves
nature will enjoy this light-hearted book. The reader
will feel relaxed and uplifted with every chapter, so
perhaps even non-naturalists should buy a copy — it
will improve their environmental awareness. This de-
lightful book is a remarkable achievement for a woman
who is over 100 years old.

Roy JOHN
2193 Emard Crescent, Ottawa, ON, Canada, K1J 6K5

By James P. Ludwig. 2013. Xlibris. Orders@Xlibris.com. Telephone 1-888-795-4274. 273 pages. Paperback 19.99 USD;
hardcover 29.99 USD; e-book 3.99 USD. Printed on demand.

The Laurentian Great Lakes of North America con-
tain 18 percent of the planet’s fresh surface water and
90 percent of the United States’ supply. The Great
Lakes region has been the major manufacturing hub
of North America, and has supplied vast quantities of
timber and minerals, particularly iron ore, to the world’s
markets. Today, agriculture, recreation and tourism,
fishing, and shipping are the important industries. The
Boundary Waters Treaty of 1909 between the govern-
ments of Canada and the United States established an
International Joint Commission and mechanisms for
resolving any disputes over waters bordering the two
countries, as they pertained to their use for domestic
and sanitary purposes, navigation, power, and irriga-
tion. The Great Lakes Water Quality Agreement of
1972/78 committed the Parties to cooperation in the
restoration and maintenance of the chemical, physi-
cal, and biological integrity of the waters of the Great
Lakes Ecosystem. These legislative tools attest to the
tremendous importance of the Great Lakes to Canada
and the United States and to planet Earth. Unfortunate-
ly, the scientific findings documenting the demise of
the Great Lakes, and the political failure to address their
implications are not well-known outside the Great
Lakes community. The Dismal State of the Great Lakes
is James P. Ludwig’s attempt to remedy that by detail-
ing his personal and professional journey in studying
the extensive damage that has been done to the Great
Lakes over his lifetime, the once-promising but even-
tual political failure to restore them, and his analysis
of why they happened and how we might fix the mess.

This is a book written by an ageing, angry man with
a fire in his belly for justice and a deep love for these
great “sweetwater seas.” It is “a call to debate the pos-
sible paths to effective change and to reignite the fires
of public involvement.” This is a book with an impor-
tant message.

The Ludwig family have documented the biological
changes in the Great Lakes for over 70 years. Jim Lud-
wig’s father and uncle, both physicians, were the first
of two generations of admittedly “compulsive bird

banders,” who each year visited the colonies of cor-
morants, gulls and terns to place aluminum bands on
the young birds. On these occasions they recorded the
catastrophic effects of a succession of chemicals and
invasive species. Trained in science, they kept good
records, understood the implications of what they were
seeing, and raised the appropriate “red flags.” Jim Lud-
wig was awarded a PhD from the University of Michi-
gan in 1968 for his thesis entitled “Dynamics of Ring-
billed Gull and Caspian Tern populations of the Great
Lakes.” He founded Ecological Research Services, a
small consulting firm, in 1969. Glenn Fox (GAF), who
also devoted his career to the study of fish-eating birds
in the Great Lakes, has collaborated with Jim Lud-
wig directly and indirectly, in research projects. Stu-
art Houston (CSH), a physician and compulsive bird
bander, has corresponded occasionally with the Lud-
wigs, father and son, since the 1950s.

Ludwig’s story consists of several threads. One deals
with the biological information; alien species, eutrophi-
cation, algal toxicity, botulism, changes in the nature,
size, and nutritional quality of the food base, and the
effects of toxic chemicals on the health, reproduction,
and survival of fish-eating birds and mammals, in-
cluding humans. Another deals with the complex and
on-going contamination by toxic chemicals; their de-
tection, identification, sources and environmental be-
haviour, and our efforts to control their release. A third
is the history of the Great Lakes fishery, its manage-
ment, and the conflicted culture of fisheries managers.
The final thread is the socio-political history surround-
ing the Great Lakes Water Quality Agreement and the
International Joint Commission, the successes and fail-
ures of numerous collective actions and Ludwig’s inter-
pretation of what went wrong. He suggests how to get
it right in the future.

Alien species. The Great Lakes have been invaded
by numerous alien species. The sea lamprey, which
infested Lake Ontario via the St. Lawrence River, and
the upper Great Lakes with the completion of the Wel-
land Ship Canal in 1932, greatly impacted Lake Trout

308

(and later introduced salmon) stocks. The Alewife, a
small herring, invaded the Great Lakes by a similar
route and became superabundant. It suffered periodic
die-offs in response to sudden changes in water tem-
perature; extraordinary numbers of their dead and dy-
ing bodies were cast up on beaches. Alewives became
a major food item for Ring-billed Gulls, whose popu-
lations increased tenfold to a million birds. Zebra and
Quagga Mussels, Round Gobies, and the Spiny Water
Flea were among many alien species introduced in
ballast water of ships from the Baltic Sea. Today, the
vast populations of these mussels filter out much of
the phytoplankton, altering energy transfer in the food
web; moribund gobies are eaten by loons and cor-
morants who then die from ingesting botulism toxin
concentrated within the gobies.

Eggshell thinning, the first of many health effects
observed in fish-eating birds. Bald Eagle productivity
dropped drastically due to eggshell thinning caused by
DDE which was widely used by the mid-1940s. The
effects persisted from 1955 until a decade after wide-
spread use of DDT was banned in 1972. In 1972, a
high percentage of cormorant eggs failed to hatch.
None of the eggs in the second clutch hatched that
year. This was attributed to severe DDE-induced egg-
shell thinning [cormorants incubate by standing on their
eggs]. Cormorants and bald eagles were the most af-
fected by DDT, gulls the least.

Crossed bills in cormorant chicks warn of devel-
opmental toxins in Great Lakes fish. In June 1983, on
Little Gull Island in northern Green Bay, Ludwig found
5 of 460 cormorant chicks had crossed bills, the first
he had seen, an unusually high incidence of a devel-
opmental defect. By June 1988, he had collected 41
more cross-billed cormorant chicks, but only 1 from
Canadian colonies. He also saw numerous chicks with
eye, leg and hip deformities. These findings suggested
that biologically significant amounts of developmental
toxins were present in Great Lakes food webs. Inten-
sive collaborative studies involving teams of analyti-
cal chemists and toxicologists from Michigan State
University, Ehime University in Japan, a teratologist
and biologists from the U.S. Fish and Wildlife Serv-
ice and the Canadian Wildlife Service suggested that
nearly 90% of the variability in egg viability among
colonies was explained by TCDD-EQs (tetrachloro-
dibenzo-p-dioxin equivalents) in the eggs alone. The
Michigan Department of Natural Resources initially
encouraged these studies and provided the funding
for the work in 1986-88. However, the arrival of a new
director and strong criticism from MDNR biologists
who were furious over findings released to the public
which could reduce fishing license sales resulted in
withdrawal of MDNR support. Fortunately, Dow Chem-
ical and Upjohn Pharmaceuticals (two companies who
used, made, and discharged a wide variety of chemi-
cals), the McGregor Foundation of Detroit, and Con-
sumers Power Corp stepped up to provide finances to

THE CANADIAN FIELD-NATURALIST

Vol. 128

complete the studies, which provided very strong cause-
effect linkages between these biological effects and
dioxin-like synthetic chemicals.

Cosmos the Cormorant, poster child for toxic chem-
ical regulation. Much public interest and political sup-
port was generated by Cosmos, a cormorant chick with
a severely crossed bill that Ludwig rescued from an
island in northern Lake Michigan in June 1988. Lud-
wig made himself a promise that he “would do every-
thing possible to keep Cosmos alive, and he would
handle the public relations for the movement to ban
toxic chemicals and clean-up Great Lakes’ Areas of
Concern with high contamination.” Cosmos made 16
TV appearances in the United States and Japan and
testified by her presence in numerous state and federal
legislative committees, and accompanied Ludwig on
27 evening presentations to environmental and teach-
ers groups in five of the Great Lakes states. Ludwig
writes “I think it safe to claim Cosmos in her single
short year of life did more to propel the issue of toxic
chemicals to the forefront of political agendas in Michi-
gan and Wisconsin than all the other environmental
activists and scientists were able to achieve through
lobbying or explaining their findings.” Cosmos’s lega-
cy lives on in Japan where she has become the symbol
of the dangers of unregulated toxic chemical use. She
was the cover image on the fifth grade textbook on pol-
lution and climate change that is used in all Japanese
public schools.

Bird Banding reveals population level impacts of
chemical contaminants. Ludwig’s family began band-
ing Caspian Terns and other colonial waterbirds in
1922 and continued to do so until 1995. Jim periodi-
cally trapped nesting adults in some of these same
colonies 1966-1992. By 2008, he had amassed 7000
encounter records of Caspian Terns banded as chicks
in Great Lakes colonies. This immense and unique data
set has proven very useful. As early as 1971, he found
that nearly half of adults in Lake Michigan colonies
were hatched in Canadian colonies which was not the
case in 1966 and 1967. When Ludwig analyzed the
7000 encounters, he found, that relative to birds raised
in Canadian colonies, survival to adult age in the most
contaminated colonies in Saginaw Bay and Green Bay
was 20% and 30%, respectively, while that of colonies
with intermediate contamination in northern Lake
Michigan was 59%. This pattern in relative survival to
breeding age was almost identical to that of relative
immunocompetence in chicks of this species that Keith
Grasman, a Ph D student from Virginia Tech meas-
ured in these same colonies 1989-1992. In Saginaw
and Green Bay colonies the immunocompetence of
chicks was less than 50% of that in the cleaner Cana-
dian colonies in Lakes Huron and Superior and was
inversely correlated with TCDD-EQs in the eggs from
these colonies. Caspian Terns banded as nesting adults
in Canadian colonies survived twice as long as adults
banded in Saginaw Bay colonies, 5 years longer than

2014

those from Green Bay colonies, and 4.2 years longer
than those from northern Lake Michigan colonies. Pre-
vious investigations had found a significant negative
correlation between recruitment of terns in these colo-
nies and circulating blood concentrations of total PCBs.
PCB concentrations in the blood of adults from the
Canadian colonies were half those in blood of terns
nesting in Saginaw and Green Bay colonies. It is not
surprising that Ludwig makes a strong pitch for long-
term studies and adequate funding for investigators.

Toxic chemical contamination. Ludwig does a good
job of describing and explaining the universal role of
energy transfer and bioaccumulation in natural systems
and how trace amounts of synthetic chemicals are sim-
ilarly transferred and bio-accumulated to toxic con-
centrations. A great many persistent toxic chemicals
pollute the waters of the Great Lakes. Mercury, poly-
cyclic aromatic hydrocarbons, the insecticide DDT,
hexachlorobenzene, and the polychlorinated biphenyls
(PCBs) and dioxins have probably been the most im-
portant. Like important amino acids and hormones in
animal bodies, the most prominent synthetic toxic com-
pounds have a backbone of six carbon bipheny!] rings.
Synthetic compounds containing two biphenyl rings
to which chlorine, fluorine, or bromine are attached to
some of their carbon atoms are the compounds that
have caused the greatest damage. Their toxicity varies
by a million-fold with the number and position of the
halogen constituent on the biphenyl rings. The most
potent of these compounds is 2,3,7,8-tetrachloro-
dibenzo-para-dioxin (TCDD). Toxicologists rate the
toxicity of other halogenated aromatic hydrocarbons
relative to TCDD and express the total toxicity of ex-
tracts from tissues and environmental media as TCDD-
Equivalents (TCDD-EQs). They act via a specific
receptor present in cells: the aryl hydrocarbon (AH)
receptor, a transcription factor involved in expression
of genes. TCDD-like compounds alter thyroid function
and other metabolic processes, cause birth defects and
permanent neurological damage in embryos and young
animals, are endocrine disrupters, and immunomodu-
lators. Some or all of these effects have been seen in
fish, amphibians, reptiles, birds, and mammals includ-
ing humans. Much of what is known about the envi-
ronmental effects and dangers of these chemicals has
been learned from investigations conducted in the Great
Lakes. Several studies have shown effects on motor and
behavioural development of infants and lower intelli-
gence quotients of older children of mothers who ate
significant amounts of fish from Lakes Michigan and
Ontario. The severity of these effects was associated
with the mother’s PCB exposure.

The Fishery. Throughout their history, the Great
Lakes have been important for fish, and to people who
depend on them for food, subsistence, income, and
recreation. The relative importance of these activities
has varied over time. Unsustainable commercial fish-
ing activity decimated the Lake Sturgeon in the 1880s

Book REVIEWS

309

and the Blue Pike in the 1960s. Atlantic Salmon pop-
ulations in Lake Ontario were also hurt by overfishing.
However, although not accepted by many fisheries
biologists, there is clear scientific evidence that Lake
Trout populations in Lake Ontario and probably else-
where in the Great Lakes were greatly affected by
TCDD-like chemical contamination. Research pub-
lished in 2003 showed that the extirpation of native
Lake Trout from Lake Ontario during the three decades
following WWII was the result of TCDD-like contami-
nants which killed all embryos of the wild Lake Trout
for over 20 years, extinguishing the stock. Today the
dominant salmonids in the Great Lakes are various
species of introduced Pacific salmon whose popula-
tions are maintained with hatchery stock, and are har-
vested mainly by sport fisherman. Recent estimates
place the value of the sport fishing industry to the
Great Lakes economy at 4 billion dollars, and the com-
mercial fishery at | billion dollars. There is a clear con-
flict between natural resource managers who benefit
from the sale of fishing licenses and public health au-
thorities who are issuing consumption advisories for
fish taken in badly contaminated locations around the
Great Lakes. Ludwig believes this conflict is one of the
reasons that funding for health effects research and
monitoring disappeared, and that a culture of “good
news only” (“One must never allow bad news to be
publicized!”) pervades governments around the Great
Lakes today.

The greatest legacy of the Boundary Waters Act of
1909 was the creation of the binational International
Joint Commission. The Great Lakes Water Quality
Agreement of 1972 committed the Parties to coopera-
tion in the restoration and maintenance of the chemi-
cal, physical, and biological integrity of the waters of
the Great Lakes Ecosystem. For many years the Com-
missioners, Parties, and Great Lakes citizens worked
hard at achieving those objectives, supported by sound
science. However, as public concern increased concern-
ing the health effects of persistent toxic substances, the
Parties adopted a defensive position, limiting support
for research, monitoring and information dissemina-
tion. The International Commission’s work was sub-
verted. The objective of virtual elimination of persist-
ent toxic substances was regarded as too expensive
by successive governments. Hence their policy focus
became a vague, multi-causal, “Ecosystem Approach”
which redirected funding and concern away from the
study of biological effects, remediation, and science-
based policy. The result was a series of regulatory
blunders, avoidance of assigned duties, and far too
much reliance on use of models and simplistic indi-
cators. No surprisingly, the chemicals of concern and
their biological effects persist.

Ludwig’s book includes his 1995 editorial in the

Journal of Great Lakes Research (21:159-160), sug-

gesting that the Science Advisory Board of the Inter-
national Joint Commission had degenerated into a

o

310

means to achieve a politically acceptable consensus.
What else has gone wrong in government? As Al Gore
said, “The private foxes have been put in charge of the
public hen houses.” In the USA, the US Environmen-
tal Protection Agency, created by Richard Nixon in
late 1970 and the Department of the Interior have been
“largely emasculated,” “intellectually corrupted,” and
“a form of ecological insanity” according to Ludwig.
No one and no agency is responsible for management
of the whole resource of the Great Lakes ecosystem;
there is a hodge-podge of funding, too many barriers
to effective communication, incredible redundancy,
very poor accountability, and no clear place for the
public to redress grievances. Sound, evidence-based
science is required but no longer encouraged or sup-
ported. Although the lifetime and multi-generational
needs of the commonweal are far more important than
money accumulation by the individual and corpora-
tions, this is incompatible with the neoliberal politi-
cians of today.

Jim Ludwig is an “insider” who suggests how to rec-
tify the current disastrous situation. Society needs wise
and informed advocates with ecological knowledge,
who are committed to change. We need to discard free-
market policies of neglect, passivity and deception, and
commit to good water quality by restoring powers of
the original Great Lakes Water Quality Agreement and
the International Joint Commission. Critical analyses

THE CANADIAN FIELD-NATURALIST

Vol. 128

must no longer be suppressed and ignored. If only
Ludwig’s facts and insight would direct the authori-
ties to appropriate action!

The Dismal State is correctly titled, but is a disap-
pointment in several ways. Its importance deserved a
University press, careful external editing, better organi-
sation, more headings, an index, a more complete list
of acronyms, and a better binding for the paperback
version (twenty two-sided pages of CSH’s paperback
review copy have already come loose). Sadly, Lud-
wig’s writing will seem too personal and too political
for some tastes. However, he has accurately document-
ed what has gone wrong. If nothing else, his testimony
will remain as a valid historical record.

The Dismal State of the Great Lakes has not yet
achieved the distribution it deserves. I offer a promis-
ing possibility for consideration: that each regional
nature club invest a hundred dollars to help the Great
Lakes’ future. Each could order e-book copies for five
of its members to pass around and donate one copy of
the permanent hardcover version to the adjacent high
school, college or University library. We are doing the
latter. Ludwig’s valuable information assuredly de-
serves to be more widely distributed.

GLEN Fox (GAF)! and C. Stuart Houston (CSH)

'877 Ridley Boulevard., Ottawa, ON, Canada, K2A 3P4
863 University Drive, Saskatoon, SK, Canada, S7N 0J8

2014

NEw TITLES

Prepared by Roy John

r Available for review * Assigned

NEw TITLES 311

Currency Codes — CAD Canadian Dollars, USD US. Dollars, EUR Euros, AUD Australian Dollars.

ZOOLOGY

* North American Amphibians: Distribution and
Diversity. Edited by David M. Green, Linda A. Weir,
Gary S. Casper, and Michael Lannoo. 2014. California-
Princeton Fulfillment Services, 1445 Lower Ferry
Road, Ewing, NJ, USA, 08618. 352 pages, 75.00 USD,
Cloth.

Dam Builders: The Natural History of Beavers and
their Ponds. By Michael Runtz. 2014. Fitzhenry &
Whiteside Limited, 195 Allstate Parkway, Markham,
ON, Canada, L3R 4TS8. 254 pages, 35.00 CAD, Cloth.

The Bee: A Natural History. By Noah Wilson-Rich.
2014. Princeton University Press, 41 William Street,
Princeton, NJ, USA, 08540-5237. 224 pages, 27.95
USD, Cloth.

Phillipps’ Field Guide to the Birds of Borneo:
Sabah, Sarawak, Brunei, and Kalimantan (3"¢ Edi-
tion). By Quentin Phillipps, and Karen Phillipps. 2014.
Princeton University Press, 41 William Street, Prince-
ton, NJ, USA, 08540-5237. 384 pages, 35.00 USD,
Paper.

American Birding Association Field Guide to Birds
of Colorado. By Ted Floyd. 2014. Scott & Nix, Inc.,
150 West 28" Street, Suite 1900, New York, NY, USA,
10001. 320 pages, 19.93 USD, Paper.

American Birding Association Field Guide to Birds
of New Jersey. By Rick Wright. 2014. Scott & Nix,
Inc., 150 West 28th Street, Suite 1900, New York, NY,
USA, 10001. 368 pages, 24.95 USD, Paper.

Common Birds of Nunavut. By Mark Mallory. 2014.
Inhabit Media Inc., P.O. Box 11125, Iqaluit, NU, Cana-
da, XOA 1HO. 32 pages, 19.95 CAD, Paper.

* Illustrated Checklist of the Birds of the World —
Volume 1 (Non-passerines). By Josep del Hoyo, Nigel
Collar, David Christie, Andrew Elliott, and Lincoln
Fishpool. 2014. HBW and BirdLife International, Lynx
Edicions, Montseny, 8, 08193 Bellaterra, Barcelona,
Spain. 904 pages, 185.00 EUR, Cloth.

The World of Birds. By Jonathan Elphick. 2014. Fire-
fly Books Ltd., 50 Staples Avenue, Unit 1, Richmond
Hill, ON, Canada, L4B 0A7. 640 pages, 75.00 CAD,

Cloth.

Prairie Dog Empire: A Saga of the Shortgrass Prai-
rie. By Paul A. Johnsgard. 2014. Barnes and Noble,
Inc., P.O. Box 111, Lyndhurst, NJ, USA, 07071. 243
pages, 22.95 USD, Cloth.

* The Amazing World of Flyingfish. By Steve N.
G. Howell. 2014. Princeton University Press, 41 Wil-
liam Street, Princeton, NJ, USA, 08540-5237. 64 pages,
12.95 USD, Cloth.

The Illustrated Handbook of Fossils: A Practical
Directory and Identification Aid to More Than 300
Plant and Animal Fossils. By Steve Parker. 2014.
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USA, 07071. 160 pages, 18.99 CAD, Cloth.

In Search of Lost Frogs. By Robin Moore. 2014.
Firefly Books Ltd., 50 Staples Avenue, Unit 1, Rich-
mond Hill, ON, Canada, L4B 0A7. 256 pages, 35.00
CAD, Cloth.

Hummingbirds. By Ronald Orenstein. 2014. Firefly
Books Ltd., 50 Staples Avenue, Unit 1, Richmond Hill,
ON, Canada, L4B 0A7. 256 pages, 35.00 CAD, Cloth.

* Life on the Rocks — A Portrait of the American
Mountain Goat. By Bruce Smith. 2014. University
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Boulder, CO, USA, 80303. 176 pages, 34.95 USD,
Cloth.

Hoofed Mammals of British Columbia — 2" Edi-
tion. By David Shackleton. 2013. Royal BC Museum,
675 Belleville Street, Victoria, BC, Canada, V8W 9W2.
272 pages, 25.95 CAD, Paper.

*A Field Guide to the Larger Mammals of Tanzania.
By Charles Foley, Lara Foley, Alex Lobora, Daniela
De Luca, Maurus Msuha, Tim R. B. Davenport, and
Sarah Durant. 2014. Princeton University Press, 41
William Street, Princeton, NJ, USA, 08540-5237. 320
pages, 29.95 USD, Paper.

Marmot Biology — Sociality, Individual Fitness,
and Population Dynamics. By Kenneth B. Armitage.
2014. Cambridge University Press, Shaftesbury Road,
Cambridge, UK, CB2 8BS. 410 pages, 120 USD,
Cloth.

The Passenger Pigeon. By Errol Fuller. 2014. Princeton
University Press, 41 William Street, Princeton, NJ,
USA, 08540-5237. 184 pages, 29.95 USD, Cloth.

* A Feathered River Across the Sky: The Passenger
Pigeon’s Flight to Extinction. By Joel Greenberg.
2014. Bloomsbury Publishing Inc., 1385 Broadway,
5th Floor, New York, NY, USA, 10018. 304 pages,
26.00 USD, Cloth.

*The Fish in the Forest — Salmon and the Web of

Life. By Dale Stokes. 2014. University of California
Press, 155 Grand Avenue, Suite 400, Oakland, CA,
USA, 94612-3758. 172 pages, 29.95 USD, Cloth.

Creatures of the Deep — In Search of the Sea’s
“Monsters” and the World They Live In — 2"4
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ples Avenue, Unit 1, Richmond Hill, ON, Canada,
L4B 0A7. 208 pages, 39.95 CAD, Cloth.

Shrewdunnit: The Nature Files. By Conor Jameson.
2014. Pelagic Publishing Ltd., PO Box 725, Exeter,
UK, EX1 9QU. 300 pages, 14.99 GBP, Cloth.

* A Sparrowhawk’s Lament — How British Breed-
ing Birds of Prey Are Faring. By David Cobham.
2014. Princeton University Press, 41 William Street,
Princeton, NJ, USA, 08540-5237. 256 pages, 35.00
USD, Cloth.

Guide to Minnesota’s Reptiles and Amphibians.
By John J. Moriarty, and Carol D. Hall. 2014. Univer-
sity of Minnesota Press, Suite 290, 111 Third Avenue
South, Minneapolis, MN, USA, 55401. 400 pages,
39/951 CAD, Cloth:

Biology and Conservation of North American Tor-
toises. By D. Rostal, E. McCoy, and H. Mushinsky.
2014. John Hopkins University Press, 2715 N. Charles
Street, Baltimore, MD, USA, 21218. 190 pages, 69.95
USD, Cloth.

The Trilobite Book: A Visual Journey. By Riccar-
do Levi-setti. 2014. The University of Chicago Press,
1427 East 60" Street, Chicago, IL, USA, 60637. 288
pages, 49.30 CAD, Cloth.

Woodpeckers of the World — A Photographic Guide.
By Gerard Gorman. 2014. Firefly Books Ltd., 50
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L4B 0A7. 528 pages, 49.95 CAD, Cloth.

THE CANADIAN FIELD-NATURALIST

Vol. 128

BOTANY

Carnivorous Plants of Australia Magnum Opus
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Carnivorous Plants of Australia Magnum Opus
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Carnivorous Plants of Australia Magnum Opus
Volume 3. By Allen Lowrie. 2014. 61 Redfern Natural
History Productions, Lake Drive, Hamworthy, Poole,
Dorset, UK, BH15 4LR. 463 pages, 34.99 GBP, Cloth.

A Great Lakes Wetland Flora: A Complete Guide to
the Wetland and Aquatic Plants of the Midwest. By
Steve W. Chadde. 2012. Barnes and Noble, Inc., P.O.
Box 111, Lyndhurst, NJ, USA, 07071. 692 pages, 44.86
CAD, Paper.

* Plantes de milieux Humides et De Bord De Mer —
du Québec et Des Maritimes. By Martine Lapointe.
2014. Editions Michel Quintin, 4770, rue Foster, Water-
loo, QC, Canada, JOE 2NO0. 456 pages, 34.95 CAD,
Paper.

* Plants of Southern Ontario. By France Royer, and
Richard Dickinson. 2014. Lone Pine Publishing, 87
East Pender Street, Vancouver, BC, Canada, VSA 1S9.
528 pages, 19.76 CAD, Paper.

* Trees of Eastern North America. By Gil Nelson,
Christopher J. Earle, and Richard Spellenberg. 2014.
Princeton University Press, 41 William Street, Prince-
ton, NJ, USA, 08540-5237. 720 pages, 29.95 USD,
Paper, 65.00 USD, Cloth.

* Trees of Western North America. By Richard Spel-
lenberg, Christopher J. Earle, and Gil Nelson. 2014.
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The Glory of the Tree — An Illustrated History. By
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Common Mosses of the Northeast and Appalachi-
ans. By Karl B. McKnight, Joseph R. Rohrer, Kirsten
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ton University Press, 41 William Street, Princeton, NJ.
USA, 08540-5237. 392 pages, 24.95 CAD, Paper.

2014

OTHER

Algonquin Park: A Portrait — The landscape, wild-
life and ecology of an iconic Canadian treasure.
By Jan and Martin Rinik. 2014. Formac Publishing,
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216 pages, 34.95 CAD, Cloth.

Essentials of Conservation Biology — 6 Edition.
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USA, 01375-0407. 603 pages, 94.95 USD, Cloth.

* Bird Lady — A Lifelong Love Affair with Birds.
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184 pages, 18.57 CAD, Paper.

Botanical Illustrators Handbook. By Sally Pinhey.
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Ramsbury, Wiltshire, UK, SN8 2HR. 128 pages, 16.99
GBP, Paper.

* Ancient Pathways, Ancestral Knowledge — Eth-
nobotany and Ecological Wisdom of Indigenous
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J. Turner. 2014. McGill-Queen’s University Press,
1010 Sherbrooke West, Suite 1720, Montreal, QC,
Canada, H3A 2R7. 1056 pages, 100.00 CAD, Cloth.

Extinction and Evolution — What Fossils Reveal
About the History of Life. By Niles Eldredge. Firefly
Books Ltd., 50 Staples Avenue, Unit 1, Richmond Hill,
ON, Canada, L4B 0A7. 256 pages, 45.00 CAD, Cloth.

Running Silver: Restoring Atlantic Rivers and their
Great Fish Migrations. By John Waldman. 2013.
Lyons Press (Globe Pequot Press), 246 Goose Lane,
P.O. Box 480, Guilford, CT, USA, 06437. 284 pages,
29.95 USD, Paper.

Tales from Gombe. By Anup Shah, and Fiona Rogers.
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Richmond Hill, ON, Canada, L4B 0A7. 320 pages,
69.95 CAD, Cloth.

Drawing and Painting Insects. By Andrew Tyzack.
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Lane, Ramsbury, Wiltshire, UK, SN8 2HR. 192 pages,
32.95 CAD, Cloth.

NEw TITLES 313

Wildlife Photographer of the Year — 50 Years —
How Wildlife Photography Became Art. Edited by
Rosamund Kidman Cox. 2014. Firefly Books Ltd.,
50 Staples Avenue, Unit |, Richmond Hill, ON, Cana-
da, L4B 0A7. 256 pages, 49.95 CAD, Cloth.

* Wallace, Darwin, and the Origin of Species. By
James T. Costa. 2014. Harvard University Press, 79
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Saving Turtles — A Kid’s Guide to Helping Endan-
gered Creatures. By Sue Carstairs. 2014. Firefly
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ON, Canada, L4B 0A7. 64 pages, 9.95 CAD, Paper.

Weird Birds. By Chris Earley. 2014. Firefly Books
Ltd., 50 Staples Avenue, Unit 1, Richmond Hill, ON,
Canada, L4B 0A7. 64 pages, 9.95 CAD, Paper.

Weird Frogs. By Chris Earley. 2014. Firefly Books
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Canada, L4B 0A7. 64 pages, 9.95 CAD, Paper.

Monsters of the Deep. By Camilla de la Bédoyere.
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Creatures of the Night. By Camilla de la Béedoyere.
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Firefly Encyclopedia of Animals. By Philip Whit-
field. 2014. Firefly Books Ltd., 50 Staples Avenue,
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pages, 19.95 CAD, Paper.

Firefly Encyclopedia of Dinosaurs and Prehistoric
Animals. By Douglas Palmer. 2014. Firefly Books
Ltd., 50 Staples Avenue, Unit 1, Richmond Hill, ON,
Canada, L4B 0A7. 256 pages, 19.95 CAD, Paper.

Minutes of the 135'* Annual Business Meeting of the
Ottawa Field-Naturalists’ Club January 14, 2014

Place and time:

Chairperson: Fenja Brodo, President

Fletcher Wildlife Garden, Ottawa, Ontario, 7:00 pm

Over 30 attendees spent the first half-hour reviewing the minutes of the previous ABM, the financial report
and the OFNC committees’ annual reports for 2012-2013. The meeting was called to order at 7:30 pm with

some opening remarks from the President.

1. Minutes of the Previous Annual Business
Meeting
It was moved by Annie Bélair and seconded by
Rémy Poulin that the minutes of the 134" Annual
Business Meeting be accepted as distributed.

Carried

2. Business Arising from the Minutes
Nil.

3. Communications Relating to the Annual
Business Meeting
Nil.

4. Treasurer’s Report by Ken Young

Ken started by explaining that the OFNC uses a
“fund accounting” system. This means that the revenues
and expenses for some OFNC activities are segregated;
each of these is a fund. A side effect is that the financial
statements do not include a single table presenting all
the revenues and expenses of the Club. Instead there are
separate statements for different types of fund. There-
fore Ken distributed a separate table showing a state-
ment of operations and changes in fund balances for
all OFNC funds.

All the funds except one are “restricted”. This means
that the revenues of the fund can only be spent on the
purpose established for the fund. One fund is “‘exter-
nally restricted”. That is the Macoun Camfield Endow-
ment, which results from a legal agreement made be-
tween the donor (the Camfield family, in memory of
Martha Camfield who passed away in 2010) and the
OFNC. This money in this fund is used to earn interest.
Half of the interest is reinvested to maintain the value
of the fund, and half is used to develop the Macoun
Field Club. The other restricted funds are based on re-
strictions that were established internally, by the OFNC.

There is one unrestricted fund, the “General Fund”,
which accounts for most revenues and expenses of the
Club.

These funds, then, are used to organize the revenues
and expenses of the OFNC. In the financial statements,
details about the General Fund are on page 3, the inter-

nally restricted funds are on page 4 and the externally
restricted fund is on page 5. The Statement of Financial
Position on page 2 is for the aggregate of all funds; it is
equivalent to the Balance Sheet of a for-profit business.
The Statement of Cash Flows on page 6 is also for all
funds taken together.

Starting in 2014, a new internally restricted fund
will be created for the Fletcher Wildlife Garden.

Ken then drew our attention to the financial results.
Net revenues for 2013 were positive for a second year,
so the Club is in a good position financially.

As for the 2014 budget, it was approved by Council
in November 2013; it is attached to the minutes of that
meeting and is available on the OFNC website where
minutes of Council meetings are kept, at http://ofne.ca
/council/minutes.php.

Fenja thanked Ken for his presentation, and for
his hard work as treasurer.

Motion: It was moved by Ken Young and second-
ed by Rob Alvo that the financial statements be accept-
ed as a fair representation of the financial position of
the Club as of September 30, 2013.

Carried

5. Nomination of the Accountant

Moved by Ken Young and seconded by Barbara
Chouinard that the accounting firm of Welch LLP be
nominated to conduct a review of the OFNC’s accounts
for the fiscal year ending September 30, 2014.

Carried

6. Committee Reports

The annual committee reports serve as a record of
what the OFNC does throughout the year. A copy had
been distributed to attendees at the beginning of the
meeting.

Moved by Annie Bélair and seconded by Ernie
Brodo that the reports be accepted as distributed.

Carried
Some notes:

a. The Birds Committee’s annual report states that
“members of the committee were active as volun-

2014

teers with the very successful BirdLife Interna-
tional World Congress held in Ottawa in the sum-
mer of 2013.” Fenja added that in December
2013, lan Davidson, member of OFNC Council
and executive director of Nature Canada, present-
ed Nature Canada’s “Affiliate Award” to formally
recognize the OFNC’s contribution to the phe-
nomenal success of the congress.

b. As of January 2014, the Excursions and Lectures
Committee will be called the Events Committee.

c. The Finance Committee’s annual report reflected
the fact that at the time of writing the report, the
OFNC financial statements were not finalized and
it was thought that a small deficit may be the re-
sult for the period. Now that the financial state-
ments are finalized, the result is a profit. This is
the reason for the discrepancy.

d. Dan Brunton pointed out that the Publications
Committee’s annual report states that the “chal-
lenge of getting The Canadian Field-Naturalist
(CFN) back on schedule has been virtually
achieved.” He announced that issue 127(4), the
last issue of 2013, was electronically sent to sub-
scribers on the day of this meeting, meaning the
CFN is indeed back on schedule. The crowd ap-
plauded to congratulate the CFN publishing team.

7. OFNC’s Twitter Accounts and Blog

Natalie Sopinka, new member of the Education
and Publicity Committee, introduced herself and de-
scribed some of the new social media initiatives that
were launched:

The OFNC now has three Twitter accounts: one
for the CFN, one for the Fletcher Wildlife Garden and
one for the OFNC in general.

A WordPress-based blog has been created (http:
//ofac.wordpress.com) for reports of monthly meetings
and events, as well as local nature-related information.
It is being monitored by members of the Education
and Publicity Committee. An email was distributed to
OFNC members to announce its launch and an article
on the subject will be published in Trail & Landscape.

She invited everyone to follow the Twitter ac-
counts and the blog, and said that anyone is welcome
to write entries for the blog.

8. Report of the Nominating Committee
A. MacKenzie

Relevant Excerpts from the OFNC Constitution

(revised February 2000)

Article 8 — “The Council shall consist of the of-
ficers of the Club and up to eighteen additional mem-
bers, all members of the Club.”

Atticle 12 — “The officers of the Club and other
members of the Council shall be elected annually at
the Annual Business Meeting. The nomination of suf-
ficient persons for election to the various offices and
membership of the Council shall be the responsibility

MINUTES OF THE 135™ ANNUAL BUSINESS MEETING

ws)
Nn

of the Nominating Committee, which shall act in the
manner prescribed in the By-Laws.

The Council shall, at the earliest possible date,
appoint chairs and members of Standing and ad hoc
committees and Editor and Business Managers, as
required for club publications.”

OFFICIAL DUTY
President

lst Vice-President
2nd Vice-President
Recording Secretary
Treasurer

NOMINATED OFFICERS
Fenja Brodo

Henry Steger
Eleanor Zurbrigg
Annie Belair

Ken Young

NOMINATED OTHER MEMBERS (in alphabetical order)
Daniel Brunton Don Hackett

Carolyn Callaghan Diane Kitching

Barbara Chouinard Ann MacKenzie

Julia Cipriani Karen McLachlan Hamilton
Owen Clarkin Lynn Ovenden

Barry Cottam Rémy Poulin

lan Davidson

Motion: Moved by Ann MacKenzie and second-
ed by Diane Lepage that this slate of nominees be ac-
cepted as members of the Council of the OFNC for
2014.

Carried

Fenja thanked David Hobden, Diane Lepage and
Jeff Skevington who will not be returning to Council
in 2014: she welcomed Julia Cipriani, now chair of the
Events Committee, who is rejoining Council.

316

9. Review of OFNC Constitution and
Bylaws for New Regulations

Ann MacKenzie, Chair of the Constitution Com-
mittee, gave a presentation on the likely implications
to the OFNC of the pending changes to the Ontario
Not-for-Profit Corporations Act. Changes are anticipat-
ed to our classes of membership, to the Council and
with respect to the notification and voting related to
the Annual Business Meeting. It is expected that a re-
vised constitution will be ready for members approval
at the January 2016 Annual Business Meeting. The
club is working with lawyers and will notify members
when there is a revised version for their consideration
and comment.

THE CANADIAN FIELD-NATURALIST

Vol. 128

10. In Remembrance

The OFNC noted with sadness the passing during
2013 of three members who had made a notable con-
tribution to the club and the naturalist community:

Gordon Pringle — died in February 2013. A key
member of the Birds Committee and the Birds Records
Subcommittee, which he chaired.

Violetta Czasak — died in May 2013. She named
the OFNC as sole beneficiary of her estate.

Bob Bracken — died in July 2013. A great natu-
ralist of the Ottawa area who started out as a member
of the Macoun Field Club.

11. Adjournment
Moved by Henry Steger and seconded by Rob
Alvo that the meeting be adjourned at 8:45 pm.

Carried

ANNIE BELAIR
Recording Secretary

2014

MINUTES OF THE 135™ ANNUAL BUSINESS MEETING

S17

The Ottawa Field-Naturalists’ Club 2012-2013 Annual Reports

Awards Committee

The Awards Committee manages the process to annually
recognize those OFNC members and other qualified persons
who, by virtue of their efforts and talents, are deserving of
special recognition. In 2013, nominations were received and
evaluated (see awards criteria http://www.ofne.ca/awards.php),
seven nominations were recommended to Council for approv-
al, and biographies were written for publication in the Club’s
journals. The awards were presented at the annual Soiree in
April. The recipients’ names, type of award(in brackets) and
rationale for recognition follow below:

* Dr. J. Bruce Falls (Honorary Member) — For his im-
mense contribution to the investigation, documentation
and protection of natural features and landscapes in On-
tario and throughout Canada.

* Peter Hall (Honorary Member) — For his work on butter-
flies among other species, the Fletcher Wildlife Garden
and biodiversity conservation, and for over 30 years of
service to the Club.

* Jay Fitzsimmons (Member of the Year) — For his work
managing the electronic publishing of The Canadian
Field-Naturalist (CFN) and also for his enthusiastic and
far-reaching promotion of the journal.

* Connie Clark (George McGee Service) — For over 10
years of active service with the Fletcher Wildlife Garden,
leading walks for the Club, committee work and writing
Trail & Landscape articles.

* David Seburn (Conservation Award — Member) — For
significant efforts for over a decade in turtle research and
conservation in the Ottawa area.

* Biodiversity Conservancy International and the National
Capital Commission (Conservation Award — Non-
Member): PT Dang (President BCI), Eva Katic (NCC)
— For the sand dune conservation project at the Pinhey
Forest in Ottawa.

* Diane Lepage (Anne Hanes Natural History Award) —
For her independent study of and publication in Trail &
Landscape on the moths of the Larose Forest.

Committee members: Irwin Brodo, Julia Cipriani, Christine

Hanrahan, Ann MacKenzie

ELEANOR ZURBRIGG, Chair

Birds Committee

The Birds Committee organized the Fall Bird Count (2012)
and along with the Club des Ornithologues de l’Outaouais
participated in the 2012 Christmas Bird Count. The Peregrine
Falcon Watch 2013 had a mixed season. The pair at the Data
Center had a successful year fledging their one chick. How-
ever, once again, the downtown pair’s eggs did not hatch. The
Bird Record Sub-committee met during the year to review
rare bird reports and is putting the finishing touches on a
new checklist for Ottawa. This project was stalled by the
passing of long-time committee member Gordon Pringle who
was a key player in the development of the checklist. Members
of the committee were active as volunteers with the very suc-
cessful BirdLife International Congress held in Ottawa in the
summer of 2013. Committee members are also involved in
bringing the Ontario Field Ornithologist’s Annual Conference
to Ottawa for 2014. The committee continues to operate a
series of bird feeders throughout the Ottawa Greenbelt. The
committee is also in discussion with FLAP (Fatal Light Aware-
ness Program) to place a chapter in Ottawa. This program

deals with how to avoid the damage caused by bird/window
collisions.

Curis TRAYNOR, Chair

Conservation Committee

The OFNC conservation committee was officially re-acti-
vated in late 2012, following a time of hiatus. In the past year,
the CC has tackled a range of conservation issues in Ontario.
We have written several letters on conservation matters, such
as the issue of cottage lease extensions in Algonquin Park, the
issue of overly enthusiastic mowing practices of meadow areas
by the municipal authorities, and a plan to expand a hydro dam
in the town of Almonte downstream from the sensitive Ap-
pleton Wetland ANSI. Other similar projects have been initi-
ated such as the impact of housecat predation on wildlife, the
implications of a possible oil spill from rail or pipelines on our
local ecosystems, the turtles being stranded on the road in the
vicinity of Mud Lake and a bioblitz near a proposed landfill
site near Carlsbad Springs.

A major focus of the new chair is flora conservation, and to
this end a few “personal” projects have been initiated. These
include: public walks with emphasis on the ecology of and the
how-to of plant identification, and the current status of uncom-
mon, rare, or declining native plants in Eastern Ontario and
Western Quebec. In this vein, an inventory of the flora of the
Fletcher Wildlife Garden has been initiated and is expected to
be completed late 2013 or early 2014.

Many other talented and dedicated members of the OFNC
CC continue to work on their own personal conservation pro-
jects. A committee meeting will be held in late 2013 to brain-
storm about increasing CC member engagement, and new ap-
proaches to preserving the ecosystems and indigenous species
of our region.

OWEN CLARKIN, Chair

Education and Publicity Committee

The primary duty of this committee is to make the Club, as
well as natural history, known to the community. It is respon-
sible for the club’s publicity, providing speakers/walks as re-
quested by outside groups, considering special educational
projects, assisting with membership drives and coordinating
the sale of club materials.

Committee members have diverse skills. We make things
and sell them, we write, we edit, we create displays and fly the
club banner at local events, we ask people to give nature talks.
We have not excelled at PR but there is hope: a new member
of the committee will focus on publicizing OFNC events and
accomplishments. During the past year, the committee expe-
rienced a change in chair, lost a few valued members and
gained some new ones. We met 6 times, about every two
months.

2012 was a big year for sales of club materials such as bird
and butterfly checklists, lanyards, 10x loupes and books. Five
new creations bear the club logo: a banner, a bookmark, an
OFNC sign atop the entryway sign to the Fletcher Wildlife
Garden, a lanyard and a lens-wipe (the latest toy). For next
year we are considering how to standardize the use of OFNC’s
logo and brand across many club activities.

We brought OFNC displays to the Soiree in April, the Lac
Deschenes Bird Fair at Andrew Haydon Park in May and an
Environment Fair at Place Vincent Massey in June. New this
year was a regional selection process for OFNC’s sponsorship

of a youth who attended Ontario Nature’s Youth Summit in
September.

Many club members share their natural history expertise
with other community groups. We especially thank the gen-
erous souls who said “yes” to requests made directly to
OFNC:

* Dave Moore and Bev McBride led a nature walk to Mud
Lake for visitors from Friendship Force in June. Dave
also presented a birding talk to the Seniors Program at
the Unitarian Church in September.

¢ Eleanor Thomson gave a nature talk to the women’s
group at the Lord Lansdowne Residence.

¢ Joan Harvey visited Uplands Catholic School to advise
the students about extending their butterfly garden.

* Jeff Skevington and Carolyn Callaghan were judges for
the OFNC special awards at the Ottawa Regional Sci-
ence Fair.

LYNN OVENDEN, Chair

Excursions and Lectures Committee

Over the course of 2013, the excursions and lectures com-
mittee coordinated 40 hikes, 5 workshops, 9 monthly meet-
ings, the annual awards celebration and the annual business
meeting. In addition to bringing back many tried and true
events and leaders, several new events were promoted.

The hikes and workshops covered a wide breadth of topics,
including: general natural history (6), birds (15), insects (8),
spiders (1), plants (8), amphibians and reptiles (3), mammals
(1), fish (1), nature photography (1), lichens (1) and fungi (1).
Highlights included workshops on invasive plants, lichens,
nature photography, arthropods of your home and crane flies,
as well as a tour of our national insect collection, an overnight
trip to the Kazabazua sand-plain, our annual butterfly count
and an exploration of Brewer Pond life. Monthly meetings
continued to be held in the Museum of Nature and included
talks on birds (5), mammals (1), conservation (1), canoeing
(1) and salamanders (1). Committee members included Rob
Alvo, Holly Bickerton, Julia Cipriani, Hume Douglas, Jakob
Mueller and Jeff Skevington (chair). If you have any new
ideas for events or would like to be a leader, please contact
Jeff Ghskevington@gmail.com) or other members of the
committee.

JULIA CIPRIANI, Committee Member

Finance Committee

As a result of the major changes to the OFNC’s financial
reporting systems over the last couple of years, Council con-
tinues to benefit from the now smoothly running financial
reporting system. With the revision of the OFNC’s detailed
chart of accounts last year, information is now more accurate
and detailed with respect to the club’s revenues and expen-
ditures. The financial reports provided by the OFNC Treasurer
several times a year are a useful tool for Council and com-
mittees.

The Finance Committee met several times during the
2012/2013 fiscal year to discuss several issues. In March, the
Finance Committee proposed a new methodology in allocat-
ing investment income. This was done to make the allocation
of interest income less complicated at year end. This new
methodology was approved by Council, the Camfield Family
and the OFNC’s auditor.

In May, the Finance Committee was asked to update Coun-
cil on the current status of honoraria in the OFNC. This infor-
mation was presented to Council, and a motion was made to

THE CANADIAN FIELD-NATURALIST

Vol. 128

resume the honorarium paid to the CFN Manager. This motion
was passed by Council.

Lastly, in September, the 2014 budget discussions resulted
in a motion to provide FWG with their 2013 budgeted funds.
However, in the future, no funds will be transferred to FWG
without prior approval through the budgetary process. This is
to bring FWG in line with all the other OFNC committees.
Budgets are determined by each respective committee, and
Council approves the budget. Payment is then made upon ap-
proved expenses against the budget. This process provides
greater transparency and allows for the timely matching of
revenues and expenses.

The club is continuing to run a deficit though it is project-
ed to be lower this year than in recent years. With an improved
financial reporting system, it is easier to identify the issues.
There are areas that the Finance Committee intends to review
in the near future for Council’s consideration. This would in-
clude issues such as the uncollectible subscriptions invoices,
the decrease in OFNC membership numbers, the possible
restriction of access to the electronic CFN to only paid mem-
bers and the declining CFN institutions subscription revenue.

BARBARA CHOUINARD, Chair

Fletcher Wildlife Garden

This is been a busy year for the Fletcher Wildlife Garden.
All of the usual activities have proceeded apace, along with a
number of other activities, events and achievements. Many of
these have involved other groups. Several subcommittees have
worked on a number of changes to how things get done at the
FWG. This report presents highlights of these as they occurred
throughout the year.

Our year kicked off on April 7 with a Spring Fling, inspired
by the successful end-of-2012 potluck for volunteers. This was
more than a social event, as group and habitat managers pro-
vided volunteers with work plans and the FWG treasurer gave
an overview of the financial situation. On April 9, the first
meeting in several years between Agriculture and Agri-Food
Canada and FWG representatives proved productive, with a
number of issues being addressed. We have had a very posi-
tive working relationship with AAFC and continue to main-
tain this relationship through other means, such as the Central
Experimental Farm Advisory Committee (CEFAC).

Shell grant: FWG’s application for funding to dredge the
pond and do related work, approved by council and AAFC,
was not accepted by the Shell FuellingChange program. Fur-
ther effort is in abeyance pending the results of an environ-
mental assessment by Environment Canada.

FWG Volunteer Groups: The Butterfly Meadow group and
the Friday morning group continued to expand in scope
through the hard work of volunteers under the able direction
of their managers. The Tuesday Invasive Species Group was
much reduced in numbers and hours of operation in a take-
stock year, trying to assess the value of its efforts in regard to
DSV. This highly successful invasive plant is a limiting factor
in our efforts to improve and maintain the habitats. Rather than
attempting to cover the FWG as a whole, specific areas were
targeted for intensive effort. Brush cutters were tested for ef
fectiveness and AAFC mowed a couple of monocultural stands
of DSV in July. Inventories and maps of invasive shrubs and
trees are being prepared. The October annual meeting of the
Ontario Invasive Plant Council provided, as in the past, useful
information on both current research and attempts by other
organizations to control invasives. The Monarch Way Station
was the focus of milkweed planting work bees in June and

2014

September. Planting, DSV removal and development of the
Bill Cody Fern Garden continued in the Ash Woods. A job
description for Habitat Managers is in final stages of prepara-
tion. Outreach to potential volunteers through our membership
in Volunteer Ottawa has been more consistent and successful
this year.

Activities involving Outside Organizations: A number of

organizations participated in special work projects and other
events. These included two full days from the Stewardship
Rangers, a return visit from PricewaterhouseCoopers, and
80 students from a Carleton University environmental studies
class. Walks and tours were undertaken by several outside
organizations, including the Canadian Wildlife Federation and
the Ottawa Horticultural Society. The FWG was a site for the
inaugural Ottawa Sustainability Tour, although we had more
volunteers on hand as greeters than tour participants. A poten-
tially useful liaison with a professor in the Carleton University
Biology department was established. The OFNC Bug Day,
held at the Interpretation Centre/Backyard Garden, was a roar-
ing success.

Research: The FWG has been the object of new research
this year. Researchers from the Invasive Species Centre in
Sault Ste Marie and the University of Guelph Microbiology
Department are conducting several experiments in attempts
to understand the soil chemistry of a number of ‘mystery’ cir-
cles, found at various locations. These circles are characterized
by a lack of DSV within and a perimeter of small, yellowed
DSV plants, with normal DSV growth beyond. Even more sig-
nificantly, the FWG is slated to be one of the test sites for the
release in 2014 of larvae of the moth Hypena opulenta that
feed on the leaves of DSV. Special notice must be given to
completion this year of Renate Sander-Regier’s PhD thesis on
the FWG, “The Power of a Small Green Place: A case study
of Ottawa’s Fletcher Wildlife Garden.”

Communications: The newsletter has been resurrected and
revamped by our new editor; the photo galleries on PBase
continue to be well received; the website is recovering from a
loss of data; social media including Facebook and twitter pro-
vide further outreach.

Interpretation Centre: A job description was drawn up for
the new role of Interpretation Centre manager. Two OFNC
members volunteered and they started in September.

Finances and first budget: The annual plant sale, our major
fundraising effort, was very successful again this year, raising
about $4000.00. While questions remain regarding the rela-
tionship between OFNC and FWG funds, recent attempts to

MINUTES OF THE 135™ ANNUAL BUSINESS MEETING

319

clarify them have resulted in the first ever annual budget for
the FWG.

Management: The management committee continues to
operate with a rotating chair. This is working reasonably well
in regard to month-to-month business. However, the roles of
the chair and the MC and other matters such as longer-term
planning and council’s role with regard to the FWG are up for
discussion. The lease and collaborative agreements between
AAFC and the OFNC are to be renewed by April 1, 2014. Pre-
liminary discussions between the parties have been positive.
Our year wraps up with a fall potluck on November 24.

BARRY COTTAM, Committee Member

Macoun Field Club Committee

The Committee held one planning meeting ahead of the
Macoun Club’s start-up in September and put together the
month-to-month program by telephone and e-mail. An up-to-
date schedule and illustrated record of the weekly activities
was maintained on its website (macounfieldclub.ca). Commit-
tee members supervised or gave presentations at 19 indoor
meetings and led 15 regular field trips during the school year;
they also led another four special trips, two jointly with the
OFNC and two with SOS Dunes (Pinhey Sand Dunes). Indoor
sessions were held in the Fletcher Wildlife Garden’s Interpre-
tation Centre. Most field trips took place either at the Macoun
Club’s nature-study area in Stony Swamp or on private prop-
erties in Lanark County. The Committee produced issue no. 67
of the Club’s annual publication (Zhe Little Bear), and the sev-
enth and eighth annual notebooks of sightings in the Stony
Swamp study area (bound in hard cover), all for distribution
to members.

RosBert E. Lee, Chair

Membership Committee

The distribution of the membership for 2013 on September
30, 2013 is shown in the table below, with the corresponding
numbers for 2012 shown in brackets. “Others” represent, for
the greatest part, affiliate organizations that receive compli-
mentary copies of the Club’s publications. The decrease in
total membership of 28 continues a long-term overall trend
but one that follows an average reduction of 21 per year dur-
ing the period from 2006 to 2012. Local membership (within
50 km of Parliament Hill) decreased to 617 from 633 in 2013
and 2012, respectively.

Henry STEGER, Chair

———————————————— _—"—“"“—O|?5°° > O00“

CANADIAN USA OTHER TOTAL
2013. =—s«.2012 2013-2012 2013 —-2012 2013-2012
ee era ee eet
ivi 324 (351) 12 (13) 0) (0) 336 (364)
cera 288 (285) 0 (1) (1) 289 (287)
Student* 1] (10) 0 (0) 0 (0) 11 ol
T and L ps (3) 0 (0) 0 (0) 2 (2)
Honorary 21 (21) 0 (0) 0 (0) = (21)
Life 45 (47) l (3) l (1) ri pani
Other 23 (23) I (0) I (1) 2 (24)
TOTAL 714 (740) 15 (17) 3 Q) 732 (760)
324 = (351) I 3) 0 (0) 336 ~~ (364)

EEE

*Student Membership was initiated in Year 2012 and is intended to encourage interest in natural history among high school

and university students.

ee)
Ww
=

Publications Committee

Publications Committee members in 2013 were Carolyn
Callaghan, Paul Catling, Jay Fitzsimmons, Sandra Garland,
Tony Gaston, Karen McLachlan Hamilton, William Halliday,
Elizabeth Morton, Frank Pope, Jeff Saarela and Dan Brunton
(chair). We met formally twice during the year to discuss a
wide variety of issues and to provide advice and information
to both the OFNC Council and to the editorial teams. A great
number of informal communications were also conducted
electronically amongst committee members on a variety of
publication topics, primarily in regards to The Canadian
Field-Naturalist.

The challenge of getting The Canadian Field-Naturalist
back on schedule has been virtually achieved. This is the
result of several years of tremendous effort by the current and
former editorial teams. Editor-in-Chief Carolyn Callaghan
has overseen the publication of five issues of the journal this
year, 126(3) through 127(3). The final issue of the 2013 vol-
ume is to be posted and circulated within the first few weeks
of 2014. With the journal entirely back on schedule, we ex-
pect the publication of four issues in 2014.

As many manuscripts had been received by October 2013
as had been received in all of 2012, which already had seen a
20% increase over recent years. We take this as confirmation
of the journal’s increasingly appeal for the appropriate and
timely documentation of important field-oriented natural sci-
ences research. Great strides have also been made by Journal
Manager Jay Fitzsimmons and others in the unglamorous but
critical area of publication production, with efficiencies
achieved with indexing and improved printing turn-around
times. Colour images continue to enhance the appearance of
more papers than was possible in years past.

All in all, progress has been most encouraging. The full
achievement of a ‘normal’ publication schedule for The
Canadian Field-Naturalist will now allow us to put more
focus on logistical and administrative improvements. That will
simplify production and make life easier both for the mem-
bers and subscribers receiving the journal and for the largely
volunteer effort required to make these happen.

Editor Karen McLachlan Hamilton and her team produced
the four issues of 7rail & Landscape in their usual timely and
efficiently manner. Trail & Landscape fulfills an important
newsletter function and also continues to document important
ecological information on the regional biodiversity and its con-
servation needs.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Discussions of a possible special publication addressing the
biodiversity of an important Ottawa Valley natural area were
initiated in 2013. This may proceed to the production of a
stand-alone document in 2014.

DANIEL F. BRUNTON, Chair

Treasurer’s Report

Financial Position: At the time of writing, the Financial
Statements have not been finalized. However, it appears that
revenues will exceed expenses.

Insurance: This year, our insurance coverage was expand-
ed to include Directors’ & Officers coverage as well as Com-
mercial General Liability.

Committee Budgets: This year more attention was paid to
the preparation of the annual budget. As a result the budget
presents a more comprehensive forecast of OFNC revenues
and expenses than in the past.

Violetta Csazak Bequest: The Club received a bequest
from Violetta Czasak. I have been appointed as executor of
the estate. We are proceeding with the sale of her condomini-
um, but in the current real estate market this is a slow process.

Routine Events: Most of the Treasurer’s duties continue
from one year to the next. They include:

¢ Filing the Charities Information Report with the Canada
Revenue Agency;

¢ Preparing T4A returns for people receiving honoraria
or contract fees from the Club;

* Obtaining Proof of Insurance certificates for Club
events;

¢ Updating the Club’s information with our bank, the
Canadian Imperial Bank of Commerce, and our online
payments processor, PayPal:

* Depositing cheques received and writing cheques to
pay expenses;

¢ Making financial reports to Council and Committees,
such as for The Canadian Field-Naturalist, the Seed-
athon, the Fletcher Wildlife Garden and the Macoun
Field Club.

I would like to express my appreciation for all the assistance
that I have received from Frank Pope, the past Treasurer, and
Ann MacKenzie, who handles the investment activities of the
Club.

KEN YOUNG, Treasurer

2014

Review Engagement Report

MINUTES OF THE 135™ ANNUAL BUSINESS MEETING 32]

To The Members of THE OTTAWA FIELD-NATURALISTS’ CLUB

We have reviewed the statement of financial position

of The Ottawa Field-Naturalists’ Club as at September

30, 2013 and the statements of operations, changes in
fund balances and cash flows for the year then ended.
Our review was made in accordance with Canadian
generally accepted standards for review engagements
and accordingly consisted primarily of enquiry, ana-
lytical procedures and discussion related to informa-
tion supplied to us by the club,

A review does not constitute an audit and conse-
quently we do not express an audit opinion on these
financial statements.

Based on our review, nothing has come to our atten-
tion that causes us to believe that these financial state-
ments are not, in all material respects, in accordance
with Canadian accounting standards for not-for-profit
organizations.

CHARTERED ACCOUNTANTS
Licensed Public Accountants

Week LLP

Ottawa, Ontario
December 23, 2013

The Ottawa Field-Naturalists’ Club
Statement of Financial Position
September 30, 2013

2013 2012
ASSETS
CURRENT ASSETS
Cash and cash equivalents
(note 4) $55,862 $156,201
Short-term investments (note 4) ~ 43,817
Amounts receivable 24,380 16,138
Prepaid expenses 633 1,403
80.875 PH Wf syo}.)
LONG-TERM INVESTMENTS (note 4) 537,456 386,170
$618 331 $603 729
LIABILITIES AND FUND BALANCES
CURRENT LIABILITIES
Accounts payable and
accrued liabilities $3 582 $2,283
Deferred revenue 16,683 21,239
20.265 QBS 22
LifE MEMBERSHIPS (note 5) ~ 1,240
FUND BALANCES
General fund 317,208 303 378
Internally restricted funds 244 376 239.689
Martha Camfield
endowment fund __ 36,482 __ 35,900
598,066 _ 578.967
$618,331 $603 729
Approved by Council:
eR, HN ee ERR CLE rime President
PO OE Te Ree Cae ROR Ret Treasurer

(See accompanying notes)

PREPARED WITHOUT AUDIT

eS)
Nw
i)

THE CANADIAN FIELD-NATURALIST

The Ottawa Field-Naturalists’ Club
Statement of Operations and Changes in Fund Balance

— General Fund

Year Ended September 30, 2013

REVENUES
Membership fees
Donations and bequests

The Canadian Field-Naturalist

— subscription revenue

— author charges
Fletcher Wildlife Garden
Pelee Trip

Interest income
Advertising

Other

EXPENSES

OPERATING:
Affiliation fees
Bookkeeping
Courier and postage
Insurance
Interest and bank charges
Membership
Professional fees
Website

General and miscellaneous

ACTIVITY:

The Canadian Field-Naturalist

Fletcher Wildlife Garden
Awards committee
Soiree — net

Birds Committee
Donations

Education and publicity
Excursions and lectures
Macoun Club

Pelee Trip

Trail and Landscape

TOTAL EXPENSES
Net REVENUES

FUND BALANCE,
BEGINNING OF YEAR

FUND BALANCE,
END OF YEAR

2013

$ 28,703
27,149

26,153
28,234
4,395

13,744
1,675
1,828

131,881

450
4,94]

359
1,245
1,400
1,000
2,800
3,500
3,452

19,147

118,051

13,830

303,378

$317,208

2012

$ 30,308
21,954
32,463
27,984

5,684
14,375
12,465

680
145,913

114,993

30,920
272,458

$303,378

Vol. 128

The Ottawa Field-Naturalists’ Club

Statement of Changes in Fund Balance — Martha
Camfield Endowment Fund

Year Ended September 30, 2013

2013 2012
Fund Balance, Beginning of Year $ 35,900 $ 35,268
Interest 582 632
Fund Balance, End of Year $ 36,482 $ 35,900

Note: The interest above represents half of the interest generated by
the fund, which is reinvested in the capital of the fund. The other half
of the interest is recognized as interest revenue of the Macoun Fund
and is made available for the use of the Macoun Field Club.

The Ottawa Field-Naturalists’ Club
Statement of Cash Flows
Year Ended September 30, 2013

2013 2012
Cash Flows from Operating
Activities
Net revenues — all funds $ 1s-sil7 $. 33.275
Adjustments for:
Amounts receivable (8,242) 17,345
Investments (16,031) (15,956)
Prepaid expenses 770 (1,113)
Accounts payable and
accrued liabilities 1,299 (11,329)
Deferred revenues (4,556) 7,518
Life memberships (1,240) (2,120)
(9.483) 27,620
Cash Flows from Financing
Activities
Purchase of investments (137,178) (120,000)
Proceeds from maturity
of investments 45,740 46,043
(91,438) (73,957)
Cash Flows from Financing
Activities
Endowment interest reinvested 582 632
Increase (Decrease) in Cash
and Cash Equivalents (100,339) (45,705)
Cash and Cash Equivalents
at Beginning of Year 156,201 201,906
Cash and Cash Equivalents
at End of Year $ 55,862 $ 156,201

Note: Cash equivalents include fixed income investments maturing
within three months

(See accompanying notes)

PREPARED WITHOUT AUDIT

2014 MINUTES OF THE 135" ANNUAL BUSINESS MEETING 323

The Ottawa Field-Naturalists’ Club

Statement of Operations and Changes in Fund Balances — Internally Restricted Funds
Year Ended September 30, 2013

General Anne Hanes De Kiriline
Reserve for Manning Seedathon Memorial Lawrence Macoun 2013 2012
Contingencies Fund Fund Fund Fund Fund Total Total
Revenues
Donations $ —- §$ 100) S 1130) "Ss - § — $ 1,425 $ 2655 $ 3,500
Interest : 3,888 = - 582 4.470 6,163
~ 3,988 1,130 - - 2,007 TAZS 9,663
Expenses
CFN author support ~ -- ~ - - - 6,051
Donations ~ — ~ ~ 313
Seed ~ 1,110 - ~ 1,110 944
Macoun activities - - - _ 209 E255
Other - - is - - 75
~ 1,110 75 - ess} 2,438 7,308
Net Revenues (Expenses) - 3,988 20 (75) - 754 4.687 2355
Fund Balances,
Beginning of Year 100,000 119,803 788 596 13,384 5,118 239,689 237,334

Fund Balances, End of Year $100,000 $123,791* $808 oS $ 13,384 $ 5,872 $244,376 $239,689

* includes principal of $100,000 plus undistributed income of $23,791.

(See accompanying notes)

PREPARED WITHOUT AUDIT

THE CANADIAN FIELD-NATURALIST

Vol. 128

The Ottawa Field-Naturalists’ Club
Notes to the Financial Statements
Year Ended September 30, 2013

1. Purpose of Organization and Tax Status

The Ottawa Field-Naturalists’ Club (the “club”) is a regis-
tered charitable organization incorporated under the Corpora-
tions Act of the Province of Ontario. The club promotes the
appreciation, preservation, and conservation of Canada’s nat-
ural heritage, encourages the investigation, publishes the
results of research in all fields of natural history, and diffuses
the information to the public and supports and cooperates with
other organizations engaged in preserving, maintaining and
restoring environments of high quality for living things.

The club is a registered charity, and is exempt from income
taxes by virtue of section 149(1)(f) of the Income Tax Act
(Canada).

2. Significant Accounting Policies

Basis of accounting

These financial statements have been prepared in accor-
dance with Canadian accounting standards for not-for-profit
organizations.

Revenue recognition
(i) The club follows the deferral method of accounting
for contributions. Restricted contributions are rec-
ognized as revenue in the year in which the related
expenses are incurred. Unrestricted contributions are
recognized as revenue when received or receivable
if the amount to be received can be reasonably esti-
mated and collection is reasonably assured. Endow-
ment contributions are recognized as direct increas-

es in net assets.

(ii) Membership fees are recognized as revenue propor-
tionately over the fiscal year to which they relate.
The club’s membership year is January | to Decem-
ber 31. The portion of membership fees that are re-
ceived but not yet recognized as revenue are record-
ed as deferred revenue.

(iii) Subscription revenue is recognized as revenue at
the time of shipment of the related publication. The
liability for the portion of subscription revenues in-
voiced in advance of shipment is recorded as de-
ferred revenue.

(iv) Advertising revenue is recognized in the period in
which the advertisement is published.

(v) Interest income comprises interest from cash and
cash equivalents and investments. Interest on cash
equivalents and investments is recognized over their
term using the effective interest method. Interest
income derived from the investment of restricted
contributions, where the contribution agreement
specifies that the investment income is restricted,
is accounted for the same manner as the restricted
contributions.

Fund accounting
The Club maintains its accounts in accordance with the

principles of fund accounting. Resources are classified for

accounting and reporting purposes into funds according to
the activity or object specified.

(1) General Fund

The General Fund reports revenue and expenses
relating to general operations and administration
activities.

(ii) Martha Camfield Endowment Fund

The Martha Camfield Endowment Fund was estab-
lished by the family and friends of Martha Camfield
to help continue her efforts to have children study,
understand, respect and preserve their natural envi-
ronment. Half of the interest generated by the fund
is reinvested in the capital of the fund while the other
half of the interest generated is made available only
for the use by the Macoun Field Club (the Macoun
Fund).

(iii) General Reserve

The General Reserve For Contingencies was estab-
lished by the Club to fund outstanding operating ex-
penses should the Club discontinue its operations.

(iv) Manning Fund

The Manning fund was established by a bequest, and
the interest generated is used to assist authors to pub-
lish articles in The Canadian Field-Naturalist.

(v) Seedathon Fund

The Seedathon fund collects donations from the an-
nual bird sighting event and purchases seed for the
Club’s bird feeders.

(vi) Anne Hanes Memorial Fund

The Anne Hanes Memorial fund was raised in mem-
ory of Anne Hanes, the founding editor of Trail and
Landscape, and is used to finance the annual winners
of the Anne Hanes Natural History Award.

(vil) De Kiriline-Lawrence Fund

The de Kiriline-Lawrence fund was funded by a be-
quest from the popular author of nature books, and is
supplemented by annual donations and used to sup-
port conservation efforts.

(vill) Macoun Baillie Birdathon Fund

The Macoun Baillie Birdathon fund recognizes the
donations and pledges based upon the number of
bird sightings in the one day birdathon sponsored
by Bird Studies Canada, and is used to support the
Macoun Field Club, a youth club.

Financial Instruments

The Club initially measures its financial assets and finan-
cial liabilities at fair value adjusted by transaction costs in the
case where a financial asset or financial liability is subsequent-
ly measured at cost or amortized cost. The club measures all

PREPARED WITHOUT AUDIT

2014

of its financial assets and financial liabilities at cost or amor-
tized cost.

Cash and cash equivalents

Cash and cash equivalents include highly liquid invest-
ments with maturities of three months or less.

Capital assets

Capital assets are expensed in the year of acquisition.

Donated services

The club is dependent on the voluntary service of many of

its members. As there is difficulty in determining the fair value
of voluntary services, they are not recognized in these finan-
cial statements.

Internally restricted net assets

Internally restricted net assets represent the amount ap-
proved by the Council to be set aside for special purposes.
These amounts are not available for unrestricted purposes
without the approval of Council.

Use of estimates

The preparation of financial statements in conformity with
Canadian accounting standards for not-for-profit organizations
requires management to make estimates and assumptions that
affect the reported amounts of assets and liabilities and disclo-
sures of contingent assets and liabilities at the date of the finan-
cial statements and the reported amounts of revenues and ex-
penses during the reporting period. Actual results could differ
from these estimates.

Estimates and underlying assumptions are reviewed on
an ongoing basis. Revisions to accounting estimates are rec-
ognized in the year in which the estimates are revised and in
any future years affected.

3. Financial Instruments

The club is exposed to various risks through its financial
instruments. The following analysis provides a measure of the
club’s risk exposure and concentrations as at September 30,
2013.

Credit risk

The club is exposed to credit risk resulting from the pos-
sibility that parties may default on their financial obligations,
or if there is a concentration of transactions carried out with
the same party, or if there is a concentration of financial oblig-
ations which have similar economic characteristics, that could
be similarly affected by changes in economic conditions, such
that the club could incur a financial loss. The club does not
hold directly any collateral as security for financial obliga-
tions of counterparties.

The club’s maximum exposure to credit risk represents
the carrying value of its cash, amounts receivable and invest-
ments, totalling $617,698 (2012 — $602,326).

The club’s cash is deposited with Canadian financial insti-
tutions: as a result management believes the risk of loss on
cash to be remote. The cash equivalents and investments con-
sist primarily of government bonds and guaranteed investment
certificates of Canadian financial institutions of high credit
quality. Possible changes to the credit quality of these securi-
ties exposes the club to credit risk. The club manages its expo-
sure to this risk by holding a diversified portfolio with varied
maturities. The club reduces its exposure to credit risk on its
amounts receivable by reviewing the accounts on a regular

MINUTES OF THE 135" ANNUAL BUSINESS MEETING

basis, following up on outstanding amounts and creating an
allowance for doubtful accounts when applicable.

An allowance of $1,160 for impairment has been recorded
against the amounts receivable.

Liquidity risk

Liquidity risk is the risk that the club cannot meet its debts
when they become due. Liquidity risk also includes the risk of
the club not being able to liquidate assets in a timely manner
at a reasonable price.

The club meets its liquidity requirements by monitoring
its expected future cash flow requirements and holding a sig-
nificant amount of assets that can be readily converted into
cash.

Market risk

Market risk ts the risk that fair value or future cash flows
of a financial instrument will fluctuate because of changes in
market prices. Market risk is comprised of currency risk, inter-
est rate risk and other price risk.

Currency risk

Currency risk refers to the risk that the fair value of finan-
cial instruments or future cash flows associated with the instru-
ments will fluctuate relative to the Canadian dollar due to
changes in foreign exchange rates.

Approximately $18,000 (2012 — $14,000) of club’s cash
and cash equivalents are denominated in U.S. currency. How-
ever, the club primarily transacts in Canadian dollars. As a
result, management does not believe it is exposed to signifi-
cant currency risk.

Interest rate risk

Interest rate risk refers to the risk that the fair value of
financial instruments or future cash flows associated with
those instruments will fluctuate due to changes in market inter-
est rates. The exposure of the club to interest rate risk arises
from its interest bearing assets.

The club’s cash includes amounts on deposit with Cana-
dian financial institutions that earn interest at market rates.
Fluctuations in market rates of interest on cash do not have a
significant impact on the club’s financial operations.

The club manages the interest rate risk of its cash equiva-
lents and investments by the implementation of prudent invest-
ment policies. The club’s investments in bonds mature at face
value on a staggered basis over the next twelve years. The lad-
dered structure of maturities helps to enhance the average
portfolio yield while reducing the sensitivity of the portfolio
to the impact of interest rate fluctuations. Effective interest
rates to maturity for these securities range from 2.40% to
4.36% (2012 2.58% to 4.75%).

Other price risk

Other price risk refers to the risk that the fair value of
financial instruments or future cash flows associated with the
instruments will fluctuate because of changes in market prices
(other than those arising from currency risk or interest rate
risk), whether those changes are caused by factors specific to
the individual instrument or its issuer or factors affecting all
similar instruments traded in the market.

The club is not exposed to other price risk.

Changes in risk

There have been no changes in the club’s risk exposures
from the prior year.

PREPARED WITHOUT AUDIT

eS)
Nw
ON

4. Cash And Cash Equivalents And Investments

Cash and cash equivalents are comprised of:

Cash in accounts — at fair value

Canadian Western Bank — 4.41% due October 9, 2012
CIBC GIC — 1.4% due October 12, 2012

Ontario — 4.64% due December 2, 2012

Short-term investments are comprised of:

Manitoba — 4.75% due September 5, 2013

Long-term investments are comprised of:

CIBC — 4.19% due October 31, 2014

New Brunswick — 4.30% due December 3, 2015
Home Trust GIC — 2.40% due October 12, 2016
Ontario Hydro — 4.01% due November 26, 2016
Ontario — 4.07% due December 2, 2017

Ontario — 2.58% due December 2, 2018

British Columbia — 3.74% due March 5, 2019
Newfoundland — 4.36% due January 7, 2020
British Columbia — 3.26% due August 23, 2021
Nova Scotia Power — 2.80% due February 26, 2022
Hydro Quebec — 3.12% due February 15, 2023
Manitoba — 3.82% due September 5, 2025

5. Commitments

Life memberships

The club is committed to provide for regular membership
benefits to lifetime members. Since it is not practicable to
determine the total liability associated with providing these
benefits for the rest of the lives of these individuals, the annu-
al costs are expensed as incurred. Lifetime memberships are
no longer being offered by the club. As of September 30, 2013,
there were 49 (2012 - 51) active lifetime members.

Fletcher Wildlife Garden
The Fletcher Wildlife Garden (FWG) is 6.5 hectare prop-
erty of the Central Experimental Farm in Ottawa, Ontario and

THE CANADIAN FIELD-NATURALIST

Vol. 128

2013 2012
Market Amortized Amortized
Value Cost ___Cost _
S$ 52862 $ 55,862 $ 88,793
fe BS 30,872
_ = 21,284
= = 15,252
$ 55,862 $ 55,862 $ 156,201
2013 2012
Market Amortized Amortized
Value Cost Cost
= - $ 43,817
2013 2012
Market Amortized Amortized
Value Cost Cost
$ 69,510 $ 67,782 $ 65,056
63,539 60,582 60,850
34,175 34.175 =
26,600 26,058 25,054
52,200 48,588 46,687
62,810 62,861 61,280
29,079 27,481 26,491
43,709 40,605 38,908
63,438 63.860 61,844
15,904 1726 =
38,423 41,185 =
45,936 APT NSS =
$ 545,323 $537,456 $ 386,170

is a long-term project of the club. The FWG is managed by a
club committee and maintained by club volunteers. The costs
associated with maintaining the property are approximately
2,000 hours of voluntary human resources per year, plus reg-
ular maintenance and cleaning supplies. The fair value of the
contributed human resources are not recognized in these
financial statements.

6. Comparative Figures

Certain comparative figures have been reclassified where
necessary to conform to the presentation adopted in the cur-
rent year.

PREPARED WITHOUT AUDIT

TABLE OF CONTENTS (concluded) Volume 128, Number 3

Book Reviews

ZOOLOGY: Beetles of Eastern North America — Birds of the Kenya’s Rift Valley — The Amazing World of Flyingfish
-A Field Guide to the Larger Mammals of Tanzania — A Feathered River Across the Sky. The Passenger Pigeon’s

Flight to Extinction — A Sparrowhawk’s Lament — How British Breeding Birds of Prey Are Faring — Wildlife of the
Caribbean

BoTANy: Plants of Southern Ontario, Trees, Shrubs, Wildflowers, Grasses, Ferns and Aquatic Plants

OrnHer: Bird Lady — A Lifelong Love Affair with Birds — The Dismal State of the Great Lakes

New TITLES

Minutes of the 135" Annual Business Meeting of The Ottawa Field-Naturalists’ Club January
14, 2014

Mailing date of the previous issue 128(2): 8 July 2014

2014

314

THE CANADIAN FIELD-NATURALIST Volume 128, Number 3

Articles

Home range, movements, and denning chronology of the Grizzly Bear (Ursus arctos) in west-central
Alberta KAREN GRAHAM and GORDON B. STENHOUSE

Mortality of Common Eider, Somateria mollissima (Linnaeus, 1758), and other water birds during

two inshore oiling events in southeastern Newfoundland, 2005 and 2006
Grecory J. ROBERTSON, SCOTT G. GILLILAND, PIERRE C. RYAN, JOHANNE DUSSUREAULT,
KYRAN Power, and BRUCE C. TURNER

Impact of the 2012 drought on woody vegetation invading alvar grasslands in the Burnt Lands
Alvar, eastern Ontario PAUL M. CATLING

A review of colour phenotypes of the Eastern Red-backed Salamander, Plethodon cinereus, in North

America JEAN-DAVID MoorE and MARTIN OQUELLET

Notes

Long-distance anadromous migration in a fresh water specialist: the Lake Trout (Sa/velinus namaycush)
Les N. HARRIS, JEAN-SEBASTIEN MoorE, CHRISTOPHER G. MCDERMID, and HEIDI K. SWANSON

New records of the Ogilvie Mountains Collared Lemming (Dicrostonyx nunatakensis) in central
Yukon THOMAS S. JUNG, BRIAN G. SLOUGH, DAVID W. NAGORSEN, and PilA M. KUKKA

A note on bird song: Samuel Hearne’s observations on the Snow Bunting (Plectrophenax nivalis)
Davip L. G. NOAKEsS and JEFFREY D. NOAKES

Late-winter habitat use by the Fisher, Pekania pennanti (Erxleben, 1777), in the Boreal Plains

Ecozone of northwestern Saskatchewan, Canada GILBERT PROULX
Tributes
A tribute to Warren Baxter Ballard, 1947—2012 FRANCIS R. COOK
A naturalist for all seasons: Richard Merrill Saunders, 1904—1998 PHILIP COLLINS

2014

N
nN
Lo

243

250

260

265

269

242

276

289

(continued on inside back cover)

ISSN 0008-3550

MIX

r from
responsible sources

ike FSC® C100205

The CANADIAN
FIELD-NATURALIST

Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

Volume 128, Number 4 October—December 2014

ci - ,
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patron
His Excellency the Right Honourable David Johnston, C.C.,C.M.M., C.O.M.,C.M.
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these
fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environ-
ments of high quality for living things.
Honorary Members

Ronald E. Bedford Paul M. Catling John M. Gillett E. Franklin Pope
Edward L. Bousfield Francis R. Cook Peter W. Hall Joyce M. Reddoch
Charles D. Bird Bruce Di Labio Christine Hanrahan Allan H. Reddoch
Fenja Brodo Anthony J. Erskine C. Stuart Houston Dan Strickland
Irwin M. Brodo J. Bruce Falls Theodore Mosquin John B. Theberge
Daniel F. Brunton Robert W. Nero Sheila Thomson

Michael D. Cadman

2014 Board of Directors

President: Fenja Brodo Daniel F. Brunton Barry Cottam Lynn Ovenden
15‘ Vice-President: Henry Steger Carolyn Callaghan Don Hackett Rémy Poulin

2" Vice-President: Eleanor Zurbrigg Barbara Chouinard Karen McLachlan Hamilton Eleanor Zurbrigg
Recording Secretary: Annie Bélair Julia Cipriani Diane Kitching

Treasurer: Ken Young Owen Clarkin Ann MacKenzie

To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate
P.O., Ottawa, ON, K1Z 1A2, or e-mail: ofnc@ofne.ca. For information on Club activities, go to www.ofnc.ca

The Canadian Field-Naturalist

The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this
journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.

Website: www.canadianfieldnaturalist.ca/index .php/cfn

Editor-in-Chief: Dr. Carolyn Callaghan Assistant Editor: Dr. Trina Rytwinski
Copy Editor: Sandra Garland Typographer: Wendy Cotie
Journal Manager: Dr. Jay Fitzsimmons: subscriptions @canadianfieldnaturalist.ca or via postal address above.

Book Review Editor: Roy John

Associate Editors: — Charles D. Bird Anthony J. Gaston David Nagorsen Jeffery M. Saarela
Paul M. Catling Thomas S. Jung Claude B. Renaud David C. Seburn
Francis R. Cook Donald F. McAlpine Marty Obbard Jeffrey H. Skevington
Jennifer R. Foote Garth Mowat

Chair, Publications Committee: Daniel F. Brunton

All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by e-mail:
editor @canadianfieldnaturalist.ca or postal mail (with disk): 611 Chemin Cregheur, Luskville, QC JOX 2G0; (819) 455-1087:
Book-review correspondence should be sent to the Book Review Editor by e-mail: r.john@rogers.com or postal mail: 2193
Emard Crescent, Ottawa, ON, K1J 6K5

Subscriptions and Membership:
Subscription rates for individuals are $40 (online only), $50 (print only), or $60 (print + online). Libraries and other institutions may
subscribe for $60 (online only or print only) or $90 (print + online). All foreign print subscribers and members (including USA)
must add $10 to cover postage. The Ottawa Field-Naturalists’ Club annual membership fee of $40 (individual), $45 (family), or
$20 (student) includes an online subscription to The Canadian Field-Naturalist. Members can receive printed issues of CEN for
an additional $30 per volume (four issues). For further details, including discounts for members of the Canadian Museum of Nature,
see http://ofnc.ca/member.php. The club’s regional journal, Trail & Landscape, covers the Ottawa District and regional Club
events and field trips. It is mailed to Ottawa area members, and is available to those outside Ottawa on request. It is available to
libraries at $33 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should
be sent to subscriptions @canadianfieldnaturalist.ca or mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069. Wests

ate
P.O., Ottawa, ON, KIZ 1A2 Canada. Canada Post Publications Mail Agreement number 40012

317. Return postage guaranteed.

Cover: A female Misumena vatia on a yellow lady’s slipper Cypripedium parviflorum in the Burnt C

ape Ecological Reserve.
Newfoundland. Photo by R. Perry. See pages 363-376 in this issue. : a avin

MCZ
LIBRARY

The Canadian Field-Naturalist FEB 20 205

Volume 128, Number 4

OctobersDecember ae
CIV ERS iry

The Enigma of the 10-year Wildlife Population Cycle Solved?

Evidence that the Periodicity and Regularity of the Cycle Are Driven
by a Lunar Zeitgeber

HERBERT L. ARCHIBALD
632 Tee Hi Place, Medford, Wisconsin 5445] USA; email: Herb. Archibald@gmail.com

Archibald, Herbert L. 2014. The enigma of the 10-year wildlife population cycle solved? Evidence that the periodicity and
regularity of the cycle are driven by a lunar zeitgeber. Canadian Field-Naturalist 128(4):; 327-340.

Despite nearly 100 years of research, the periodicity and regularity of the 10-year wildlife population cycle remain an enigma.
This paper presents the hypothesis that the 9.3-year nodal half-cycle of the moon is the zeitgeber (“time-giver’) of the 10-year
wildlife population cycle. The period of the population cycles of the Snowshoe Hare (Lepus americanus) and Ruffed Grouse
(Bonasa umbellus) is close to 9.3 years. These wildlife cycles have stayed closely in phase with the 9.3-year nodal half-cycle for
150 years. Population density of the Snowshoe Hare and Ruffed Grouse is inversely related to a 9.3-year cycle of the moon’s
tidal force. There is also a 9.3-year cycle of “nights without darkness” at the equinoxes, in which the full moon rises before
sunset and sets after sunrise the following morning in certain years. Snowshoe Hare and Ruffed Grouse cycles are positively
correlated with this phenomenon. The nodal cycle provides explanations for the key features of the 10-year wildlife cycle:
regularity, periodicity, amplitude, distribution, and synchrony. Population models based solely on the nodal cycle account for
62% of the variation in the Canada Lynx (Lynx canadensis) cycle and 37% in the Ruffed Grouse cycle. The mechanism(s) by
which herbivore cycles might be entrained by the lunar nodal cycle could involve a cyclic effect on factors including predation,
stress, photoperiod, phenology, temperature, cloudiness, ultraviolet B radiation, cosmic rays, and food plant quality.

Key Words: 10-year wildlife cycle; lunar nodal cycle; Snowshoe Hare; Lepus americanus; Ruffed Grouse; Bonasa umbellus;

Canada Lynx; Lynx canadensis

Introduction

The 10-year wildlife cycle has been described as an
“extraordinary precise metronome” by ecological, if not
celestial, standards (King and Schaffer 2001). Yet, des-
pite nearly 100 years of research, the periodicity and
regularity of the 10-year wildlife cycle remain an enig-
ma. “The key question of how the 10-year cyclic pattern
is produced is still unsolved” (Yan ef a/. 2013).

Numerous hypotheses, involving both endogenous
(e.g., disease, predation, food abundance) and exoge-
nous (e.g., weather, sunspots, ultraviolet radiation) caus-
es, have been proposed to explain the cycle (see sum-
maries in Murray [2003] and Zimmerman et al. [2008]).
Cyclic declines in Ruffed Grouse (Bonasa umbellus)
have been attributed to increased predation because of
a predator shift following decreases in Snowshoe Hare
(Lepus americanus) numbers (Keith 1963), but this
hypothesis has been challenged because Ruffed Grouse
tend to peak and decline before Snowshoe Hare ( Hoff-
mann 1958). The prevailing endogenous hypothesis
is that the Snowshoe Hare cycle results from the tri-
trophic interaction between predation and food sup-
plies, with predation as the dominant factor (Krebs et
al. 2001). In his review, Rusch (1989) concluded that
predation, primarily avian, is responsible for virtually all
grouse mortality and the cyclic trend in grouse popula-
tions. In their classic paper, Elton and Nicholson (1942)
concluded that the broad synchronization of the Canada
Lynx (Lynx canadensis) cycle “makes it certain that
some overriding process maintains the cycle in line over

the whole extent of Canada.” In his plant stress hypoth-
esis, Selas (2006) posited that the Snowshoe Hare cycle
is forced by the sunspot cycle, through its effects on
plant productivity caused by changes in ultraviolet B
(UVB) radiation. Sinclair et al. (1993) hypothesized that
Snowshoe Hare cycles are synchronized by the sunspot
cycle via its effect on climatic factors. However, a
causal relationship between sunspots and the Canada
Lynx cycle has been rejected on a statistical basis
(Moran 1949; Lindstrom et al. 1996). Nilssen ef ai.
(2007) demonstrated that the 11-year sunspot cycle can-
not explain the 10-year cycle of the Autumnal Moth
(Epirrita autumnata) in Fennoscandia because the two
cycles ran in-phase and then completely out of phase
during a 114-year period.

Murray (2003) suggested the possibility that an
exogenous factor could entrain or synchronize a cycle
that is driven by endogenous forces. Yan ef a/. (2013)
concluded that the 10-year hare—lynx cycle is the result
of the joint forces of both endogenous and exogenous
factors. The ideal exogenous cycle candidate would
have a consistent period that matched the wildlife cycle
and would shed light on its other unexplained features.

Snowshoe Hare and Canada Lynx populations tend
to cycle synchronously across the boreal forests of
Canada and Alaska (Krebs ef al. 2001). Recent expla-
nations of the spatial synchrony of 10-year-cycle species
across large geographic regions have generally focused
on common weather or climatic perturbations (i.e.,
Moran effect), dispersal, migration of natural enemies

327

(Liebhold e¢ al. 2004), or a combination thereof. Krebs
et al. (2013) concluded that the mobile predator hypoth-
esis offers the most likely explanation of the synchrony
and traveling waves of hare populations across north-
western North America. Analysis of the spatial syn-
chrony of Canada Lynx populations revealed three
large climatic regions in which the phase synchrony is
similar, with the continental region |—2 years ahead of
the Pacific and Atlantic (Stenseth ef al. 1999, 2004).
These authors concluded that external climatic forcing
associated with the North Atlantic Oscillation acts as a
synchronizer within each region, but they offer no expla-
nation for the inter-regional timing of the cycle. Krebs
et al. (2013) rejected the “attractive” hypothesis that
regional weather synchronizes local Snowshoe Hare
populations because of the lack of a mechanism where-
by weather could drive hare numbers.

The path of the moon across the sky is complicat-
ed by its nodal cycle. The 18.6-year lunar nodal cycle
gives rise to year-to-year differences in both tidal force
and the moon’s “altitude” (angle or height in the sky
above the horizon). The tidal force of the moon is the
difference between the moon’s gravitational force on
the side of the earth closest to the moon and that on the
side farthest from the moon (Kutner 2003). The moon
and the sun produce maximum tidal forces at the
equinoxes when lunar “declination” (angle relative to
the equator) is zero, which happens twice during the
nodal cycle or every 9.3 years, and when the sun is
directly above the equator, so that solar declination is
zero (Pugh 1966). Because the moon’s orbit is inclined
5.1° to the earth’s orbit (also known as the “ecliptic’’),
the declination of the moon’s orbit ranges from a mini-
mum of 18.3°N and S every lunar month to a maximum
of 28.6°N and S every month during the 18.6-year peri-
od of the nodal cycle. What the observer at 55°N lati-
tude would see 1s variation in the moon’s altitude from
as little as a high of 53.5° and a low of 16.5° two weeks
later (monthly swing of 37°) to, 9.3 years later, as much
as a high of 63.5° and a low of 6.5° (monthly swing of
57)

Although the nodal cycle has been known at least as
far back as Stonehenge, the biological significance of
the 9.3-year nodal half-cycle remains virtually un-
known. Archibald (1977) hypothesized that the length
and regularity of the period, approximate regional and
inter-specific synchrony, and northern geographic dis-
tribution of the 10-year wildlife population cycle arise
from the 9.3-year half period of the moon’s 18.6-year
nodal cycle. This hypothesis has been largely ignored
or dismissed as a nonsense correlation (Royama 1992),
perhaps because Archibald did not provide any evi-
dence of biological effects of the 9.3-year lunar cycle
or plausible mechanism. Selas (2014) proposed a new
variation of the plant stress hypothesis involving mod-
ulation of cosmic rays by a 9.3-year cyclic “lunar index”
(i.e., maximum declination cycle) and the 11.1-year
sunspot cycle. According to Selas, a low lunar index

THE CANADIAN FIELD-NATURALIST

Vol. 128

(i.e., maximum tidal force) increases geomagnetic
activity, which reduces incoming cosmic ray particles,
which increase protein mobilization and reduce produc-
tion of secondary metabolites in plants, thereby increas-
ing herbivore forage quality and, hence, enhancing her-
bivore performance.

The objective of this paper is to present evidence
supporting the hypothesis that the 9.3-year half period
of the lunar nodal cycle is the zeitgeber (literally “time-
giver”) of the 10-year wildlife cycle and to speculate on
mechanisms by which herbivore cycles might be en-
trained by the lunar nodal cycle.

Methods

I compiled reports of peaks in Snowshoe Hare and
Ruffed Grouse abundance (Table 1) from nine sets of
data (Table 2) of several types: density surrogates, such
as annual fur returns (MacLulich 1957) or spring drum-
ming counts (Williams e¢ al. 2004; Minnesota DNR
2011); simple lists of peak years in an area (Shorger
1947); and lists of peak abundance for multiple loca-
tions (Keith 1963; Hodges 2000a,b). In decades where
there were two or three consecutive years of similarly
high population reported in a dataset or occurring in a
species, the median was used as the value of the peak
year. None of these datasets is precisely accurate; all
have flaws and biases of various kinds. Also, peak years
are not directly comparable across species because data
are based on different types of records and are from dif-
ferent regions. Nevertheless, the mean peak years com-
puted by averaging across species provide a broad-
scale, general picture of the timing of cycle peaks over
large geographic areas during the 150-year period cov-
ered.

Predicted peak years of abundance in Table | were
computed using the formula P, = 1950.22 + 9.3n,
where P_ is the nth peak year after or before 1950.22
(date of the vernal equinox of 1950 when lunar decli-
nation reached a peak value, which happens only once
in each 18.6-year nodal cycle) and 9.3 is the period
of the population cycle (Archibald 1977).

Sun and moon rise and set times were obtained from
the Astronomical Sky Calendar (www.briancasey.org
/artifacts/astro/skycalendar.cgi) and from the United
States Naval Observatory’s Complete Sun and Moon
for One Day website (http://aa.usno.navy.mil/data/docs
/RS_OneDay.php). The number of minutes the full
moon was above the horizon at both dusk (moonrise
before sunset) and the following dawn (moonset after
sunrise) were computed for April and September. Lunar
declination data were obtained from the Jet Propulsion
Laboratory Horizons Web-Interface (http://ssd.jpl.nasa
.gov/horizons.cgi#top). Over the course of the lunar
nodal cycle, the moon’s declination varies about 5.1°
above and below the ecliptic. For each year, the ab-
solute value of the maximum declination from the eclip-
tic in the equinox month of September was determined
at SO°N, 90°W, i.e., absolute value (reported declina-

2014

ARCHIBALD: THE ENIGMA OF THE 10-YEAR WILDLIFE POPULATION CYCLE SOLVED?

329

TABLE |. Reported < aclistecinanlesitinia : :
. Reported and predicted peaks in abundance of the Showshoe Hare (Lepus americanus) and the Ruffed Grouse (Bonasa

umbellus) in northern North America.

00200000

Reported peak years of abundance

Mean peak

Snowshoe Hare Ruffed Grouse

year
2006.0 2009.0 2007.5
1998.0 1998.5 1998,2
1989.2 1989.0 1989.1
1980.0 197 Or 1979.6
1970.8 1971.8 1971.3
1959.8 1959.5 1959.6
LOSS 1949.2 1950.4
1941.6 1941.2 1941.4
1932.2 1932.6 1932.4
1923.6 1922.6 19231
1912.4 1914.2 1913°3
1904.1 1904.6 1904.4
1896.0 1898.0 1897.0
1886.5 1887.0 1886.8
1876.0 1877.0 1876.5
1865.0 1866.0 1865.5
1857.0 1857.0 1857.0
Mean (SD)

Mean period 25 a5

Note: SD = standard deviation.
*Between reported and predicted peak years.

Predicted Difference*
peak years (years)
2006.0 1.3
1996.7 15
1987.4 ey
1978.1 1.5
1968.8 25
1959.5 0.1
1950.2 0.2
1940.9 0.5
1931.6 0.8
1922.3 0.8
1913.0 0.3
1903.7 0.7
1894.4 2.6
1885.1 ey
1875.8 0.7
1866.5 lo)
1857.2 —0.2
0.9 (0.91)
9.4

TABLE 2. Sources of the reports of peak abundance listed in Table 1.

Species Date range Reference

Snowshoe Hare 1857-1899 MacLulich (1957: 296)
1900-1955 _ Keith (1963: 42)
1956-1992 Hodges (2000a: 119-120
1956-1992 Hodges (2000b: 166-168)
1993-2011 Krebs (2011: 6)

Ruffed Grouse 1857-1899 — Schorger (1947: 54)
1900-1951 Keith (1963: 57)
1949-2010 Minnesota DNR (2011: 45)
1956-2001 Williams ef al. (2004: 1138)

Location Source type

Fur returns

Peak abundance
(median years)
Peak abundance
(median years)
Peak abundance
(median years)
Density on study
area

Peak years of
abundance

Peak abundance
(median years)
Drumming counts
Drumming counts

Hudson’s Bay Watershed
Northern North America

Canada, Alaska

Minnesota, Wisconsin, Manitoba
Yukon Territory

Wisconsin

Northern North America

Minnesota
Northern Wisconsin, North Dakota

ee oOoOoOoOoqoqqowoaomnmS— ————__—qv

tion -23.5°). September maximum lunar declination
and April and September total minutes full moon above
horizon were compared for two population datasets:
Ruffed Grouse spring drumming counts conducted in
Minnesota from 1949 to 2010 (Minnesota DNR 2011)
and spring Snowshoe Hare density in the Kluane region
of Yukon Territory (Krebs 2011).

Population modeling and preparation of figures were
done using SciDaVis (http://scidavis.sourceforge.net/)
software. Modeling results were compared with 1949—
2010 Minnesota drumming counts and 1842-1918
Canada Lynx fur returns from the Northern Department
(Elton and Nicholson 1942). The classic lynx time series

is arguably the best example of a near-perfect popula-
tion cycle with a minimal amount of noise. This dataset
was not used, however, for the peak years of abundance
analysis (Table 1) because populations of Canada Lynx
are not independent of populations of their main prey,
the Snowshoe Hare.

Correlation between population size and lunar dec-
lination was measured with least squares regression
using software supplied by Wessa (Free Statistics Soft-
ware, Office for Research Development and Education,
version 1.1.23-r7, www.wessa.net), even though the
assumption of independence of observations in each
time series is clearly violated (Liebhold er a/. 2004).

330

The inadequacy of currently available correlational
measures was pointed out by Liebhold e¢ al. (2004):
“for cyclic populations, subtle differences in the tim-
ing of cycles... may result in low statistical correlation
even when populations are closely linked.”

Results

The period of the cycles of the Snowshoe Hare and
Ruffed Grouse shown in Table | is close to 9.3 years:
9.3 and 9.5 years, respectively. Selas (2014) demon-
strated that the periods of both the Canada Lynx and the
Autumnal Moth cycles are very close to the 9.3-year
lunar nodal half-cycle. One point that seems to have
been overlooked in the literature, however, is how crit-
ical the period length is. For example, if you spin back-
ward from 2006.0 (when lunar declination reached a
peak value seen only once in each 18.6-year nodal cy-
cle) using the 9.6-year period suggested by Elton and
Nicholson (1942) and Krebs et al. (2001), the predicted
peak year 15 cycles back is 1862.0, which is almost
completely out of phase with the lunar peaks in 1857.2
and 1866.5. In contrast, after 15 cycles back using the
9.3-year period, reported and predicted peaks remain in
phase, with the mean herbivore peak of 1857.0 coincid-
ing with the lunar peak at 1857.2. To become complete-
ly out of phase, a difference of 0.2 years/cycle (i.e., a
period of 9.1 or 9.5 years) takes about 23 cycles (or
about 214 years) and a difference of 0.1 years/cycle
(i.e., a period of 9.2 or 9.4 years) takes about 46 cycles
(or about 428 years); sufficient data are not available
to evaluate either of these cases.

For over 150 years, reported years of peak abun-
dance of Ruffed Grouse and Snowshoe Hare have per-
sistently stayed in phase with predicted peak years
based on the lunar model P, = 1950.22 + 9.3n (Archi-
bald 1977), differing by an average of 0.9 years (Table
1). Differences between mean reported and predicted
peak years appear to have become larger after about

t a @
Oo o o

Full moon above horizion (minutes/day)
N
oOo

1960

THE CANADIAN FIELD-NATURALIST

Vol.

1965, with the reported peaks consistently lagging be-
hind the predicted peaks.

9. 3-year lunar declination cycle

The 9.3-year cycle in the maximum lunar declination
from the ecliptic in absolute value at the equinoxes Is
shown in Figure |. The alternating higher declination
peaks are years when the moon’s maximum declination
runs below the ecliptic. Semidiurnal (result in two high
tides and two low tides per day) and diurnal (result in
one high and one low tide per day) tidal factors oscillate
in 18.6-year cycles, 180° out of phase, with diurnal fac-
tors largest when lunar declination is highest (1950,
1968, 1987, 2006) and semidiurnal factors highest
when lunar declination is minimal (1959, 1978, 1997)
(McKinnell and Crawford 2007). The semidiurnal lunar
forces are 23% less when the moon reaches its maxi-
mum declination of 28.6°. The solar semidiurnal forces
are reduced by 16% when the sun is at its maximum
declination of 23.5° at the solstices (Pugh 1996). To
sum up, tidal forces reach a maximum when the moon
is in the equatorial plane (i.e., at zero declination from
the ecliptic), which happens twice during the 18.6-year
nodal cycle or every 9.3 years.

9.3-year moonlight cycle

The full moon can be above the horizon both before
dusk and after dawn the following morning, resulting in
a “night without darkness” with light levels no lower
than full moonlight. This can occur on two successive
nights and in several months in a row. The nodal cycle
causes an 18.6-year cycle of total minutes that the full
moon is above the horizon both before dusk and after
the following dawn at the full moon nearest the autum-
nal equinox (light bars in Figure 1). The westward
regression of the lunar nodes from the autumnal equi-
nox to the vernal equinox takes 9.3 years. Then, there is
a corresponding 18.6-year cycle of total minutes that
the full moon is above the horizon both before dusk and

te uw a ~N

(saaibap) uojjeul.ep weuny

w

—

FIGURE 1. Length of time full moon (April, dark shading; September, light shading) was above the horizon both before
dusk and after the following dawn (total minutes/day) and maximum lunar declination (open circles) in September

relative to the ecliptic in absolute value (SO°N, 90°W).

2014

after the following dawn during the full moon nearest
the vernal equinox (dark bars in Figure |). Therefore,
this phenomenon occurs every 9.3 years, alternating be-
tween the vernal and autumnal equinoxes. Nights with-
out darkness occur multiple times in the years at or near
the peak of the cycle. For example, in the September
peak year 1991, there were nine nights without dark-
ness: July 26, August 24-25, September 23-24, Octo-
ber 22~23, November 21, and December 20. Although
the midnight sun near the summer solstice at higher lat-
itudes is well known, there appears to be nothing in the
literature about this nights-without-darkness phenom-
enon at the equinoxes. The only information I have
found so far about “sun and full moon both up” are
some anecdotal reports of confused homing pigeons
flying in the wrong direction, perhaps because they mis-
took the rising full moon for the sun. The 9.3-year
moonlight cycle is inversely related to the 9.3-year
lunar declination cycle (Figure 1).

Correlations with 10-year wildlife population cycles
Ruffed Grouse drumming counts in Minnesota are
positively correlated (7 = 0.26, P < 0.001, with 1-year

a ™N

ui

Ww

N

Lunar declination (degrees)
Q
CO

1950

ARCHIBALD: THE ENIGMA OF THE 10-YEAR WILDLIFE POPULATION CYCLE SOLVED? 331

lag) with September maximum lunar declination (Fig-
ure 2) and, therefore, inversely related to the moon’s
tidal force. During the equinox months of September
and March, full and new moons always coincide with
minimum lunar declination (moon over the earth’s
equator) (Brahde 1988). There is some tendency for
grouse peaks to alternate in amplitude (i.e., every other
peak is a “high” peak) (Archibald 1977). High grouse
peaks are associated with years when the moon’s max-
imum declination runs above the ecliptic. Because the
zeitgeber effect of the 9.3-year nodal half-cycle “need
not occur in more than 2—3 consecutive years per
decade” (Archibald 1977), interpretation of correlation
coefficients (which cover all years) is problematic.

Snowshoe Hare density in the Yukon Territory is also
directly correlated (77 = 0.41, P < 0.001, with 2-year
lag) with September maximum lunar declination and
inversely correlated with the moon’s tidal force (Figure
3). There is also a tendency for hare population peaks
to alternate in amplitude (Archibald 1977); higher peaks
coincide with years when the moon’s maximum dec-
lination runs below the ecliptic.

FIGURE 2. Maximum lunar declination (open circles) in September relative to the ecliptic in absolute value (S50°N, 90°W) com-
pared with spring drumming counts (closed circles) for Ruffed Grouse (Bonasa umbellus) in Minnesota, 1949-2010

(Minnesota DNR).

n

“

nw

Lunar Declination (degrees)

~

t)
1975 1960 1985 1990

Year

Figure 3. Maximum lunar declination in September (open circles
pared with spring Snowshoe Hare (Lepus americanus)

Territory, 1977-2010 (Krebs 2011).

Aysueg 22H

1995 2000 2005 2010

) relative to the ecliptic in absolute value (S0°N, 90°W) com-
density (closed circles) in the Kluane region of the Yukon

2
Lo
NR

The cyclic nights-without-darkness phenomenon at
the equinoxes generally starts to increase |—2 years
before the Snowshoe Hare peak; it peaks during the
hare decline and stays relatively high throughout the
hares’ low phase (Figure 4). The inverse relation be-
tween Snowshoe Hare density and nights without dark-
ness during the low phase is intriguing because expla-
nation of the low phase remains a puzzle (Krebs ef al.
2001). Similar to hare density, nights without darkness
generally increase following an increase in Ruffed
Grouse density and stay relatively high during the ensu-
ing low (Figure 5).

Ruffed Grouse population model

The following model of Ruffed Grouse drumming is
based on a single exogenous factor, the nodal cycle of
the moon:

Drums/stop(f) = 1.27 + 0.55sin(2z )/9.307 * (t— 1) + 1.41)
+ 0.55sin(27 )/18.6134 * (f—1)+ 0.907)

Where ¢ is the year (yyyy) with l-year lag; 9.307
and 1.41 are the period and phase angle of the nodal

Full moon above horizon (minutes/day)
N > oa]
Oo oO o

oOo

1980 1985 1990

THE CANADIAN FIELD-NATURALIST

Vol. 128

phase cycle, respectively; 18.6134 and 0.90 are the peri-
od and phase angle of the nodal amplitude cycle (Yn-
destad 2009); 1.27 is the amplitude; and 0.55 is half
the range of observed Minnesota drumming counts.

The model output matches the observed Ruffed
Grouse cycle fairly well (7? = 0.39, P < 0.001), in both
period and alternating amplitude (Figure 6).

Canada Lynx population model

Furs/year(t) = 20800 + 16540sin(21 /9.307 * ¢ + 1.417)
+ 16540sin(2z /18.6134 * (t-9.307) + 0.90z )

Where ¢ is the year (yyyy) with no lag on the 9.3-
year phase cycle and a lag of 9.307 years on the 18.6-
year amplitude cycle (to offset the higher peaks); 9.307
and 1.41 are the period and phase angle of the nodal
phase cycle, respectively; 18.6134 and 0.90 are the peri-
od and phase angle of the nodal amplitude cycle (Ynde-
stad 2009), 20800 is the amplitude and 16540 is half
the range of observed Northern Department lynx fur
returns between 1842 and 1918 (Elton and Nicholson

1942).

Ajsueap ae}

Ye Sd ‘ 0

1995 2000 2005

Year

FiGure 4. Length of time full moon (April, dark shading; September, light shading) was above the horizon both before dusk
and after the following dawn (total minutes/day; 50°N, 90°W) compared with spring Snowshoe Hare (Lepus americanus)
density (closed circles) in the Kluane region of the Yukon Territory, 1977-2010 (Krebs, 2011).

&

B

8

8

&

| ®

Full moon above horizon (minutes/day)
8

°

1970

1980

1990

daqs/swinug

2000

Year

FIGURE 5. Length of time full moon (April, dark shading; September, light shading) was above the horizon both before dusk
and after the following dawn (total minutes/day; 50°N, 90°W) compared with spring drumming counts (closed circles)
for Ruffed Grouse (Bonasa umbellus) in Minnesota, 1949-2010 (Minnesota DNR 2011). -

2014 ARCHIBALD: THE ENIGMA OF THE 10-YEAR WILDLIFE POPULATION CYCLE SOLVED? 333

1950 1960 1970

1990 2000 2010

FIGURE 6. Spring drumming counts (open circles) predicted by the lunar model compared with observed drumming counts
(closed circles) for Ruffed Grouse (Bonasa umbellus) in Minnesota, 1949-2010 (Minnesota DNR 2011).

ONE aad

3

7

FIGURE 7. Canada Lynx (Lynx canadensis) fur returns predicted by lunar model (open circles) compared with observed Canada
Lynx fur returns (closed circles), Northern Department, 1842-1918 (Elton and Nicholson 1942).

The model output matches the “classic” Canada
Lynx cycle quite well (77 = 0.62, P < 0.001) in both
period and alternating amplitude over the period 1842—
1918 (Figure 7). Note that additional decimal places in
the period values were used in the Rugged Grouse and
Canada Lynx population models because the formulas
are very sensitive to small differences in period length.

Discussion
Key cycle features accounted for

King and Schaffer (2001) list five important features
of the 10-year wildlife cycle, and a definitive explana-
tion for the cycle should account for all five points. The
9.3-year half period of the nodal cycle hypothesis pro-
vides a plausible explanation of each of these five
features:

1. Regularity: Reported peak population years have
stayed in phase with predicted peaks based on
the 9.3-year lunar cycle for over 150 years (Table
1).

2. Period: The overall mean cycle period (9.4 years)
is close to the 9.3-year period of the lunar cycle
(Table 1).

3. Amplitude: Peaks in the Snowshoe Hare cycle
tend to alternate in amplitude (Archibald 1977).

The trend is evident in the Hudson’s Bay data
from 1857 to 1896 (MacLulich 1957) and a cen-
tury later at Kluane Lake (Figure 3). Schaffer
(1984) noted alternating amplitude in the Canada
Lynx cycle from 1821-1913. Higher peaks in the
hare-lynx cycle are associated with vernal equi-
nox peaks in the lunar cycle (Figure 4) and max-
imal semidiurnal tidal factors. Ruffed Grouse
peaks also tend to alternate in amplitude but high-
er peaks in the grouse cycle are associated with
autumnal equinox peaks in the lunar cycle (Figure
5), when the moon’s maximum declination runs
above the ecliptic (Figure 2) and diurnal tidal fac-
tors are largest.

Distribution: The 10-year cycle is a northern
phenomenon restricted to latitudes above 45-
50°N in North America (Archibald 1977) and
even higher latitudes in northern Europe. North-
ern populations of Snowshoe Hares and Ruffed
Grouse are cyclic, while more southerly popula-
tions exhibit little or no periodicity (Rusch 1989;
Murray 2003). Several effects of the lunar nodal
cycle increase significantly with latitude. For
example, the full moon was above the horizon at
both dusk and dawn on the night of April 29-30,

1999, for a total of 71 minutes at 40°N, 90 min-
utes at SO°N and 131 minutes at 60°N. Latitude is
an important factor governing both the species
(semidiurnal and diurnal) and maximum ampli-
tude of tides (Pugh 1996).

5. Synchrony. Regional synchrony of populations
over wide geographic areas has been reported for
both Snowshoe Hare and Ruffed Grouse (Keith
1963), and population levels of these species tend
to be closely aligned within the same areas, with
peaks predominantly | year apart (Keith 1963).
Selas (2006) found transatlantic synchrony in
Snowshoe Hare/Mountain Hare (Lepus timidus)
and Canada Lynx/Eurasian Lynx (Lynx lynx)
series, suggesting that the cycles are triggered by
a common large-scale phenomenon. Synchrony
is to be expected under a lunar cycle hypothesis,
because lunar phenomena, such as height in the
sky, rise and set times, and gravitational pull are
similar worldwide at a given latitude, varying
only because of such factors as elevation and
topography.

Population modeling

King and Schaffer (2001) developed a tri-trophic
population dynamics model of the Snowshoe Hare
cycle with browse, hare, and predator density as de-
pendent variables and 14 parameters (but no weather
factors). They concluded that the increase phase is halt-
ed by food limitation and the decline from peak num-
bers is due to predation. Yan et al. (2013) used two-
level population models with rate of change of hare
and lynx as dependent variables and either global weath-
er indices or local temperature, rainfall, and snowfall
with various time lags. Their simulations showed that
predation and direct, as well as delayed, density de-
pendency are necessary, but not sufficient, factors for
the appearance of 10-year hare—lynx cycles and that
external climatic forcing through rainfall and snowfall
are essential to produce 10-year cycles.

Zimmerman et al. (2008) used comparative model
selection to assess the correlation of numerous combi-
nations of 13 hypothesized factors with spring drum-
ming counts for grouse in northeastern Minnesota over
a 22-year period. The hypothesized factors included
goshawk abundance (predation), Eastern Tent Cater-
pillar (Malacosoma americanum [Fabricius, 1793])
abundance (aspen defoliation), weather during the
breeding season, weather during the previous winter,
colour phase ratios (phenotypic hypothesis), fall age
ratios (recruitment), and male grouse weight (condi-
tion entering winter). The best model indicated that
grouse abundance was highest during cold snowy and
warm dry winters, presumably related to optimal snow-
roosting conditions during periods of extreme cold. As
the best model accounted for only 17% of the variation
in drumming counts, these authors concluded that sev-
eral interacting factors most likely cause the cycle but
the mechanism remains unknown.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Using a multiple regression model, Selas (2014)
found that a maximum September lunar declination
index (3-year lag), March-April North Atlantic Oscil-
lation, previous August-October North Atlantic Oscil-
lation and sunspot numbers accounted for 38% of the
variation in an Autumnal Moth index from Fennoscan-
dia covering 113 years. A similar model, which used the
Pacific-North American index instead of the North
Atlantic Oscillation, accounted for 23% of the variation
in a Canada Lynx population index for the period 1950—
2009.

The Canada Lynx model presented here, with just
the 9.3- and 18.6-year lunar parameters, accounted for
62% of the variation in the Northern Department fur
returns over 77 years (Figure 7). The Ruffed Grouse
model presented here accounted for 37% of the varia-
tion in the Minnesota Ruffed Grouse drumming counts
over 62 years (Figure 6). This suggests that both the
9.3-year and 18.6-year nodal cycles play a significant
role in lynx and grouse population dynamics. A predic-
tion model with four lunar parameters including both
the 9.3 and 18.6-year nodal periods accounted for 62%
of the variation in strong-to-violent March—August
tornadoes in the United States between 1950 and 2012
(Eppelbaum 2013). Schaffer’s (1984) power spectrum
analysis of Canada Lynx fur returns (1821-1913)
showed a fundamental period of roughly 10 years and
a second harmonic period of about 20 years. The obvi-
ous similarity in periods suggests that the “underlying
determinism” this author sought could well be the 18.6-
year nodal cycle and its 9.3-year sub-harmonic. Further
research directed toward gaining an understanding of
the interaction of the 9.3-year and 18.6-year nodal cy-
cles is clearly needed. Also apparent from these model
results is that, while density-dependence may well play
a part in nature and is a critical component of prevail-
ing models of cyclic populations, it is not necessary to
closely mimic the cyclic population dynamics of the
Canada Lynx and Ruffed Grouse.

Spatial synchrony

Regional synchrony is expected under a lunar hy-
pothesis, and the problem to be reconciled is why the
synchrony is not absolute (Krebs ef a/. 2001). Some
asynchrony can be expected between even nearby pop-
ulations because of differences in local conditions.
Selas (2006) suggested that spatial asynchrony in hare
cycles could result from asynchronous fluctuations in
plant chemistry due to regional variations in UVB lev-
els caused by differences in cloud formation or ozone
distribution. If the zeitgeber effect of the lunar nodal
half-cycle occurs for 3-4 years at the bottom of the
cycle, more regional variation in the timing of the sub-
sequent peak would be expected because of differences
in local environmental conditions and food webs. But
this explanation does not account for the tendency for
synchrony of Snowshoe Hare populations to spread out
from a north-central Canada in traveling waves.

2014

Smith (1983) found that peaks in the Snowshoe Hare
cycle began in a central Canada core area (Saskat-
chewan and Manitoba) and traveled outward in waves
with |-year and then 2-year phase lags in the peak year
occurring with increasing distance. Selas (2014) found
that the increasing phase lag of Canada Lynx popula-
tion peaks moving away from the core area was signif-
icantly negatively related to cosmic radiation levels.
In other words, Canada Lynx tend to peak first in areas
of highest radiation (i.e. the “core area of the auroral
oval”). Although it may be a contributing factor, the
cosmic ray hypothesis seems an unlikely overall expla-
nation for a metronomic population cycle because, first,
the 9.3-year lunar cycle and the 11.1-year solar cycle
will go completely in and out of phase every 103 years
and, second, Selas found that the solar signal had a
much stronger relation to geomagnetic activity than the
lunar signal.

The earlier peaking in the core area could be due to
the generally flatter topography of central Canada if the
lunar zeitgeber effect on the Snowshoe Hare involves
nights without darkness. The shadow effect of moun-
tainous and hilly terrain can be expected to produce lat-
er actual rise and earlier actual set times resulting in
nights without darkness occurring in fewer consecutive
years (hence the lags) and with less biological impact.

Regarding the search for a mechanism, one initial
question that arises is whether there is a single mech-
anism underlying all 10-year wildlife population cycles.
Several observations suggest not. The 9.3-year cycles
of the Autumnal Moth in Fennoscandia and the Douglas
Fir Tussock Moth (Orgyia pseudotsugata [McDun-
nough, 1921]) in the Pacific Northwest are almost com-
pletely out of phase with the Snowshoe Hare and Ruffed
Grouse cycles in North America (Archibald, unpub-
lished data; Selas 2014), strongly suggesting a differ-
ent mechanism. Also, the difference in peak amplitude
alternation between the Snowshoe Hare and Ruffed
Grouse cycles could result from a different mechanism.

“Correlation means little unless we see a plausible
causal relationship” (Royama 1992). The two 9.3-year
lunar cycles reported here provide the basis for specu-
lative mechanistic explanations of the 10-year wildlife
cycle. Other 9.3-year cycles resulting from the nodal
cycle may await discovery. The 9.3-year moonlight
cycle could have behavioural, physiological, chrono-
biological, or phenological effects. The 9.3-year decli-
nation cycle could have an impact on cyclic population
dynamics through effects of oceanic and atmospheric
tides on climatic factors, such as cloudiness and UVB
and cosmic radiation, which, in turn, affect food plant
quality.

Ruffed Grouse population peaks tend to follow max-
imum lunar declination by 1—2 years (Figure 2). And
grouse lows are closely associated with minimum lunar
declination, suggesting that the declination effect could
be on food quality or palatability rather than directly on
the grouse itself. The primary winter food of northern

ARCHIBALD: THE ENIGMA OF THE 10-YEAR WILDLIFE POPULATION CYCLE SOLVED?

335

Ruffed Grouse is Trembling Aspen (Populus tremu-
loides Michaux) buds and aspen use is correlated with
Ruffed Grouse density (Jakubas and Gullion 1991).
These authors found that Ruffed Grouse consistently
selected Trembling Aspen buds with relatively low lev-
els of conifery! benzoate and that high levels of this
secondary metabolite were associated with declining or
low grouse density. Captive Ruffed Grouse assimilated
24% less energy from aspen buds with high levels of
coniferyl benzoate (Guglielmo ef al. 1996). Jakubas and
Gullion (1991) concluded that the impact of a shortage
of suitable aspen in winter “may increase predation
risks and energetic costs for Ruffed Grouse.” It is inter-
esting that, during four winters between 1985 and 1990,
the coniferyl benzoate levels Jakubas and Gullion re-
ported were highly correlated (77 = 0.84) with lunar dec-
lination.

Cloud conditions have more impact in terms of atten-
uating UVB radiation than stratospheric ozone, partic-
ularly at high latitudes (Svenoe ef a/. 1995). Pertsev and
Dalin (2010) separated lunar-phase and lunar-declina-
tion effects on cloudiness; both were statistically signif-
icant with lunar declination somewhat stronger. They
found a semi-monthly (13.66-day) variation in tropos-
pheric nighttime cloudiness, with the relative amount of
cloud increasing with an increase in lunar declination
by absolute value. If this relation is extendable (Ray
2007) from the fortnightly half-period of the nodical
month to the 9.3-year half-period of the nodal cycle, the
maximum declination line in Figure 2 may also repre-
sent relative cloudiness. If so, then Ruffed Grouse den-
sity is correlated with attenuation of UVB radiation,
which is generally thought to have a positive effect on
food quality (van Asch and Visser 2007). The relation
is particularly apparent at the Ruffed Grouse’s cyclic
lows, when exposure of food plants to UVB light is
highest. Ayres (1993) linked reduced radiation result-
ing from increased cloud cover to reduced (average of
54%) secondary metabolites in food plants and their
increased palatability to herbivores. In one study, Snow-
shoe Hares showed a strong preference for shaded Alas-
ka Paper Birch (Betula papyrifera ssp. humilis [Regel]
Hultén) shoots (Bryant et a/. 1987). The mechanistic
pathway suggested here is: increased lunar declination
leads to increased cloudiness resulting in reduced UVB
radiation and reduced secondary metabolites leading to
increased herbivore use and higher herbivore density.
This scenario is similar to the UVB plant stress hypoth-
esis proposed by Selas (2006) as a possible cause of the
Snowshoe Hare cycle, except that the driving force here
is the 9.3-year lunar nodal cycle rather than the 11.1-
year sunspot cycle and the intermediary is cloudiness
rather than ozone.

Snowshoe Hare population peaks tend to precede
maximum lunar declination by 1—2 years, and increas-
ing hare abundance is closely aligned with increasing
lunar declination (Figure 4). Like grouse, hare popu-
lation lows are associated with minimum lunar decli-

336

nation, suggesting that maximum tidal force is detri-
mental, directly or indirectly, to both species. Sinclair
et al. (1988) rejected a “fluctuating secondary com-
pound hypothesis” to explain the Snowshoe Hare cycle
because phenols declined through the increase and
decline phases and increased during the low phase —
exactly the opposite of what was predicted. Boonstra
et al. (1998) posited that the low phase of the hare cycle
might be related to deterioration in food quality be-
cause of the production of secondary plant defense com-
pounds but dismissed this idea because of the failure of
the fluctuating secondary compound hypothesis .

However, suppose that lower lunar declination (max-
imum tidal force) somehow raises the level of second-
ary metabolites in food plants above the threshold that
the hare must stay beneath in order to grow, reproduce,
and survive. In this case, the low phase of the hare cycle
would continue until the lunar declination increases
enough to exceed the threshold. Further study may clar-
ify this issue.

The relative timing of the 9.3-year moonlight cycle
and Snowshoe Hare density (Figure 4) suggests a pos-
sible mechanism for the Snowshoe Hare cycle, 1.e.,
nights without darkness at the equinoxes produce a
cyclic acceleration of the effects of key factors affecting
the hare cycle: predation rate, predation risk, predator-
induced stress, and nutritional deficiency. Predation is
generally considered to be the driving factor in the hare
cycle (Murray 2003). The rate of predation on Snow-
shoe Hares is higher when the moon is full (Griffin e¢
al. 2005). Griffin and Mills (2009) found that weekly
hare survival rates were about 2% lower in spring and
fall compared with summer and winter rates. Spring
and fall are the seasons when there is a possibility of a
pelage—background colour mismatch resulting in height-
ened vulnerability to predation (Litvaitis 1991). The in-
creased duration of moonlight near the peak of the 9.3-
year lunar cycle can be expected to increase the time of
elevated predation risk. Lima and Bednekoff (1999)
suggested that animals should exhibit the greatest anti-
predator behaviour in high-risk situations that are brief
and infrequent. Snowshoe Hares forage almost exclu-
sively in twilight and at night (Griffin et a/. 2005). Lack
of darkness after sunset could result in either delayed
feeding or foraging restricted to areas of protective cov-
er but lower quality food. In either case, the resulting
reduced food intake may lead to declining body condi-
tion and reduced reproductive output (Boonstra ef al.
1998). Rongstad and Tester (1971) found that the S—10-
minute daily return of Snowshoe Hares to care for their
young appeared to be related to light intensity in the
evening twilight period; this behaviour so crucial to
juvenile survival could be affected by nights without
darkness. Chronic predator-induced stress has been
posited to decrease reproductive fitness during the peak
and decline phases of the hare cycle (Boonstra et al.
1998; Krebs et al. 2001; Sheriff et a/. 2010). Nights

THE CANADIAN FIELD-NATURALIST

Vol. 128

without darkness may contribute to increased stress due
to conflict between predator avoidance and hunger.

The cyclic predation acceleration mechanism sug-
gested for Snowshoe Hares may also be applicable to
the Ruffed Grouse cycle (cf. Figures 4 and 5). In a large
study of radio-marked grouse in northwestern Wis-
consin, Lauten (1995) found that 96% of 341 known
mortalities were due to predation or hunting. Bumann
(2002) reported the highest predation rates in Septem-
ber, March, and April. Spring and fall are dangerous
seasons for Ruffed Grouse; behaviour such as drum-
ming, territorial defense, nesting, and dispersal are like-
ly to increase predation risk at the same time that rap-
tors are migrating. Rusch and Keith (1971) found that
predation of adult males was highest during the spring
and fall drumming periods. Archibald (1976) reported
that moonlight was the major external factor stimulat-
ing drumming and found that nearly continuous noctur-
nal drumming occurred around the full moon in April.
From fall to spring, however, Ruffed Grouse activity is
highly crepuscular. Archibald (1973) found that late
winter activity, which consisted mainly of travel to and
from and feeding in aspen clones, peaked about 30
minutes before sunrise and almost precisely at sunset.
Lack of darkness after sunset and before sunrise on
nights without darkness could disrupt the timing, loca-
tion, or duration of feeding resulting in nutritional defi-
ciency, increased predation risk, or perhaps stress.

The lunar nodal cycle could also affect cyclic pop-
ulation dynamics through a cyclic impact on phenol-
ogy. Temperature and photoperiod are the two most
important phenological factors (van Asch and Visser
2007), and the lunar nodal cycle may play a role in both
cases. First, cyclic nights without darkness at the eq-
uinoxes could confuse the perception of photoperiod.
Bunning and Moser (1969) suggested that moonlight
might interfere with photoperiodic time measurement
in some plants and animals. Bowden (1973) posited that
periods of continuous illumination at full moon could
provide photoperiodic cues for insects. Interruptions in
normal circadian light cycles and the resulting disrup-
tion of normal melatonin rhythms cause widespread
effects involving reproduction, physiology, and behay-
iour (Navara and Nelson 2007). If nights without dark-
ness at the equinoxes disrupt the circadian rhythms of
Snowshoe Hares, there are several possible effects
including timing of reproductive processes and molt.
Change in day length is most rapid near the equinoxes
and increases with latitude (e.g., ~5.5 minutes/day at
60°).

Second, there is growing evidence that climatic fac-
tors are influenced by the lunar nodal cycle. The sur-
face climate of much of western and central Canada is
strongly influenced by a large-scale atmospheric vari-
ation, the Pacific-North American (PNA) teleconnec-
tion (Mysterud ef a/. 2003). McKinnell and Crawford
(2007) reported that the PNA index accounted for over

2014

70% of the variation in average air temperature at
and near Sitka, Alaska, in January. In addition, they
found that the PNA index was significantly negative-
ly correlated with the lunar nodal cycle with a 2-year
lag. Yndestad (2006) found that the North Atlantic
Oscillation winter index was correlated with the lunar
nodal cycle. Meslow and Keith (1971) found that the
onset of breeding in Snowshoe Hares was significant-
ly correlated with the intensity of illumination (mea-
sured by cloud cover) during the previous midwinter.
Major seasonal molts (which occur near the equinoxes)
in the Snowshoe Hare are controlled by photoperiod
(Murray 2003), whereas snow onset and disappearance
are governed by temperature (Kielland et al. 2010).
Because the nodal cycle may have a cyclic influence on
both factors, there is an interesting possibility that the
degree of pelage—background colour asynchrony from
year to year may be cyclic as well. The lunar nodal
cycle could conceivably affect Ruffed Grouse phenol-
ogy through an effect on the timing of drumming, mat-
ing, or dispersal as well as the synchrony of hatching
and chick food availability.

Recent research suggests that lunar declination may
be an important factor in the ecology of northern lati-
tudes. Ramos da Silva and Avissar (2005) found that the
Arctic Oscillation has been “unambiguously correlated”
with the 9.3- and 18.6-year oscillations of the lunar
nodal cycle since the 1960s. Yasuda (2009) found a sta-
tistically significant 18.6-year periodicity, synchronized
with the nodal cycle, in tree-ring chronologies in west-
ern North America over a 300-year period, and attrib-
uted this bi-decadal variability to tidal mixing. Several
authors have reported 13.66-day (fortnightly) oscilla-
tions in Arctic tidal mixing, with maximum heat flux
occurring at minimum lunar declination (Rogachev ef
al. 2001; Martin et al. 2004). According to Ray (2007),
“the tidal mixing mechanism so plainly evident at near-
fortnightly periods should extend in principle to the
18.6-year period.” Li et al. (2011) found 13.6-day and
27.3-day oscillations in atmospheric pressure fields and
atmospheric geopotential height (which is what cli-
matic indices such as the PNA teleconnection and the
North Atlantic Oscillation measure). The lunar forcing
of these atmospheric tides reaches its maximum when
the lunar declination is zero. Krahenbuhl et al. (2011)
reported that the 27.3-day maximum lunar declination
cycle influences upper tropospheric circulation by de-
forming high latitude Rossby waves, which are asso-
ciated with the formation of weather systems. Foreman
et al. (2006) estimated that the 18.6-year nodal modu-
lation causes a 19% variation in the net incoming tidal
energy flux of the Bering Sea basin. The 18.6-year lunar
nodal cycle accounted for 59% of the variance in Pacif-
ic Halibut (Hippoglossus stenolepsis) recruitment from
1927 to 1983 (Parker et al. 1995). Based on wavelet
analysis, Yndestad (2009) concluded that the biomass
variability of zooplankton, capelin, cod, and herring in
the Barents Sea was related to both the 18.6-year ampli-

ARCHIBALD: THE ENIGMA OF THE LO0-YEAR WILDLIFE POPULATION CYCLE SOLVED?

337

tude and the 9.3-year phase tide. So it seems possible
that the climatic forcing that Yan e¢ a/. (2013) found
necessary for successful modeling of the Canada Lynx
cycle was actually the effect of the underlying lunar
nodal cycle.

There is some evidence that the 10-year wildlife
cycle has changed in recent years. Krebs ef a/. (2013)
reported that the widespread regional synchrony of hare
and lynx populations found in northwestern North
America from 1970 to 1990 broke down after the mid-
1990s. These authors found that Canada Lynx popula-
tions peaked in the early 1970s, 1980-81 and 1989-9],
but there was no clear sign of a strong cyclic lynx peak
after 1995. This study shows substantially greater dif-
ferences between predicted and mean reported peak
years after about 1965 (Table 1). Long-term monitor-
ing of 10-year-cycle species is required to increase our
understanding of cyclic phenomena (Krebs ef a/. 2013)
and to assess possible links between changes in pop-
ulation cycles and global warming (Yan ef al. (2013).

Just a coincidence?

Some ecologists will contend that the correlations
presented here are merely a coincidence. The coinci-
dence involves not only period but also phase: peaks
of the Snowshoe Hare and Ruffed Grouse cycles are
aligned with peaks of the nodal half-cycle. Period,
phase, and amplitude of the Ruffed Grouse cycle in
Minnesota and the “classic” Canada Lynx cycle in
northern Canada can be mimicked by the lunar models
presented. One school of ecological thought holds that
experiments are the only way to separate causation from
spurious correlation (Lambin ef a/. 2002). Unfortunate-
ly, the 9.3-year moonlight and lunar declination cycles
are not subject to experimental manipulation in the
field. Moreover, there does not appear to be any statis-
tical method available to prove that the 9.3-year nodal
half-cycle actually causes the timing of the 10-year
cycle (Lindstrom ef al. 1996; Berryman 2002). How-
ever, without invoking an exogenous zeitgeber, the per-
sistent period, regularity, and synchrony of the 10-year
cycle are difficult to explain convincingly. The 9.3-year
moonlight cycle and 9.3-year lunar declination cycle
are plausible causative factors. Previously, it was stated
that the ideal exogenous factor candidate would have a
consistent period that matched the wildlife cycle and
would shed light on its other unexplained features. The
93-year nodal half-cycle of the moon meets these cri-
teria. Unlike sunspots, the wildlife cycle has stayed in
phase with the nodal half-cycle for 150 years despite
major disruptive influences (¢.g., Krakatoa) that might
be expected to “affect the future course of the cycle
and thus become incorporated into the future history
of the series” (Moran 1953).

Acknowledgements

Thanks to Vidar Selas for advice and encourage-
ment, and to Marlene Klemm for interlibrary loans. I
am indebted to Duncan MacLulich, Charles Elton,

338

Lloyd Keith, the Minnesota DNR, Charles Krebs and
their many colleagues for supplying the long-term data
series that are the foundation of this paper. This work
would not have been possible without the free tools
available on the internet including the Astronomical
Sky Calendar, Google Scholar, JPL Horizons, US Naval
Observatory, and Wessa Statistics Software.

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Habitat Selection and Nest Success of the Upland Sandpiper
(Bartramia longicauda) in Ivvavik National Park, Yukon, Canada

aie | . 3
VALERIE MILLER!, Erica Not!:3, Linn P. NGUYEN2, and DEVIN M. TURNER!

ImMenartment of hi

Department of Biology, Trent University, Peterborough, Ontario K9J 7B8 Canada
“Parks C anada Agency, PO Box 1840, Inuvik, Northwest Territories XOE 0TO Canada
*Corresponding author: enol@trentu.ca

Miller, Valerie, Erica Nol, Linh P. Nguyen, and Devin M. Turner. 2014. Habitat selection and nest success of the Upland Sandpiper
(Bartramia longicauda) in Ivvavik National Park, Yukon, Canada. Canadian Field-Naturalist 128(4): 341-349.

The Upland Sandpiper (Bartramia longicauda) is a grassland shorebird species associated primarily with prairie habitats in
central North America. A disjunct and poorly studied population also occurs in Yukon, Canada, and Alaska, United States.
We studied habitat selection of nesting Upland Sandpipers in Ivvavik National Park, Yukon, at the scales of microhabitat (1-m
radius around nest) and putative home range (11.3-m radius plots at nests and within 50 m of nest). At the microhabitat scale,
the Upland Sandpiper selected nest sites with lower visibility from above than that of their home range (median 91.5%, range
70-98% versus median 99.0%, range 85—100%) and less-variable composition of vegetation than at random sites within the home
range. Vegetation adjacent to the nest in the eastern quadrat was significantly shorter (mean + standard error: 10.6 + 1.55 cm) than
that in other directions around the nest (> 13 cm); nest sites and microsites within home ranges were more often hummocky than
random sites in the park. At the mesohabitat scale, Upland Sandpipers selected sites within home ranges with fewer trees than
random sites within the park (10.3 + 3.0 trees per 11.3-m-radius plot around nest versus 32.9 + 5.9 trees per 11.3-m-radius plot
in the park) and greater herbaceous cover (70.7% + 3.0% versus 56.2% + 3.7%). Despite the disproportionate use of sites with
fewer trees, more herbaceous cover, and lower vertical visibility, these factors did not relate to nest success in our sample. Of the
24 nests found in 2010 and 2011, 22 contained four eggs and 2 contained three eggs. Upland Sandpipers at this high latitude site
had a nesting success rate of 0.85 + 0.01 and 0.56 + 0.01 in 2010 and 2011, respectively, for a 21-day incubation period. Further
assessment of the selected characteristics of nesting sites will improve our ability to predict the effects of northward shrub and
tree encroachment on this grassland species.

Key Words: Upland Sandpiper; Bartramia longicauda; mesohabitat; microhabitat; nest-site selection; northern limit; shorebirds;
Yukon; Ivvavik National Park

Introduction research from the northern disjunct range (Nouvet ef

Habitat selection refers to the preference of specific
environmental attributes that differ from other sites to
enhance fitness, survival, or both (Block and Brennan
1993: Jones 2001). Nest-site selection is a top-down,
hierarchical process that begins at the regional scale fol-
lowed by home range then specific nest site (Block and
Brennan 1993; Jones 2001). Vegetation characteristics
are typically important features of nest-site selection
for birds (Rodrigues 1994; Bollinger 1995; Clark and
Shutler 1999).

We studied vegetation characteristics that may influ-
ence nest-site selection by the Upland Sandpiper (Bar-
tramia longicauda), a medium-sized, grassland-nesting
shorebird. The largest part of the geographic range of
the Upland Sandpiper is in the Great Plains of North
America, although several small, disjunct populations
occur in northwestern Canada and Alaska (Houston ef
al. 2011). In the central and southern part of its breed-
ing range, the Upland Sandpiper prefers to nest in large
open grasslands with flowering herbaceous vegetation
and avoids woody and tall, dense vegetation (Kirsch
and Higgins 1976; Ailes 1980; Bowen and Kruse 1993;
Vickery et al. 1994; Dechant ef al. 2003*). Despite
some early publications on this species (Buss 1951;
Campbell 1967), there has been a near absence of recent

al. 2008), where warmer temperatures, earlier springs,
and longer growing seasons associated with climate
change have resulted in the advancement of the tree-
line (Szeicz and Macdonald 1995; Sturm ef a/. 2001;
Moen et al. 2004).

The objectives of our study were to describe the char-
acteristics of the habitat surrounding nest sites at the
microhabitat scale and within a 50-m radius, the pre-
sumed home range of the Upland Sandpiper (Mong
2005) and to compare habitat characteristics of success-
ful and unsuccessful nests at appropriate scales. Based
on nesting habitat characteristics described for south-
ern populations (Kirsch and Higgins 1976; Ailes 1980;
Bowen and Kruse 1993; Bollinger 1995; Dechant ef
al. 2003*; Vickery et al. 2010*), we predicted that the
Upland Sandpiper would select home ranges and nest
sites in open, grassland areas with little woody cover.
We also predicted that parents with nests closer to trees,
more trees in the home range, and less vertical nest cov-
er would experience less reproductive success than
those with nests farther from trees, fewer trees in the
home range, and more vertical nest cover. Our final ob-
jective was to document nest survival of the Upland
Sandpiper in the disjunct northern population for com-

34]

342

parison with other studies from the centre of its breed-
ing range.

Study Area

We collected data within a 4-km? area surrounding
Sheep Creek (69°09'N, 140°09'W) in Ivvavik Nation-
al Park, Yukon, Canada. Sheep Creek is located with-
in the British Mountains, which run perpendicular to
the Firth River (Brooks and Lane 2011*). Vegetation in
this area was primarily a function of elevation. At low
elevations, it was dominated by sedges (Carex spp.)
and stunted White Spruce (Picea glauca [Moench]
Voss). Dense stands of White Spruce occurred on steep,
mostly south-facing slopes. Low-growing vegetation,
e.g., Eight-petalled Mountain Avens (Dryas octopeta-
la [L.]) and Entire-leaved Mountain Avens (D. inte-
grifolia Vahl) and lichens and scattered shrubs, such as
willows (Salix spp.), Glandular Birch (Betula glan-
dulosa Michaux), and alder (A/nus spp.) occurred at
high elevations. Potential predators of nests and sand-
pipers in the study area included Gray Jay (Perisoreus
canadensis), Common Raven (Corvus corax), Merlin
(Falco columbarius), Northern Shrike (Lanius excu-
bitor), and Arctic Ground Squirrel (Cite//us parryi).

Methods
Nest location

In 2010, we searched for nests from 3 to 28 June.
Nests were found opportunistically, while searching for
songbird nests for a concurrent study (Turner 2013).
Nests were monitored every 2—7 days until 2 July when
the final egg hatched. Nests were checked, if there were
no predators detected in the area, by flushing the adult
bird from the nest. In 2011, the Upland Sandpiper was
a focal species for our study. We searched for nests from
I to 25 June in three ways: incidentally, while walking
through potential home ranges; by dragging a 5-m rope,
weighted down with tin cans, between two researchers;
or with two or more researchers walking 2—3 m apart.
Although we concentrated search efforts in open areas,
we also searched areas typically not used as nesting
sites by Upland Sandpipers, such as stands of stunted
White Spruce on the flat, shallow slopes, and heavily
forested areas. At discovery, we chose one egg from
each nest and used the float stages described by Lie-
bezeit et al. (2007), assuming a 21-day incubation
period, to estimate hatch date (Houston er a/. 2011).
We monitored nests every 3-4 days during incubation
and every 2 days after the estimated hatch date until
eggs showed signs of hatching (starring or pipping),
after which we checked nests daily until hatch. The last
egg hatched on 10 July 2011. In both years, nest loca-
tions were recorded using a Garmin GPSMAP 60CSx
global positioning system unit (Garmin International
Inc., Olathe, Kansas, USA) and marked with naturally
occurring objects placed within 10 m of nests to facili-
tate monitoring.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Nest success

We considered a nest to be successful if one or more
eggs was alive or had hatched (judged by observation
of adults with chicks or clean eggshell remnants in the
nest) by the last visit. We considered nests abandoned
when no adults were present in the area, or eggs were
cold on three successive visits. We examined nests
with missing eggs for evidence of predation (tracks
or broken eggshells). We estimated daily survival of
nests using the Mayfield estimate, assuming the nest
was lost at the midpoint between the last visit when it
was active and the first visit showing it had failed
(Mayfield 1975; Johnson 1979). We compared tree
density, distance to nearest tree, and vertical cover of
successful and failed nests using two sample (tests.
We used the delta method (Powell 2007) for calculat-
ing the variance in period survival rate. We did not use
logistic exposure methods or other models for our analy-
sis of nest survival or habitat preferences because of the
small number of nests in our sample. For all analyses,
we set a at 0.05.

Habitat analysis

Most vegetation variables were assessed differently
in 2010 and 2011 because of logistical constraints, but
assessments were made within two weeks of hatching
or nest failure in both years. In 2010, we measured veg-
etation characteristics at 10 nests and 10 random points
in the park. In 2011, we also measured vegetation char-
acteristics in 16 sampling plots within 50-m radius of
the nest (“home range’), as well as 16 “nest” and 23
“park” sites. We did not document space use or move-
ment, but selected a 50-m radius to include the pre-
sumed home range (Houston ef a/. 2011). Although
Upland Sandpipers are known to breed in loose col-
onies and exhibit little territoriality, other populations
have nests more than 300 m apart (Casey ef al. 2011,
Houston ef al. 2011). To select sampling sites within
the presumed home range, a random direction and dis-
tance up to 50 m from the nest were selected, exclud-
ing sites in water. To select park sites, we randomly
chose coordinates in the study area using ArcGIS
(ESRI, Redlands, California, USA) and located sites
in the field using a GPS unit. Randomly chosen sites
were excluded if they were in water, had a high tree
density (> 80 trees with a diameter at breast height
> 2.5 cm in an 11.3-m radius), or were more than 150 m
above the treeline in dry, sparsely vegetated areas. No
nests were found in these environments in either year.

In both years, we measured variables at two scales:
microhabitat (1-m-radius plots) and mesohabitat (11.3-
m-radius plots). In 2010, we measured height of the
tallest vegetation in each cardinal direction within 5 cm
of the point (using a ruler) and the elevation of the point
(using a GPS unit and comparing with topographic
maps for accuracy) at the microhabitat scale. We also
visually estimated the proportion of the nest visible
from | m above and the percentage cover of each spe-
cies and other elements (water and bare ground) in the

2014

microhabitat. Rare and short (< | cm) herbaceous
species were ignored. At the mesohabitat scale, we
paced the distance to the nearest tree (even if it was
beyond the plot boundaries) or shrub greater than | m
in height and recorded the plant species. The number
of trees and shrubs over | m tall within the 11.3-m-
radius plot was recorded, as well as the percentage cov-
er of shrubs less than | m tall. The area of the habitat
patch containing a nest or park site that did not contain

trees was delineated by walking the inner perimeter of

the surrounding trees using the track function on the
GPS unit.

In 2011 at the microhabitat scale, we measured veg-
etation height near the nest cup and the elevation as in
2010. In addition, we visually estimated percentage
vegetation cover, identifying each species (including
dead sedges and grass), bare ground, water, litter, and
lichens. The slope within the microhabitat was catego-
rized as < 5° or > 5°. Visibility from above was esti-
mated by placing a 13-cm-diameter disc (the approx-
imate diameter of most nests in our study) in the nest
cup or at the centre of the home range or park micro-
habitat sites and estimating the proportion that was vis-
ible. The disc was brown and speckled black to resem-
ble egg colouration. We recorded whether the site was
on a hummock or flat ground. We also recorded the
height of the tallest vegetation within the microhabitat
radius (to the nearest mm), the species, whether it was
dead or alive, and its direction and distance from the
nest.

At the mesohabitat scale, we recorded the number
and species of living and dead trees over | m tall, the
number and species of shrubs over or equal to | m tall,
and the percentage cover of shrubs less than | m. Trees
less than 30 cm in height with a single stem were des-
ignated as seedlings and counted. The distance to the
nearest tree or shrub over | m (even if outside the plot
boundaries) in each cardinal direction (using an esti-
mated 10° swath) was paced and the species was re-
corded, as well as the overall nearest tree and its direc-
tion. Percentage cover of the herbaceous layer, rock and
bare ground, lichens, and water were visually estimat-
ed. Slope within the mesohabitat was visually estimat-
ed in the same categories as above.

Statistical analysis

All data were tested for normality and homogeneity
of variances using Statistica v. 7.0 (StatSoft Inc., Tulsa,
Oklahoma, USA). Data were transformed to meet the
assumptions of parametric testing if necessary. If as-
sumptions of normality were not met after transforma-
tion, we used non-parametric tests. We used ANOVA
to determine whether vegetation height around the nest
varied as a function of direction. We then calculated the
average vegetation height adjacent to the nest for fur-
ther analyses. Data for nest and park plots from 2010 at
both microhabitat and mesohabitat scales were com-

pared using /-tests.

MILLER ET. AL: HABITAT SELECTION AND NEST SUCCESS OF THE UPLAND SANDPIPER

343

Data for nest, home range, and park plots from 2011
at both microhabitat and mesohabitat scales were com-
pared using ANOVA (for continuous data); equality of
medians was compared using y? and Kruskal-Wallis
tests (for nonparametric data) and y? tests (for cate-
gorical data). Variables that were significantly corre-
lated were removed from analysis. For example, height
of tallest vegetation adjacent to the nest was correlated
with the tallest vegetation within the microhabitat; thus,
only the latter was retained for analysis. Before analy-
sis, at the mesohabitat scale, we combined variables
related to the number and percentage cover of shrubs
and trees (e.g., numbers of trees over and under | m
were combined). As results were similar for the com-
bined and individual variables, we report results for
combined variables only.

To assess differences in the composition of herba-
ceous plants at the microhabitat scale in both years,
we used PERMANOVA, a permutational distribution-
free method for assessing similarities in communities
(Anderson 2005) and non-metric multidimensional
scaling (MDS) plots with fourth-root transformed data
using the Bray—Curtis similarity index. As vegetation
cover was assessed differently in 2010 and 2011, this
analysis was conducted separately by year. We exclud-
ed plant species that were present at fewer than three
sites, as rare species have little effect on the Bray—
Curtis distance, but increase stress. Stress represents
distortion in MDS plots; a value under 0.2 is consid-
ered a good visualization of the data (Clarke and War-
wick 2001; Clarke and Gorley 2006). We also com-
bined similar species, such as Louseworts (Pedicularis
spp.), to reduce the number of variables. We identified
15 species in 2010 and 39 species or genera in 2011.
PERMANOVA requires a balanced design, so we ran-
domly selected 16 of the park sites for comparison in
PAC

We first assessed the similarity of the dispersion
between the nest, home range, and park sites using
PERMDISP (Anderson 2005). Despite a significant
difference in the dispersion between the groups for
both 2010 and 2011 (see Results), we elected to com-
plete the PERMANOVA analysis to explore the data.
For both PERMDISP and PERMANOVA, we used
4999 permutations.

To represent relations among nest, park, and home
range sites graphically, we used PRIMER v. 6.13
(PRIMER-E Ltd., Plymouth, UK) to create MDS plots.
The axes of these plots do not relate to the original
variables, but the proximity of points within the plots
gives a relative measure of similarity (¢.g., closer points
are more similar, Proctor ef a/. 2012). A hierarchical
cluster analysis using group averages created a cluster
overlay on the plot. Resemblance levels of 20, 40, 60,
and 80% were used with a slackness value of 75%.

We used f-tests to compare successful and failed
nests (combining years to increase power) in terms of
three habitat characteristics (vertical visibility, distance

344

to the nearest tree and number of trees in the meso-
habitat).

Results
Microhabitat

Vegetation height adjacent to nests varied as a func-
tion of direction, with vegetation significantly lower
east of the nests than in other directions (/’, ,,= 3.13,
P=0.03); mean height of vegetation (+ standard error):
north, 13.2 + 1.5 cm; east, 10.6 + 1.6 cm; south, 13.4 +
1.2 cm; west, 15.4 + 1.5 cm. In 2010, neither average
tallest vegetation adjacent to nests nor vertical visibility
differed between nest and park sites (Table 1). Using
PERMANOVA, we found a marginally significant dif-
ference in vegetation composition between nest and
park sites (F’, ,.= 2.05, P = 0.08), and a significant dif-
ference in average within-group dissimilarities (F’, ,.=
8.64, P= 0.01). Vegetation cover around nest sites had
greater similarity (61.6%) than at park sites (45.7%).
Based on the MDS plot, stress was 0.16 and similarities
between the majority of the points were over 40%.

Among the nests found in 2011, vertical visibility
differed significantly between site types, with nest sites
less visible than home range or park sites (Table 2).
No other continuous microhabitat variables differed
significantly between the three site types. Over half of
all nests were on a gradual slope, but this proportion
was not significantly different than that of home range
and random park sites (Table 2). Nests were, on aver-
age, at an elevation of 254 m (similar to the value from
2010, Table 1), a value that also did not differ signifi-
cantly from home range or random sites (Table 2). Nest
and home range sites were significantly more likely to
be in hummocky habitats (> 80%) than random park
sites (47.8%).

In 2011, as in 2010, we found no significant differ-
ences in composition of the plant communities at nest,
home range and park sites (PERMANOVA, WN = 48,
F, ,,= 1.25, P= 0.24). The MDS plot shows that sim-

"247 ie cee
ilarity of plant composition between points is over 40%

THE CANADIAN FIELD-NATURALIST

Vol. 128

and nest sites are concentrated toward the upper centre
of the MDS plot (Figure 1). There were significant dif-
ferences in within-group vegetation composition for all
three site types (PERMDISP, F, ,,= 6.45, P = 0.002),
with nest sites more similar than both park and home
range sites, which were not different (Table 3).

Mesohabitat

Upland Sandpipers nested in Arctic and alpine tundra
and wet sedge meadows, although they were restricted
to drier locations in the latter. Nests occurred in both
small and large clearings within the scattered stunted
White Spruce. Most random park sites and nearly all
home range locations were on gradual slopes, reflect-
ing the hilly nature of the study area. Water in the park,
home range and nest mesohabitat plots was relatively
rare (< 25%; Table 2). The openings used for nesting
were small (< 0.1 ha), but these did not vary signifi-
cantly in size from openings in the random park sites
(Table 1).

Vegetation characteristics at this scale were similar
between nest and park sites in 2010 (Table 1). In 2011,
park sites had significantly more trees and less herba-
ceous cover than either home range or nest sites, which
did not differ from each other (Figure 2, Table 2). None
of the remaining variables differed across site types.

We found no significant differences between suc-
cessful and unsuccessful nests in vertical visibility
(t,, = 1.18, P= 0.25), distance to the nearest tree (t,, =
0.63, P = 0.53), or number of trees in the mesohabitat
(t,, = -0.92, P = 0.37). Inter-nest distances were 288 +
45 m in 2010 and 234 + 42 m in 2011.

Nest survival

In 2010, we found 10 nests, eight with four eggs, one
with three eggs and one with two chicks. In 2011, we
found 16 nests, 14 with four eggs, one with three eggs
and one after the chicks had hatched and left the nest.
In 2010, eight nests hatched successfully, one nest con-
tained whole broken shells on the last visit, and one nest
was abandoned, while in 2011, nine were successful,

TABLE 1. Comparison of habitat characteristics at micro and meso scales between Upland Sandpiper (Bartramia longicauda)

nest sites and random “park” sites at Sheep Creek, Ivvavik National Park, 2010.

—_——————— ——— —"—"“"—_————
Mean (+ SE) or median (range)

Habitat characteristic Nest (n = 10) Park (n = 10) 1e
Microhabitat

Elevation, m* 249.5 (239-293) n/a —_
Height of adjacent vegetation, cm RS ise 14) NOME 17 0.83

Vertical visibility, %* 91.5 (60.0—97.0) 82.5 (0.0—-100.0) 0.85
Mesohabitat

Area of clearing, m? 492.8 + 261.3 474.6 + 405.1 0.97
Distance to nearest tree, m 3.4+0.9 4.5+1.0 0.41
Shrub cover, % 4S 278 V2.4 2S 0.59
Number of trees over | m tall+ 17.1+8.6 24.3 + 5.0 0.48

ooo
N

ote: n/a = not applicable, SE = standard error.

*Results are presented as median (range) as the data do not conform to assumptions of normality.
+Log,, transformation used for analysis; means presented as untransformed data.

2014 MILLER ET. AL: HABITAT SELECTION AND NEST SUCCESS OF THE UPLAND SANDPIPER

Ww

4

Nn

= Ag?) % aric hahite ~haracterictice ¢ 1Cr, : j ; ]
TABLE 2, Comparison of habitat characteristics at micro and meso scales among Upland Sandpiper (Bartramia longicauda)
nests, random home range sites and random park sites at Sheep Creek, Ivvavik National Park, 2011.
aS 05—00200qa ee

Mean (+ SE), median (range), or proportion (%)

Home range

Habitat characteristic Nest (1 = 16) (n= 16) Park (n = 23) P
Microhabitat

Elevation, m* 254 (240-356) 260 (247-450) 254 (238-359) 0.30
Tallest vegetation, emt 30.4 + 1.8 44.6+6.3 54.7 + 13.1 0.11
Distance to tallest vegetation, cm 62.7 7.6 74.3 +5.9 79.2 +5.3 0.17
Vertical visibility, %* 91.5 (70.0-98.0) 99.0 (85.0-100.0) 99.0 (82.0—100.0) 0.0004
Proportion (%) on hummock 14/16 (87.5) 13/16 (81.3) 11/23 (47.8) 0.01
Proportion (%) on gradual slope 11/16 (68.8) 7/16 (43.8) 11/23 (47.8) 0.30
Mesohabitat

Herbaceous cover, % ROh 7 23.0 GIRSEE Sz Son2c= 3.7 0.045
Lichen cover, %§ 2D te Oe, 1.6+0.6 Dy iets (7 0.62
Bare ground, %+ 2.920656 6105 2.5 Srl = 176 0.81]
Shrub cover, % LS. eee 2.9) DS ee Aes 22,5 2:9 0.55
Number of shrubs over | m* 0.0 (0.0—2.0) 0.0 (0.0-35.0) 0.0 (0.0-18.0) 0.20
Number of saplings* 0.0 (0.0-4.0) 0.0 (0.0-4.0) 0.0 (0.0-38.0) 0.18
Proportion (%) on gradual slope 11/16 (68.8) 7/16 (43.8) LH/Z3KGS 9) 0.14
Proportion (%) with water 3/16 (18.8) 4/16 (25.0) 2/23 (8.7) 0.37
Number of trees$ NOE S10 9.4+3.3 32.93 5.9 0.004

Note: SE = standard error.

*Results are presented as median (range) as the data do not conform to assumptions of normality.
+Inverse transformation; means presented as untransformed data.

tSignificant differences between site types.

Site
& Home range
@ Nest
@ sC~Park

% Similarity

FicurE 1. Non-metric multidimensional scaling plot of vegetation in the microhabitat at Upland Sandpiper (Bartramia
longicauda) nests, random home range sites and random park sites at Sheep Creek, Ivvavik National Park, Yukon,
Canada, 2011. Using the Bray—Curtis similarity index, the cluster overlay has resemblance levels of 20, 40, 60, and
8( % and a slackness of 75%. Close points indicate sites with more similar plant composition. Stress was 0.18.

TABLE 3. Comparison of within-group similarity of vegetation at Upland Sandpiper (Bartramia longicauda) nests, random
home range sites and random park sites at Sheep Creek, Ivvavik National Park, 2011, using PERMDISP.

Bray—Curtis index (average

Site type within group similarity), % Comparator t P

Nest 66.3 Home range ay (0.009
Home range 56.8 Park 4.0 jee
Park 54.6 Nest 0.7 0.520

a re

346

125

Frequency

N
\
N
N
N
N
N
N
\
\
N
N
N
N
N
N
N
N
N
N
N
N
N

Oe i=20" “21-30

—

THE CANADIAN FIELD-NATURALIST

31-40

Vol. 128

@ Nest
NS Home range
O Park

>40

Number of trees/plot

FIGURE 2. Frequency distribution of number of trees in the mesohabitat (11.3-m-radius plots) at Upland Sandpiper (Bar-
tramia longicauda) nest and home range sites and random park sites at Sheep Creek, Ivvavik National Park, Yukon,

Canada, 2011.

seven were unsuccessful, and the fate of two nests
remained uncertain. Mayfield estimates of daily nest
survival were 0.989 + 0.007 (n = 10) in 2010 and
0.976 + 0.010 (n = 14, excluding the two uncertain
nests) in 2011 for period survival rates of 0.851 + 0.135
and 0.558 + 0.118, respectively, assuming a 21-day
incubation period.

Discussion

Of the many habitat variables we compared in this
study, only a few affected the choice of nest site by
Upland Sandpipers. As predicted, Upland Sandpipers
nested in home ranges with fewer trees and greater
herbaceous cover than found generally in this tree-
line region, although trees in the park were also rela-
tively sparse (32.4 trees/ha). Upland Sandpipers placed
their nests at sites with more vertical cover than avail-
able in their putative home ranges. The composition
of low-lying species of Arctic plants within the nest
sites was less variable than in the home range, suggest-
ing that a site with a relatively homogeneous plant com-
munity is sought. Most nests were placed in areas dom-
inated by sedges, with a mixture of Mountain Avens,
Lapland Rosebay (Rhododendron lapponcium [L]
Wahlenberg), mosses, and willows. None of the vari-
ables we studied predicted nest success of the Upland
Sandpiper, although our sample size of nests was small.

Vegetation east of the nests was 2—5 cm shorter than
in other directions. Upland Sandpipers probably entered
the nest cup consistently from the east, facing into the
predominant northwest wind, which would result in
some trampling of vegetation. Orienting into the wind
both reduces ruffling of the feathers (Gochfeld 1978)
and gives incubating bird greater cover from wind,
thus reducing conductive cooling.

Use of grasslands and other treeless habitats is a
characteristic of the Upland Sandpiper in the southern
and central part of its range (Kirsch and Higgins 1976;
Ailes 1980; Bowen and Kruse 1993; Dechant ef al.
2003*). In North Dakota, occupancy of mixed-prairie
grassland by Upland Sandpipers is negatively correlat-
ed with the proportion of woodland. Therefore, Upland
Sandpipers are designated a woodland sensitive species
(Grant et al. 2004). Trees may provide perches for
avian predators, as well as for their prey, increasing the
potential for higher abundance and activity of preda-
tors associated with woody vegetation. Simultaneous-
ly, trees reduce visibility and diminish the ability of
incubating adult Upland Sandpipers to sense these pred-
ators (Johnson and Temple 1990; Winter et al. 2000:
Coppedge et al. 2001; Chapman et al. 2004; Grant et
al. 2004; Graves et al. 2010). Although Upland Sand-
pipers avoided trees at the scale of their nests and home
ranges in the Yukon, they did use trees in the landscape
for perching and vocalizing while we were in the area
of their nests. Shrub cover in northern Yukon may not
have influenced Upland Sandpipers, as the majority
of the shrubs in this habitat were short (< 20 cm), not
much taller than the herbaceous layer, and scattered
in distribution. In addition, during the nesting season,
the shrubs had limited to no foliage, thus reducing the
effect on visibility.

Upland Sandpipers selected nesting sites with greater
vertical cover or less visibility than random sites with-
in the home range. Vertical or overhead cover, mostly
by grasses and sedges in our study, can increase nest
success by reducing visibility, especially for avian pred-
ators, and reduce egg loss from predation (Dwernychuk
and Boag 1972; Martin 1993; DeLong et al. 1995:
Stokes and Boersma 1998; Dion et al. 2000). The ben-

2014

eficial effect of overhead cover is especially important

during incubation recesses when the movement of

adults can make the nests more susceptible to avian
predators (Smith e a/. 2012). While incubating, Upland
Sandpipers have a highly cryptic plumage, and the ad-
ditional grasses and other plants that protruded around
the nest cup probably decreased their conspicuousness
even more. Overhead cover can also help to retain
warmth around the nests (With and Webb 1993). As
mean temperatures ranged from —4.2° to 26.9° C dur-
ing incubation in Ivvavik National Park (Government
of Canada 2014), features that reduce temperature stress
on the incubating Upland Sandpipers may improve
reproductive success.

Variation in plant species cover in the microhabitat
around nests was lower than at random sites in the
home range or park, although there was no significant
difference in the amount of species cover. The reduced
variation at nests suggests that Upland Sandpipers
are avoiding sites at the extremes. For example, the
maximum amount of bare ground at nesting sites was
9.0%, whereas at home ranges, it was 91.0% and at
park sites 28.5%. Upland Sandpipers did not use sites
that were wet, where Eguisetum spp. were common, or
dry, rocky sites, where Dryas spp. were common. Bare
ground could occur at either dry sites where rocks and
exposed ground were common or wet sites where the
water had receded. In the future, vegetation should be
categorized by habitat type (i.e., wet, dry, etc.) or struc-
ture, rather than by individual species, as it is unlikely
that Upland Sandpipers can distinguish between plant
species of similar height and shape.

Nest success (56-85%) in this northern breeding
population was at the high end of the range reported
from other more southerly locations (see review in
Garvey et al. 2013). This observation, albeit based on
a small sample size, is consistent with predictions re-
garding a latitudinal effect on predation risk (McKin-
non et al. 2010). Contrary to our predictions, nest suc-
cess was not influenced by either distance to trees or
number of trees at the mesohabitat scale, nor vertical
cover. This may have been due to the small sample size
of failed nests or because of a diverse predator com-
munity. When there is a diverse predator community
that uses various cues and searching methods to find
nests, it can result in no adaptive signal of habitat selec-
tion (Braden 1999, Dion ef al. 2000). Further explo-
ration of the relation between predator abundance and
an encroaching treeline is needed in this northern loca-
tion. In the prairies, Clay-colored (Spizella pallida) and
Vesper Sparrows (Pooecetes gramineus) have greater
nest survival near forest edges, as their main predator
(the Thirteen-lined Ground Squirrel, /ctidomys tride-
cemlineatus) is primarily found in grassland interiors
(Grant et al. 2006). By contrast, in predominantly for-
ested areas, many species experience lower reproduc-
tive success near edges (Burke and Nol 2000). The
nesting success of Upland Sandpipers in the Canadian

MILLER ET. AL: HABITAT SELECTION AND NEST SUCCESS OF THE UPLAND SANDPIPER

347

prairies appears to be predominantly influenced by nest
age and not by landscape features (Garvey ef a/. 2013),
a result that is similar to our findings.

We have provided the first detailed description of the
habitat of nesting Upland Sandpipers at the northern
edge of their range. As in the prairie regions, Upland
Sandpipers in the northern Yukon breed in home ranges
with less tree cover and greater herbaceous cover than
what is randomly found. Within lvvavik National Park,
Upland Sandpipers are restricted to inland, sparsely
treed regions (Brown ef al. 2007). In addition, within
the park, Upland Sandpipers nest in flat valleys and
along shallow slopes, as the mountains are too rocky
for this grassland species and the steep, south-facing
slopes have dense spruce cover. As early as the middle
of the last century, Buss (1951) suggested the potential
for habitat loss for Upland Sandpipers in the southern
Yukon because of encroachment of vegetation into the
clearings used for breeding. Given the already small
openings (< 0.1 ha) in the sparsely forested landscape
where Upland Sandpipers nest, habitat loss as a result
of northward shrub and tree encroachment with cli-
mate warming is a real threat to this population’s per-
sistence.

Acknowledgements

We are extremely grateful for funding provided by
Parks Canada, the Wildlife Management Advisory
Council (North Slope), the Natural Sciences and Engi-
neering Research Council and its Undergraduate Stu-
dent Research Award program, and the Northern Sci-
entific Training Program. We are especially grateful to
Kayla Arey, whose plant identification expertise was
invaluable. We thank the Parks Canada’s Western Arc-
tic Field Unit employees and volunteers for assistance
with logistics and fieldwork. We also thank Brett San-
dercock for detailed comments on an earlier version
of this manuscript.

Documents Cited (marked * in text)

Brooks, G. R., and L. S. Lane. 2011. A guide to the land-
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Dechant, J. A., M. F. Dinkins, D. H. Johnson, L. D. Igl,
C. M. Goldade, B. D. Parkin, and B. R. Euliss. 2003.
Effects of management practices on grassland birds: Up-
land Sandpiper. Northern Prairie Wildlife Research Center,
Jamestown, North Dakota, USA. Accessed 22 September
2014. http://www.npwrc.usgs.gov/resource/literatr/grasbird
/upsa/upsa.htm.

Vickery, P. D., D. E. Blanco, and B. Lépez-Lants. 2010.
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22 September 2014. http://www.whsrn.org/sites/default
/files/file/Upland_Sandpiper_Conservation_Plan_v1.1_
10 _02-28.pdf.

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Received 13 June 2013
Accepted 30 August 2013

Cory’s Bittern, xobrychus “neoxenus” exilis: Morph, Mutant,
or Mixed Bag?

PAUL SMITH

Fauna Paraguay, Encarnacion, Departamento Itaptia, Paraguay and Para La Tierra, Reserva Natural Laguna Blanca, Municipalidad
de Santa Barbara, Departamento San Pedro, Paraguay; email: faunaparaguay@gmail.com

Smith, Paul. 2014. Cory’s Bittern, Leobrychus “‘neoxenus”’ exilis: morph, mutant, or mixed bag? Canadian Field-Naturalist 128(4):
350-357.

Cory’s Bittern (Lxobrychus “neoxenus”’ exilis) is an enigmatic form, usually referred to as a rare dark colour phase or morph
of Least Bittern (/xobrychus exilis [Gmelin, 1789]). A review of the literature indicates that birds identified as Cory’s Bittern
show considerable plumage variation and commonly exhibit multiple plumage anomalies. Temporal and spatial clumping of
records shows that they are of irregular occurrence; thus, they do not meet the criteria for recognition as a colour morph.
Cory’s Bittern should thus be viewed as a colloquial name that refers to any one of a number of abnormally dark-plumaged,

genetically undefined and phenotypically heterogeneous Least Bitterns.

Key Words: Cory’s Bittern; /vobrychus neoxenus; Ardetta neoxena; Least Bittern; Lcobrychus exilis; leucism; melanism;

Ontario; Florida

Introduction

Since its description as Ardetta neoxena (Cory 1886a,
b), Cory’s Bittern has captured the attention of ornithol-
ogists on account of its extreme rarity, striking plumage
and disjunct distribution. Despite the confident asser-
tion that the form is “without doubt perfectly distinct
from any other known species” (Cory 1886a), doubts
over its specific identity were soon expressed. After
examining a number of specimens and publishing a
series of papers (Scott 1889, 1891), Scott (1892a) stated,
“I feel impelled to record the strong impression that I
entertain, that it will ultimately be found to be a color
phase of Botaurus exilis.” Sharpe (1894) expressed the
controversial opinion that “Ardetta neoxena of Cory
seems to be founded on very old individuals of A. exilis,
in which the rufous tips to the quills and the light stripes
on the back have disappeared with age or wear,” al-
though on examination of a specimen, he later afford-
ed it full species status (Sharpe 1898). Bangs (1915), on
the other hand, referred to Cory’s Bittern as a “case
of nothing more or less than erythrism.”

Bent (1926) concurred with the impression that
Cory’s Bittern is conspecific with Least Bittern (Jxo-
brychus exilis), but warned, “It should not be called a
color phase of a dichromatic species, as it occurs too
rarely and irregularly.” Hellmayr and Conover (1948)
indicated that “neoxenus” was synonymous with exilis
and followed Bent in describing it as a “melano-ery-
thristic mutation.”

In a thorough review of the state of Cory’s Bittern
knowledge, Pittaway and Burke (1996) speculated that
“Cory’s Least Bitterns seen today probably result from
a rare recessive allele in the population of typical birds”
and that “The lack of intermediate morphs between
typical and Cory’s Least Bittern suggests a single gene

having two alleles for colour: dominant (typical) and
recessive (Cory’s).” The authors lamented the lack of
available genetic material and highlighted the specu-
lative nature of this hypothesis. This theory has subse-
quently been adopted in the modern literature (Hancock
and Kushlan 1984; Gibbs et al. 1992; Sibley 2000),
perhaps stemming in part from a widespread but in-
correct understanding of the term “morph.”

Polymorphism is correctly defined as “the coexis-
tence in one interbreeding population of two or more
distinct and genetically determined forms, the least
abundant of which is present in numbers too great to be
due solely to recurrent mutation” (Ford 1945). Poly-
morphism can occur in two forms: transient (where an
advantageous gene spreads through the population)
and balanced (where it is maintained at a fixed level
by a balance of selective agencies). The definition “ex-
cludes continuous variation... and the appearance of
heterozygous mutants subject to elimination by selec-
tion” (Ford 1955). As this definition excludes both
plumage abnormalities (as suggested by Bent 1926)
and the segregation of rare recessives (suggested by Pitt-
away and Burke 1996), the status of Cory’s Bittern re-
quires clarification.

This paper is a critical review of the published data
that aims to re-examine the status of “neoxenus.” with
the hope that a pilot genetic study will one day be con-
ducted to confirm or dismiss the hypotheses proposed.

Spatial Distribution

Cory’s Bittern is known from very few documented
records, most of which are from eastern North America
before 1900 (Palmer 1976: Pittaway and Burke 1996:
Sibley 2011) (Figure 1). Despite the range of the Least
Bittern in the Americas — stretching from southern
Canada to northern Argentina (Hancock and Kushlan

350

2014

1984) — the 38 documented North American records
are of the migratory subspecies exilis and are concen-
trated around the Great Lakes and Florida, with 22
records from Ontario; 7 from Florida; 2 each from
Michigan, Ohio, and Illinois; and 1 each from Massa-
chusetts, New York, and Wisconsin (Brewster 1902:
Kumlien ef a/. 1948; Palmer 1976; Pittaway and Burke
1996; Sibley 2011). Additional undocumented sight
records are listed by Pittaway and Burke (1996) and
Sibley (2011) but, with the exception of a pair of recent
reports by the same observer (D. Arbour, of Red Slough,
Oklahoma — 15 August 2001 and 14 May 2012), they
do not expand the known distribution.

All three South American records are of the resident
subspecies erythromelas and are confined to a compar-
atively small area of southern Brazil (Sao Paulo State
and Mato Grosso do Sul [Teixeira and Alvarenga 1985;
Sibley 2011]) and adjacent Paraguay (Asuncion [Clay
et al. in press]). Though only one of the individuals was
collected (Teixeira and Alvarenga 1985), the other two
were documented by video (Sibley 2011) and photo-
graph (Clay ef al. in press) respectively. The South
American subspecies are non-migratory, but under-
take local movements in response to water conditions
(Martinez-Vilalta and Motis 1992).

Temporal Distribution

All Ontario specimens of Cory’s Bittern were tak-
en in the breeding season between May and Septem-
ber (Cross 1892; Brown and Brewster 1893; Fleming
1902: Pittaway and Burke 1996; Sibley 2011), and 21
of the 22 specimens from Ontario were taken in a 10-
year period between 1890 and 1900, with a single addi-
tional specimen taken in 1913. Pittaway and Burke
(1996) list undocumented sight records from Ontario
in “about 19207 1923, 1927; 1928, 1932; 1939, 1941,
1950, “in the 1950s,” 1973, and 1981. Cory’s Bittern
has thus not been documented in the “stronghold” of
its range since 1913 and not reported there since 1981.
Images of the specimens held in the Royal Ontario
Museum can be consulted online at www.jeaniron.ca
/2011/corys.htm.

Six of the first seven specimens of Cory’s Bittern
were taken in Florida, although the description of the
type specimen taken in Florida in 1885 lacks a precise
date (Cory 1886a): five specimens were taken between
9 July 1889 and 15 August 1891 (Scott 1889; Cory
1891; Chapman 1896). Although Scott (1892b) de-
scribes it as occurring “regularly though probably in
small numbers at Lake Flirt and Lake Okeechobee,”
the next report from Florida was not until Bent and
Copeland (1927) saw a bird on 7 April 1925. There
are no further Florida reports until an undocumented
sight record by J. Brunner in April 2003 (Sibley 201 1).
There have been no documented records of Cory’s Bit-
tern in Florida, the second core area of the range, for
almost a century. All records correspond approximate-
ly to the breeding season.

SMITH: STATUS OF CoRY’S BITTERN

The first Ohio specimen of Cory’s Bittern was col-
lected on 25 May 1907 in a “large colony of Least
Bitterns” (Ruthven 1907), and the second, possibly a
migrant bird, was photographed after flying into a
building at Youngstown on 3 October 1949 (Pittaway
and Burke 1996). In Illinois, Carpenter (1948) took a
specimen on | June 1909 and another was collected by
Eifrig (1915) on 23 May 1914. Cherrie (1896) report-
ed the only specimen from Wisconsin, a male in full
plumage on 22 May 1893, while Watkins (1895) re-
ported a “fine male” from Michigan on 8 August 1894,
which was followed by another Michigan male taken
on 14 May 1904 (Taverner 1905). Brewster (1902) doc-
uments the only Massachusetts record, collected in a
“yard” on 18 May 1901. Allen (1913) includes a pho-
tograph of a female taken at the Ithaca marshes, New
York on 17 May 1913, although Bull (1985) was later
unable to trace this specimen. Two modern sight rec-
ords from Oklahoma — 15 August 2001 and 14 May
2012 — were also during the breeding season, but far
removed from the traditionally understood “core range.”

South American records account for almost half of
the seven worldwide records since the 1950s (and all
of the documented records), but none of the records
prior to that date (Teixeira and Alvarenga 1985; Sibley
2011; Clay et al. in press). All South American records
have been taken in the austral autumn during April
and May. Consequently, all dated reports of Cory’s Bit-
tern are between April and October, with no reports
from anywhere in the entire range between November
and March. Wintering grounds, migration routes, and
whether these differ from those of “normal” birds are
all unknown.

Variation

Cory (1886a) described his Ardetta neoxena as “Top
of the head, back, and tail dark greenish black, showing
a green gloss when held in the light. Sides of the head
and throat rufous chestnut, the feathers on the back of
the neck showing greenish black tips. Breast and under-
parts nearly uniform rufous chestnut, shading into dull
black on the sides; wing-coverts dark rufous chestnut;
under wing-coverts paler chestnut. All the remiges
entirely slaty plumbeous. Under tail-coverts uniform
dull black.” In addition he noted the presence of two
white flank feathers, which he attributed to “albinism.”
Subsequent published descriptions of specimens as-
cribed to neoxena, however, show considerable diver-
gence from this type description; some examples are
summarized below.

Chapman (1896) noted great variation in the plu-
mage of Cory’s Bittern: “only three of my ten speci-
mens of neoxena are alike.” In comparing it to typical
exilis, he added, “there is no regularity in the substi-
tution of colors; hence these birds differ not alone in
color, but also in pattern of coloration.” Significantly,
he added, “Nor do the known cases of dichromatism
among Herons give us any ground for asserting that

Cyows THE CANADIAN FIELD-NATURALIST Vol. 128

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FiGURE |. Distribution of the Least Bittern, xobrychus exilis (grey shading; BirdLife International and NatureServe 2013),
and reports of Cory’s Bittern (/. “neoxenus” exilis) in North and South America documented in 1880-1900 (triangles),
1900-1920 (circles), 1940-1960 (squares) 1960-1970 (cross), and 2010-2013 (plus signs). 1. Lake Okeechobee area.
Florida, USA (Cory 1886a; Scott 1892a); 2. Toronto area, Ontario, Canada (Brown and Brewster 1893): 3. Lake
Koshkonong, Wisconsin, USA (Cherrie 1896); 4. Jackson County, Michigan, USA (Watkins 1895): 5. Scituate. Plymouth
County, Massachusetts, USA (Brewster 1902); 6. St Clair Flats, Michigan, USA (Taverner 1905): 7. Toledo, Ohio, USA
(Ruthven 1907); 8. Rob Roy Marsh, Aurora, Illinois (Carpenter 1948) and Fox River, 40 miles northwest of Chicago,
Illinois, USA (Eifrig 1915); 9. Ithaca Marshes, New York, USA (Allen 1913); 10. Youngstown, Ohio, USA (Pittaway and

Burke 1996); 11. Sao Paulo, Brazil (Teixeira and Alvarenga 1985); 12. Pantanal, Mato Grosso do Sul, Brazil (Sibley
2011); 13. Asuncion, Paraguay (Clay et al. in press). ‘

2014

neoxena is a dichromatic phase of exilis.”. However,
while admitting that he could not “explain this unusual
degree of variation,” he opted instead to ignore it and
consider Cory’s Bittern a distinct but presumably high-
ly variable species. Although seven of his ten speci-
mens showed “melanistic or albinistic markings or both
combined,” Chapman (1896) did not consider the pos-
sibility that the form represented a heterogeneous range
of phenotypic variations caused by diverse plumage
abnormalities.

A male specimen from Florida (no. 44 806) is de-
scribed as “completely melanistic” (Chapman 1896),
while Carpenter (1948) reported a Cory’s Bittern from
Illinois that was “extremely albinistic yet decidedly
melanistic,” in which the pale areas of the plumage
were entirely white, several irregular black patches
were present, and there were only scattered chestnut
feathers on the underparts. Allen’s (1913) specimen
from New York, however, showed no “albinistic ten-
dencies.” A sight record reported by Alex Lucas on 2
August 1939 along the Otonabee River at Rice Lake,
Peterborough County, Ontario, was described as ““choco-
late-colored all over” (Pittaway and Burke 1996).

The second known specimen of Cory’s Bittern re-
ported by Scott (1891) from Florida (no. 3237) was
described as “deep greenish-black” on the “entire upper
surface,” with this colour “also the general tone of the
belly, sides, and flanks, though a few whitish and some
chestnut feathers are mixed” and “bright chestnut” only
on the “lower surface of the neck and throat.” Further-
more, the greater, median, and mid lesser coverts are
“dark like the back at the bases but shade into deep
chestnut terminally.”

A male specimen (no. 11 449, collection of W. E.
D. Scott), also reported by Scott (1892a), is also dark
greenish-black “both above and beneath,” differing
yet again in that the “feathers of the lower neck alone
having traces (edgings) of dark chestnut, their centres
being of the greenish black tone prevailing through-
out the bird.” In the same paper (no. 11 451) is said
to be “very like Mr. Cory’s original bird save that there
are traces on the sides of the back of lines, much ob-
scured, but of a general brownish tint, thus approach-
ing Botaurus exilis.” Cherrie’s (1896) Wisconsin
specimen, a male in full breeding plumage, “agrees
minutely above (with the type specimen)” but below
“the throat and neck are just a trifle paler chestnut,
and there is just a little more white on the abdomen.”
Brewster’s (1902) Massachusetts specimen had the
“flanks, abdomen and under-tail coverts, slaty brown
tinged with reddish.”

In addition to other black markings on the head, the
second specimen from Michigan reported by Taverner
(1905) had a “triangular patch of black” below and
behind the eye that reached from the bill before fading
out on the ear coverts. Fleming’s (1902) adult male (no.
14, collection of the Provincial Museum of Toronto)
has chestnut sides of the head and the breast and ab-

SMITH: STATUS OF CoRY’S BITTERN

353

domen black “slightly tinged with chestnut” but pos-
sesses “a clearly-marked black line running from the
gape almost to the back of the head, on both sides,...
[that] nearly divides the chestnut on the sides of the
head.”

Among the three modern South American records,
similar variation is present. Teixeira and Alvarenga’s
(1985) bird did not show any greenish gloss on the
black areas of the plumage and had entirely black sec-
ondaries. The specimen from Paraguay does show the
greenish gloss on the black areas of the plumage, but
possesses the typical yellowish bill colouration of “nor-
mal” exi/is, not the “mostly blackish brown” associated
with Cory’s Bittern (Pittaway and Burke 1996), and has
a black medial line down the centre of the foreneck
(Clay ef al. in press).

Fleming (1902) reports a young male (no. 15, col-
lection of the Provincial Museum of Toronto) with
“peculiar” feet so that “the green of the tarsus and
toes is curiously mottled and blotched with reddish
brown,” leading to the speculation that “it may prove
that the feet and legs of the nestlings are not colored as
in the adult.”” However, the colour of the legs is not
mentioned in the description of the type and in the
subsequent literature is variably described as “black-
ish to brown” (Eifrig 1915), “dark(er) olive-green”
(Sibley 2011), and brownish without yellow tinges
(Teixeira and Alvarenga 1985), while the photograph
in Clay et al. (in press) shows legs that resemble those
of typically plumaged birds.

The morphometrics of Cory’s Bittern have received
scant attention. Chapman (1896) provided limited data
for a small number of specimens that he examined, but
elected not to discuss the results in his text. Presum-
ably he was unable to detect any obvious diagnostic
characteristics in his data.

Assortative Mating

In an attempt to explain the high rate of partial leu-
cism in specimens of Cory’s Bittern, Pittaway and
Burke (1996) offered assortative mating resulting in
inbreeding as a potential, if “unlikely” explanation.
Thus, it may be inferred that the other highly variable
plumage characteristics exhibited by the birds are to
be treated as normal “morph” plumage. With no more
likely explanation consistent with a morph hypothesis
offered to explain the strong link with leucism, it seems
relevant to address the evidence for assortative mating.

The tendency for Cory’s Bittern to pair with other
Cory’s Bitterns was first reported by Scott (1891) re-
counting a letter received by him from Mr. J. F. Menge
of Fort Meyers, Florida, regarding a nest discovered
on 8 June 1890. Although the letter mentions four
young birds in the nest, Mr. Menge did not provide a
description of them and alludes to the parent birds,
again without description. Never having seen these
birds, Scott relies solely on this secondhand account.
Fleming (1901) refers to a female on a nest, but again
there is no description of her and no mention of her

354

mate. The assumption clearly made by the author is
that Cory’s Bittern is a valid species and, hence, the
mate must be of the same species, an assumption sup-
ported by Ames (1901) who, describing the same bird
and eggs, refers to it as Ardetta neoxena. Bent and
Copeland (1927) recount another secondhand report
from Mr. Oscar E. Baynard, who found a nest in Flori-
da in 1927 and stated that “the young are always as
black as Clapper Rails and that both adults are always
dark-colored, evidence supporting the view that Cory’s
Least Bittern is a distinct species.” The certainty ex-
pressed is at odds with the conclusions that may be
reasonably drawn from an undocumented report of a
single nest. Perhaps the only actual evidence of asso-
ciation in these birds is of a male and a female col-
lected at Ashbridge’s Bay, Toronto on 12 July 1900,
although there is nothing to suggest that these birds
were breeding or had even formed a pair beyond the
collection date. In addition, a single pair of mated
Cory’s Bitterns would not be considered proof of
assortative mating.

Although there is no real evidence that Cory’s =
Cory’s pairs are the norm, there are also no published
reports of mixed pairs as occurs commonly in other
polymorphic herons (Hancock and Kushlan 1984). In
fact the data available are too limited and unreliable
to draw any conclusions at all.

Assortative mating as a consequence of mate choice
based on morph appearance is extremely rare in birds
and has been documented in the wild in very few avian
taxa. In some polymorphic geese, for example, most
birds select a mate with colour or pattern similar to
that of the family in which they were raised (Cooke
1978; Abraham et al. 1983). However, even in these
instances a considerable percentage of the pairs are
mixed (15-18% in the Snow Goose, Anser caerule-
scens), rendering secondhand statements that Cory’s
Bittern always mates with like highly suspect. Kalmus
and Maynard-Smith (1966) and Seiger (1967) go so
far as to state that sexual imprinting leading to absolute
assortative mating in a species with two morphs would
be a speciating mechanism.

Early authors convinced of the species status of
neoxena and the collectors who supplied them with
these valuable birds may be forgiven for assuming or
even promoting the idea of like with like pairs. Clear-
ly, early collectors were aware of the financial bene-
fits that species status brought, and Pittaway and Burke
(1996) mention at least one well-known Ontario col-
lector, George Pearce, who destroyed specimens of
normal exi/is to promote the idea of species status and
even dyed normal birds to sell to collectors. Perhaps
not coincidentally, the same collector is the author of
a supposed sight record of breeding birds at Lake Erie
in 1923 (Pittaway and Burke 1996).

Pittaway and Burke (1996) do not assume assorta-
tive mating (although the references they cite do) and
refer to assortative mating of any kind as “unlikely.”

THE CANADIAN FIELD-NATURALIST

Vol. 128

However, adopting the more likely scenario leaves us
without an explanation as to why, if it is indeed a
morph, Cory’s Bittern shows such high levels of leu-
cism and such random plumage variation overall.

Discussion

Although many specimens attributed to Cory’s
Bittern have shown only superficial similarity to the
description of the type (Carpenter 1948), the extraor-
dinary degree of plumage variation exhibited by spec-
imens of Cory’s Bittern seems to have been largely
overlooked by modern authors.

Cory’s Bittern specimens referred to as “partially
albinistic” are more properly described as “partially
leucistic” (van Grouw 2006). The enzyme tyrosinase,
necessary for the chemical processes that produce
melanins in vertebrates, is present in leucistic individ-
uals but absent in albinistic ones; hence, partial albinism
cannot exist (van Grouw 2006). Partial leucism is not
an indicator of genetic health, as it results from an
inherited disturbance disorder of pigment transfer dur-
ing which deposition of melanin in the feather cells
fails to occur (van Grouw 2006). According to Ket-
tlewell (1973), “Melanic, melanistic and melanochroic
forms refer to heterogeneous and genetically quite
indeterminate groups.” Furthermore multiple types of
plumage aberrations in a single individual are often
indicative of a genetic defect influencing several pig-
mentation systems (Buckley 1982; Davis and Blumin
2012). The irregular and varied plumage types seen
in Cory’s Bittern are thus explicable in the context of
plumage aberrations resulting from genetic defects and
are not consistent with the “distinct and genetically
determined forms” required for recognition of poly-
morphism (Ford 1945, 1955). The variation observed
is sO great, in fact, that it is not even possible to at-
tribute it to a single plumage aberration. Specimens
ascribed to neoxenus seem to exhibit a random assort-
ment of eumelanistic, leucistic, and phaeomelanistic
aberrations, and potentially others, each of which may
be influenced by any number of genes and pathways.

The localized spatial and temporal distributions of
the birds led Pittaway and Burke (1996) to suggest that
these represent places “where random processes al-
lowed the Cory’s morph to become temporarily estab-
lished, because of chance colonization by a few indi-
viduals with the trait.” However this begs the question:
where did these colonizing individuals come from?
The alternative and mutually exclusive explanation
offered by the same authors is that Cory’s is “an older
form that is now at a selective disadvantage and has
been replaced by the typical morph” (Pittaway and
Burke 1996). However, this ignores the fact that the
records clump temporally as well as spatially. In the
context of a morph hypothesis, the first statement
approximates transient polymorphism, but selective
advantage is not consistent with an association with
plumage abnormalities, nor is the latter scenario con-

2014

sistent with the summary statement “if our ideas about
the genetics are correct, it is likely that Cory’s Least
Bitterns will turn up from time to time” (Pittaway and
Burke 1996).

Aware of the potential for confusion between muta-
tion and true polymorphism, Ford (1945) states that
“any given gene subject to adverse selection must al-
ways be infrequent in the population; because being
constantly eliminated, it is dependent for its existence
upon mutation.” Therefore, contrary to Pittaway and
Burke’s (1996) speculation, balanced polymorphism
does not allow for temporary establishment of morphs
but, in fact, demands permanence “in marked contrast
to the distribution of rare genes maintained, ultimately,
by mutation pressure” (Ford 1945).

Ford (1945) notes, “A balanced polymorphism in
which the variation involved is environmental is gen-
erally the product of rather exceptional conditions.”
There is no indication that there is anything exceptional
about the environmental conditions in any of the three
main foci of records (Ontario, Florida, southeast South
America) when compared with the rest of the range
(Scott 1892b), nor do these areas have anything obvi-
ously in common that would favour the joint expres-
sion of this morph in such widely dispersed geographic
locations. It should also be noted that the spatial pat-
terns of occurrence observed in Cory’s Bittern do not
conform to distributional patterns exhibited by any oth-
er polymorphic Ardeid (Bent 1926).

A possible explanation for the observed spatial dis-
tribution that appears never to have been proposed is
that the concentrations of specimens may be, at least in
part, an artifact of observer effort. Ashbridge’s Marsh,
Toronto (now defunct), where most specimens were
collected (Fleming 1906), was frequently used for
shooting “so that the bird, though of retiring habits,
could scarcely have chosen a more frequented piece of
marsh” (Hubert Brown 1894 quoted in Pittaway and
Burke 1996, page 38). With all the specimens in this
area collected between 1890 and 1900, plus an addi-
tional one in 1913, and a similar pattern of temporal
clumping seen in Florida, it seems pertinent to repeat
the question first posed by Carpenter (1948): how did
the morph evade detection prior to these dates? And
given the high value of such specimens, why did their
collection stop so abruptly? Thus, while the spatial
distribution may conceivably be explained as an arti-
fact of observer effort, it seems that temporal distri-
bution cannot.

A recessive allele theory may be partly consistent
with the available data on the rarity of Cory’s Bittern,
but it fails to explain temporal patterns, plumage vari-
ation, and the association with partial leucism. Although
a single rare recessive allele might be expected to pro-
duce a single rare phenotype, rare but heterogeneous
mutations would result in a variety of different but rare
phenotypes that would be temporally and perhaps spa-

SMITH: STATUS OF CoRY’S BITTERN 35

wn

tially clumped. Observed patterns are thus consistent
with recurrent mutations.

More important, regardless of whether mutation or
recessive alleles is accepted as an explanation, in nei-
ther case are the resultant plumages correctly termed
morphs. Consequently the basic premise of Pittaway
and Burke’s (1996) rare recessive allele theory, even
if correct, falls outside the definition of polymorphism
(Ford 1955), which excludes “segregation of rare reces-
sives.

Conclusion

Cory’s Bittern does not comply with Ford’s (1945)
definition of polymorphism: it has not been geneti-
cally determined, it shows extraordinary phenotypic
variation, and it is not present in numbers too great to
be due solely to recurrent mutation.

The rarity of these aberrations, coupled with a ten-
dency for observers to label any Lrobrychus exilis that
exhibits abnormally dark plumage characteristics with
the name Cory’s Bittern, is responsible for the enor-
mous difficulties faced by those who try to define the
plumage characteristics of the “morph.” The most
familiar phenotype, described by Sibley (2011), and
understood today to be the typical Cory’s Bittern, un-
duly generalizes the range of variation exhibited by
birds ascribed to Cory’s in the literature. The assump-
tion of a single plumage type representing Cory’s Bit-
tern is an unfortunate byproduct of a general (if uncon-
scious) acceptance by modern ornithologists that Cory’s
Bittern is a valid colour morph and, hence, must sub-
scribe to a general form.

Pittaway and Burke (1996) state, “The almost com-
plete disappearance of the Cory’s Least Bittern... (are)
important losses of genetic diversity and habitat to the
Least Bittern.”” However, if, as seems likely, diverse
and potentially unrelated factors, some of which may
even be deleterious, are responsible for the variation,
then no such loss of important genetic diversity is tak-
ing place. The appearance and disappearance of a
localized population over a short period of time may
be attributed to the lower fitness of such abnormally
plumaged individuals (Buckley 1982; Slagsvold er al.
1988: Ellegren ef al. 1997; Rutz et al. 2004). Their
loss is to be expected.

Consequently, I propose that, following a line of
thought that began with Bent (1926) and Carpenter
(1948), Cory’s Bittern should be viewed as a colloquial
name referring to any one of a number of abnormally
dark-plumaged and phenotypically heterogeneous Least
Bitterns and not as a valid colour morph.

Acknowledgements

I thank Matt Garvey, Andrew Vitz and Jon Green-
law, who provided information on old records in their
states. Rob Clay and Tony Gaston read through the
manuscript before submission and provided extreme-
ly useful comments. I am extremely grateful to Hugo

356

Cabral for providing the map for this paper. Two anony-
mous reviewers greatly improved the manuscript with
their thorough revision and comments.

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Received 5 October 2013
Accepted 9 February 2014

Use of a Marsh Dominated by the Introduced European Lake Sedge,
Carex acutiformis, by Highly Localized Native Butterflies

PAUL M. CATLING!:3 and BRENDA KosTIUK2

‘Environmental Health, Biodiversity, Saunders Building, Central Experimental Farm, Agriculture and Agri-Food Canada, Ottawa,
Ontario K1A 0C6 Canada

*170 Sanford Ave., Ottawa, Ontario K2C 0E9 Canada

’Corresponding author: catlingp@agr.gce.ca

Catling, Paul M., and Brenda Kostiuk. 2014. Use of a marsh dominated by the introduced European Lake Sedge, Carex acuti-
formis, by highly localized native butterflies. Canadian Field-Naturalist 128(4): 358-362.

To determine whether native butterflies had colonized a marsh in Ottawa that was entirely dominated by the invasive alien
European Lake Sedge (Carex acutiformis), we surveyed two adjacent stands of the sedge and surrounding habitats. Dion
Skipper (Euphyes dion), Mulberry Wing (Poanes massasoit), Broad-winged Skipper (P. viator), and browns (Lethe spp-)
were all abundant in the introduced sedge, but absent from surrounding habitats. This is the first report of the use of invasive-
dominated wetland by native Canadian butterflies. Reduced nectar resources because of dominance of the invasive species
over native nectar-producing plants did not prevent significant colonization. The known restriction of the butterflies to native
Lakebank Sedge (Carex lacustris) as a larval host plant, but its absence in the area, coupled with dominance of its close relative,
European Lake Sedge, provides strong circumstantial evidence of the use of the latter as larval food. This report doubles the
number of recently localized native butterflies that have been able to increase their distribution by switching to habitat dominated
by invasive plants.

Key Words: Ottawa; invasive alien; European Lake Sedge; Carex acutiformis; larval food plant; Euphyes dion; Poanes massasoit:

Poanes viator, Lethe eurydice; localized butterflies; invasive management; food plant switch

Introduction

Invasive alien plants destroy habitat for butterflies
and other native insects by displacing specific food
plants. The invasion by Scotch Broom (Cytisus scopar-
ius [L.] Link) in southeastern Vancouver Island pro-
vides a good example (Baron and Backhouse 1999). On
the other hand, invasive plants have vastly increased
the abundance of some native insect species that were
rare and local only a few decades ago (Catling ef al.
1998). In all cases of beneficial effects of invasive alien
species, which are often dominant, the habitats have
been disturbed, early successional or both (e.g., road-
sides, old fields).

In the Stony Swamp Conservation Area west and
southwest of Bells Corners in the city of Ottawa, an
extensive area of sedge marsh is entirely dominated by
the invasive alien, European Lake Sedge (Carex acu-
tiformis Ehrh.) with scattered trees of Eastern White
Cedar (Thuja occidentalis L.), Red Maple (Acer rubrum
L.), and Ash (Fraxinus sp.). On 22 July 2013, we no-
ticed that the native North American Dion Skipper but-
terfly (Euphyes dion [W. H. Edwards, 1879]) was com-
mon and widespread in this marsh (Figure 1). Use of
invasive-dominated wetland by localized (sedentary)
native butterflies had not been reported previously. To
evaluate the extent of the use of this wetland by the skip-
per and other marsh butterflies, we completed a short
survey, the results of which are reported here.

Study Area
The main marsh, stand 1, is approximately 4.2 ha in
extent and is located at 45.3143, -75.8516 in the city of

Ottawa. Bounded by Robertson Road on the north and
the Trans Canada Trail on the south, it is situated on a
plateau of sandstone where water levels were apparent-
ly raised leading to the death or partial death of cedar
woods and the submergence of open mesic meadow fol-
lowed by colonization by the European Lake Sedge. The
adjacent stand 2, to the north across Robertson Road,
at 45.31440, -75.85337, has a treeless open part which
is 0.1 ha. It has apparently also undergone an increase
in water level and is equally dominated by European
Lake Sedge as is the understory of the surrounding ash
swamp. The dominance of the introduced sedge in these
two adjacent stands is remarkably complete. The plants
form a thick thatch of dead material along with dense
leaves that reach a height of 1-1.5 m and exclude most
light. Indeed over the entire area any other graminoid is
extremely rare and other herbs (mostly ferns) account
for less than 0.1% of the cover (Catling and Kostiuk
2003, Table 1). The marsh is sometimes dry for much
of the year, but filled with water to a depth of 20 em
in spring and early summer.

Methods

To determine the extent to which native butterflies
were localized in the area of the introduced sedge, we
surveyed the two stands as well as adjacent areas to the
north, south, east, and west. This survey was carried out
on 24 July 2013 between 10:30 a.m. and 2 p.m.; the
temperature was 18—22°C. Half an hour was spent at
each of six sampling sites and approximately equal
areas of 1.2 ha were sampled at each site. To the north
of the European Lake Sedge stands are areas of marsh

358

2014 CATLING AND KosTiUK: MARSH DOMINATED BY EUROPEAN LAKE SEDGE, BY BUTTERFLIES 359

so ntvanlie oh OY § ates

a aff ay - ny

FiGuRE |. Extensive marsh in Ottawa dominated by European Lake Sedge (Carex acutiformis). Inset: male Dion Skipper
(Euphyes dion) on leaf of European Lake Sedge. Photo: P. M. Catling.

TABLE |. Vascular plants, by frequency and percentage cover, recorded in 645-m? plots along transects through an open wetland
dominated by European Lake Sedge (Carex acutiformis), near Bells Corners, Ottawa. Cover is the percentage of leaf surface
in | m? of ground surface; thus, 200% means that | m? of ground contains 2 m? of leaf surface.

Frequency Cover
Species (%) (%)
European Lake Sedge, Carex acutiformis Ehrh. 100 200
Reed Canary Grass, Phalaris arundinacea L. 1.70 0.79
Marsh Fern, Thelypteris palustris Schott var. pubescens (Lawson) Fernald 1.24 0.12
Five-leaved Virginia Creeper, Parthenocissus quinquefolia (L.) Planchon ex DC. 1.08 0.10
Red-osier Dogwood, Cornus sericea L. 0.46 0.04
White Elm, U/mus americana L. 0.15 O77
White Birch, Betula papyrifera Marshall 0.15 0.77
Willowherb, Epilobium sp. 0.15 0.15
Crested Wood Fern, Dryopteris cristata (L.) A. Gray 0.15 0.01
Common Elderberry, Sambucus canadensis L. 0.15 0.10
Sensitive Fern, Onoclea sensibilis L. 0.03 0.03
Northern Lady Fern, Athyrium filix-femina (L.) Mertens var. angustum (Willdenow) G. Lawson 00.03 0.10

Nee ee eee SS, TEE eee
Source: Catling and Kostiuk 2003.

dominated (98%) by Broadleaf Cattail (7ypha latifolia straw (Galium album Mill.), Early Goldenrod (Solida-
L.), Reed Canary Grass (Phalaris arundinacea L.), and go juncea Aiton), Tufted Vetch (Vicia crac aL) and
Purple Loosestrife (Lythrum salicaria L.) and wood- Oregano (Origanum vulgare L.). The east side was
lands. To the south, on the south side of the Trans Cana- _ largely woodland with few open areas.

da Trail is marsh strongly dominated by Reed Canary The list of species found (Table 2) underestimates
Grass and woodland. To the west on higher ground — the numbers, because any individuals that could not
were dry, open meadows and semi-open woodlands be accurately identified were not listed. Close-focus
of Jack Pine (Pinus banksiana Lamb.), Sugar Maple _ binoculars were helpful, and some species were con-
(Acer saccharum Marsh.) and Eastern White Cedar. _ firmed after capture in nets. Care was taken not to
The meadows were dominated by Awnless Brome count an individual twice by noting direction of flight
(Bromus inermis Leyss.), Poverty Oatgrass (Danthonia and moving quickly along the census path.

spicata [L.] Beauv. Ex Roem. & Schult.), White Bed-

Vol. 128

360 THE CANADIAN FIELD-NATURALIST

TABLE 2. Butterflies recorded in a survey of six adjacent areas in a marsh in Ottawa on 24 July 2013. Numbers are underes-
timates, because individuals that could not be accurately identified were omitted and extra care was taken not to count an

individual twice by noting direction of flight and moving quickly along the census path.
ee eee ————

Species

Delaware Skipper, Anatrytone logan logan (W. H. Edwards 1863) 2 —
Common Wood Nymph, Cercyonis pegala nephele (W. Kirby 1837)

Clouded Sulphur, Colias philodice (Godart 1819)

Dion Skipper, Euphyes dion (W. H. Edwards 1879)

Dun Skipper, Euphyes vestris metacomet (T. Harris 1862)
Northern Pearly-eye, Lethe anthedon (A. Clark 1936)
Lethe eurydice (L.), Eyed Brown

Lethe spp.

Viceroy, Limenitis archippus archippus (Cramer 1775)
Pearl Crescent, Phycioides tharos tharos (Drury 1773)
Cabbage White, Pieris rapae (L.)

Mulberry Wing, Poanes massasoit (Scudder 1863)
Broad-winged Skipper, Poanes viator (W. H. Edwards 1865)
Eastern Comma, Polygonia comma (T. Harris 1842)

Hickory Hairstreak, Satyrium caryaevorum (McDunnough 1942)

Stand 1 Stand2 North South [East West

Nw |
|
|
S)

Great Spangled Fritillery, Speyeria cybele cybele (Fabricius 1775) 2 | | | 3 a

Results

The survey revealed 15 species of butterflies (Table
2). Recording of Lethe species was confused in the field
and the only voucher specimen retained was referable
to Eyed Brown (L. eurydice [L.]). We have listed the
Lethe species as one eurydice and 11 Lethe sp. We be-
lieve that the largest source of error occurred with the
Dion Skipper, and we suspect that the actual numbers
of this fast-flying species were up to twice those re-
corded because individuals were difficult to observe in
waving beds of sedge. The data for other species and
other habitats are thought to be reliable. Although the
surrounding habitats were not highly productive on the
day in question, there are some definite patterns of
restriction (Table 2). Mulberry Wing (Poanes massas-
oit [Scudder, 1863]), Broad-winged Skipper (Poanes
viator |W. H. Edwards, 1865]), browns (Lethe sp.) and
Dion Skipper were all abundant in the introduced sedge
but absent in surrounding habitats. Common Wood-

nymph (Cercyonis pegala nephele |W. Kirby, 1837])
and Dun Skipper (Euphyes vestris metacomet [T.
Harris, 1862]) were abundant in field habitats to the
east and west, but mostly absent from the marshes.

Discussion
Wetland: a new invasive habitat

Habitats dominated by invasive alien species that
are used by more or less sedentary eastern Canadian
butterflies (Table 3) include weedy, disturbed areas;
old fields; woody second growth: and roadsides. This
report is the first case of the use of an invasive-alien-
dominated wetland by native Canadian butterflies
that have a reputation for sedentary behaviour and
are widely reported to be highly localized.

A similar situation occurs in the coastal marshes in
New Jersey, where later instar larvae of the Rare
Skipper (Problema bulenta [Boisduval and Le Conte,
1837]) have been found on Common Reed (Phrag-

TABLE 3. Localized native eastern Canadian butterflies that have expanded locally or broadly by switching to habitat dominated
by an invasive alien plant that, in most cases, serves as larval food.

TTT 0 ee ————————______

Species

Introduced food plant and/or community dominant

Henry’s Elfin, Callophrys henrici (Grote and Robinson 1867)

Common Ringlet, Coenonympha tullia inornata
(W. H. Edwards 1861)

Eastern Tailed Blue, Cupido comyntas (Godart 1824)
Wild Indigo Duskywing, Erynnis baptisiae (W. Forbes 1936)

Dion Skipper, Euphyes dion (W. H. Edwards 1879)

Silvery Blue, Glaucopsyche lygdamus couperi (Grote 1873)
Browns, Lethe spp. including L. eurydice (L.)

Mulberry Wing, Poanes massasoit (Scudder 1863)
Broad-winged Skipper, Poanes viator (W. H. Edwards 1865)

Glossy Buckthorn Frangula alnus P. Mill. L.
(Catling et al. 1998)

Kentucky Bluegrass, Poa pratensis L.

(Eberlie and Hess 1980)

Clovers (Layberry et al. 2014)

Crownvetch, Securigera varia (L.) Lassen

(Layberry et al. 2014)

Lesser Pond Sedge, Carex acutiformis Ehth. (this article)
Alfalfa, vetches, clovers (Catling and Layberry 2013)
Lesser Pond Sedge (this article) ;

Lesser Pond Sedge (this article)

Lesser Pond Sedge (this article)

SS

2014 CATLING AND KostIUK: MARSH DOMINATED BY EUROPEAN LAKE SEDGE. BY BUTTERFLIES 36]

mites australis [Cav.] Trin. ex Steud.; Chazal and Hob-
son 2002), presumably the introduced subspecies aus-
tralis, which dominates some of these marshes (P.
Catling personal observation). The strong association
of the Broad-winged Skipper with wetlands dominated
by Common Reed in New England (Nakamura and
Cooper 2005; Stichter 2014) provides another exam-
ple since it is the European subspecies australis that
dominates these wetlands (personal observation).

A significant addition to recently localized occupants
of invasive habitat

Many butterflies native to Canada switched to intro-
duced plants as larval food before 1900 and have al-
ways been considered widespread. Examples includ-
ed the Black Swallowtail (Papilio polyxenes Fabricius,
1775) feeding on Queen Anne’s Lace (Daucus carota
L.), the Clouded Sulphur (Colias philodice Godart,
1819) feeding on Alfalfa (Medicago sativa L.), the
American Copper (Lycaena phlaeas {Linnaeus, 1761])
feeding on Sheep Sorrel (Rumex acetosella L.), and
the Silver Spotted Skipper (Epargyreus clarus [Cramer,
1775]) feeding on Black Locust (Robinia pseudoaca-
cia L.). Any of these may have been rare and local,
but their switch to introduced species occurred so early
in the settlement period that their original distributions
and food plants were not recorded and remain a mys-
tery.

In other cases, the switch to a new larval food has
been more recent, has involved previously localized
species, and has sometimes been accompanied by sub-
stantial range expansion. Examples include the in-
crease in Henry’s Elfin (Callophrys henrici (Grote and
Robinson) in Eastern Ontario accompanying the inva-
sion of Glossy Buckthorn (Frangula alnus P. Mill.,
Catling et al. 1998) and the similar effect of introduced
legumes on the Silvery Blue (Glaucopsyche lygdamus
couperi Grote, Catling and Layberry 2013; Layberry
et al. 2014). A switch in food source and spread has
been recent in some cases. For example the Wild Indi-
go Duskywing (Erynnis baptisiae (Forbes)) has only
recently adopted introduced Purple Crown-Vetch (Cor-
onilla varia) and areas dominated by it along highways
in eastern Canada (personal observation), although its
use of Purple Crown-Vetch in the United States dates
back to 1979 (Shapiro 1979). Other Hesperiidae, such
as the Hobomok Skipper (Poanes hobomok [T. Harris,
1862]), Long Dash (Polites mystic [W. H. Edwards,
1863]), Peck’s Skipper (P. peckius [W. Kirby, 1837]),
and Tawny-edged Skipper (Polites themistocles |La-
treille, 1824]), may have switched to introduced Ken-
tucky Bluegrass (Poa pratensis L.) as larval food and
colonized areas dominated by it long ago, but infor-
mation is lacking. Some species of Erynnis may also
have switched to invasive clovers, but information in
eastern Canada is incomplete. Regardless, the four new
species occupying wetlands strongly dominated by an

invasive plant almost doubles the number of local-
ized butterflies that have recently switched to inva-
sive habitats (Table 3).

Although races that have switched to non-native lar-
val food plants may be much less sedentary than those
confined to the native food plants from which they
developed, many are still more localized than wander-
ing. This requires more study, but clearly rare and local
species have increased and in some cases become wide-
spread and abundant by using an invasive plant habitat.

Potential use of European Lake Sedge as a larval

food plant

The butterfly species found to be confined to the
invasive-dominated marsh are often very localized and
do not wander far from the native broad-leaved sedges
that serve as their larval food plant (P. Catling person-
al observation; Layberry e¢ a/. 1998). As no native
sedges were available in the marsh surveyed, it seems
likely that European Lake Sedge was used by larvae
as a food plant. Dion Skipper, Broad-winged Skipper,
and Appalachian Brown are all reported to use the
native Lakebank Sedge (Carex lacustris Willd.) as lar-
val food (Layberry et a/. 1998); Lakebank Sedge is
closely related to the introduced European Lake Sedge
(Reznicek and Catling 2002). This is the first circum-
stantial evidence of the use of an introduced sedge as
larval food by native Canadian butterflies.

Limitations of domination by European Lake Sedge

In the marsh surveyed, nectar-producing herbs were
rare and evidently outcompeted by European Lake
Sedge. Swamp Milkweed (Asclepias incarnata L.) was
sparse over the entire area, but its flowers were much
visited by Dion Skippers, Mulberry Wings, and Broad-
winged Skippers. Nectar may be a limiting factor, but
the apparently high abundance of butterflies indicates
that this has not prevented colonization.

Conclusions

We report for the first time the use of wetlands dom-
inated by an invasive alien plant species by native Can-
adian butterflies with a reputation for sedentary behav-
iour. Our observations have almost doubled the number
of butterfly species that have been able to increase their
distribution by switching to habitat dominated by inva-
sive plants. There is circumstantial evidence of the
use of an introduced sedge species as larval food by
native Canadian butterflies. Limited nectar resources
resulting from invasive dominance over native nectar-
producing plants did not prevent significant coloniza-
tion by localized native butterflies.

A number of aspects require further study, including
the impact of limited nectar resources, precise deter-
mination of the Lethe species present in the marsh,
and proof of the use of European Lake Sedge as a lar-
val food plant.

362

Acknowledgements

Ross Layberry and Peter Hall provided helpful re-
views of this report. Don Lafontaine kindly checked
the identification of voucher specimens.

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Received 28 January 2014
Accepted 13 March 2014

Spiders of the Southern Taiga Biome of Labrador, Canada

R. C. Perry!3, J. R. PICKAVANCE?, and S. PARDY!

Wildlife Div isin, Department of Environment and Conservation, Government of Newfoundland and Labrador, 117 Brakes
om Cove, Corner Brook, Newfoundland and Labrador A2H 7S1 Canada

“The > Provincis vs C ave “2 ane S 2c 7 . ;

e Rooms Prov incial Museum, 9 Bonaventure Avenue, St. John’s, Newfoundland and Labrador AIC 3Y9 Canada
*Corresponding author: robperry@gov.nl.ca

Perry, R. C., J. R. Pickavance, and S. Pardy. 2014. Spiders of the southern Taiga biome of Labrador, Canada. Canadian Field-
Naturalist 128(4): 363-376. :
Ad hoc collections of spiders were made in August and early September 2003 and pitfall trap collections were conducted from
June to October in 2004 and 2005 in southern Labrador. These collections represent the first systematic spider sampling for the
most easterly area of mainland Canada. In total, 161 species in 15 families were identified to genus and species and 16 were inde-
terminate. Of the identified species, 89 were new records for Labrador and, of those, 16 species were new records for the province.
In total, 94 species (58.4%) have Nearctic distributions and 67 species (41.6%) are Holarctic. No Palearctic species were found.
Our study brings the number of spider species recorded in the province of Newfoundland and Labrador to 377 (213 in Labrador).

Key Words: Arachnids; Spiders; Eastern Canada; Labrador; species distribution

Introduction sampling occurred, is part of the taiga biome, which is

The distribution of the flora and fauna of Labrador _ typified by very cold winter temperatures, a lengthier
is largely undocumented, and this is particularly true growing season and more precipitation relative to the
for spiders. Provincial spider lists have been limited — tundra biome. In general, the soils in the taiga biome
to the Newfoundland portion of the province (Hackman are acidic and lack important nutrients such as nitrogen
1954: Pickavance and Dondale 2005), where Paquin et and phosphorus. It is dominated by coniferous trees,
al. (2010) reported 361 species; in Labrador, only 124 especially Balsam Fir (Abies balsamea [L.] Miller)
species have been identified. and Black Spruce (Picea mariana [Miller] Britton,

The completion of the Trans-Labrador Highway in — Sterns & Poggenburgh). Paper Birch (Betula papyrifera
2003 created an east-west corridor transecting southern Miller), Trembling Aspen (Populus tremuloides Mi-
Labrador and allowing access to vast tracts of previous- — chaux), and American Mountain-Ash (Sorbus ameri-
ly inaccessible old-growth forest (Figure |). Given the — cana Marshall) are the most common deciduous trees.
absence of biodiversity data for this territory, under the There are also large expanses of wetlands, especially
auspices of the Government of Newfoundland and Lab- bogs and fens, as well as numerous rivers, lakes, and
rador, we took advantage of this opportunity to collect ponds. Nested within the taiga biome are seven distinct
and identify spiders and insects in the area. Our work ecological regions (Meades 1990).We ensured each was
resulted in the first systematically collected data on the —_ yepresented when spider trap-sampling stations were
diversity of spiders in Labrador, important baseline data chosen (Table 1, Figure 1). Descriptions of each ecore-
to which the results of future studies may be compared. _ gion, based on Meades (1990) follow, along with brief

descriptions of sampling sites within each ecoregion.

Study Area

Labrador is the mainland portion of the Canadian
province of Newfoundland and Labrador. It is situated
in northeastern North America between 52° and 60°N
and 56° and 64°W and encompasses approximately
293 000 km2, about 3% of Canada’s total land mass
(Anderson 1985). The current Labrador spider fauna
likely colonized Labrador after the Wisconsin Glacial
Episode (the last retreat of the Laurentide glacier) ap-
proximately 24 000 years ago (Dyke et al. 2002). The
prevailing winds move from west to east and coloniza-
tion probably occurred primarily through wind disper-
sal (i.e., ballooning) from elsewhere in North America
as well as through introductions associated with the

arrival of Europeans on the Labrador coast. | ot
Labrador is contained within two distinct biomes: areas. Traps were placed in areas with clumps of lichens

tundra and taiga. Northern Labrador is found in the tun- — (C/adina spp.), laurels (Kalmia spp.), and Labrador Tea
dra biome, while southern Labrador, the area where our (Rhododendron groenlandicum. ).

Forteau Barrens

This ecoregion is located at the southeastern tip of
Labrador, near the Strait of Belle Isle (Figure 1). The
region is characterized by low hills covered by Black
Spruce, slope bogs, and barrens. The area is subject to
strong winds, frequent storms, cool, rainy summers,
and relatively mild winters. Annual precipitation is
about 1000-1250 mm and annual snowfall averages
3.5-4.5 m. July temperatures average 12°C and the
growing season ranges between 100 and 120 days. Wet
soils and climate limit the growth of trees. Barrens are
thickly covered with lichens.

For trap sampling in this ecoregion, we chose an al-
pine heath with dwarfed black spruce and some boggy

ee)

63

364 THE CANADIAN FIELD-NATURALIST Vol. 128

Survey Locations
Spider Collections
4 Ad-hoc Collections
@ Pitfall Trap Collections
Roads
es] Large Lakes and Reservoirs
Labrador
Newfoundiand
~ Quebec
Ecoregions where sampling occured
1 - Forteau Barrens
2 - Coastal Barrens (Okak-Battle Harbour)
3 - Mid-Boreal Forest (Paradise River)
BI 4 - String Bog (Eagle River Plateau)
WE 5 - Low Subarctic Forest (Mecatina River)
6 - High Boreal Forest (Lake Melville)
7 - Mid-Subarctic Forest (Lake Michikamau)

60°N

58°N

| Labrador Sea

56°N

Lake Melville

52°N

56°W

FiGuRE |: Ecoregions and sites where spiders were collected in southern Labrador in 2003 usi i
r 2 sing ad hoc mett
in 2004 and 2005 using pitfall traps (circles). : Se ee

2014 PERRY ET AL.: SPIDERS OF THE SOUTHERN TAIGA BIOME OF LABRADOR

TABLE |. Locations and ecoregions where spiders were collected using ad hoe methods (2003) and pitfall traps (2004 and

2005) in southern Labrador.

Location (site)
Red Bay

St.Lewis

Port Hope Simpson
Muskrat Falls (A)
Birch Stand (B)
Middle Brook
Ossak Camp (C)
Labrador West (D)
Ad hoe site |

Ad hoe site

Ecoregion
Forteau Barrens (1)
Coastal Barrens (2)
Mid-Boreal Forest (3)
High Boreal Forest (6)
High Boreal Forest (6)
Low Subarctic Forest (5)
Mid-Subarctic Forest (7)
Mid-Subaretic Forest (7)
Low Subarctic Forest (5)
Low Subarctic Forest (5)

Longitude, °N

365

Latitude, °W

2
Ad hoc site 3
Ad hoc site 4
Ad hoe site 5

String Bog (4)
String Bog (4)
Mid-Boreal Forest (3)

56.4069 51.9106
55.7057 52.3960
56.2660 52, 5185
60.7844 53.2606
60.9197 53.2294
63.1429 53.3785
65.0129 53.4233
65.2952 53.4125
60.4374 52.6097
60.2627 52.9495
59.7182 52.8255
58.8275 53.0876
57.6660 53.056

Coastal Barrens

This ecoregion is found on a narrow band of the
coast extending from Napaktok Bay south to the Strait
of Belle Isle and containing exposed headlands, some
sheltered inlets, and several islands (Figure 1). It has a
low subarctic climate with cool summers and a grow-
ing season of approximately 100-120 days. Annual
precipitation is 1000-1300 mm. Winters are very cold,
with an average snowfall of 3.0-4.0 m. The dominant
vegetation is Empetrum spp., and forest stands occur
only in valleys.

Our trap-sampling station was located at the southern
end of the ecoregion in a dwarf shrub barren dominat-
ed by lichens, some laurels, and Labrador Tea as well as
stunted Black Spruce. Traps were placed near boggy
sites, in areas with lichen, and at the bottom of rock out-
crops.

Mid-Boreal Forest

This ecoregion is located in southeastern Labrador,
near the Paradise River (Figure 1). It is characterized by
bedrock outcrops and productive, closed-crown forests
composed of Black Spruce and Balsam Fir. Hardwoods,
such as Paper Birch and Pin Cherry (Prunus pensyl-
vanica L. f.), can also be found, as well as raised bogs
in valleys. A boreal climate prevails with cool to warm
summers, short cold winters, a growing season of 120—
140 days, and annual precipitation between 1000 and
1300 mm (mean snowfall 4.0—5.0 m).

In this ecoregion both trap sampling and ad hoc
collection were carried out. Trap-sampling plots were
located near the town of Port Hope Simpson in mixed
stands of Balsam Fir and Black Spruce. Some hard-
woods were also present. Traps were set in moss.

String Bog

This ecoregion corresponds largely with the Eagle
River Plateau, which is 500-600 m above sea level and
consists of extensive string bogs containing numerous
open pools surrounded by fen vegetation. Summers are
cool and winters are very cold. Annual precipitation 1s
1000-1200 mm (mean snowfall approximately 5.0 m).

Vegetation in the area consists of scrub black spruce,
Labrador Tea, and Splendid Feather Moss (Hylocomi-
um splendens [Hedw.] Schimp. in B.S.G.). Sporadic
eskers support open lichen woodlands dominated by
Black Spruce. Speckled Alder (Alnus incana [L.]
Moench) can be found along most watercourses and
lakes. Only ad hoc sampling of spiders was done in this
ecoregion.

Low Subarctic Forest

This ecoregion, located primarily in southern Lab-
rador, is characterized by broad river valleys and rolling
hills covered by shallow till, drumlins, and eskers.
Summers are short and cool and winters are long and
very cold. The growing season is approximately 120—
140 days and annual precipitation is 1000-1300 mm
(annual snowfall 3.5—5.0 m). Open Black Spruce for-
ests are the dominant vegetation. String bog complexes
cover extensive areas throughout the region.

Ad hoc and trap sampling were done in this eco-
region. The trap-sampling site was an open Black
Spruce forest with a thick mat of lichens covering the
forest floor. Understory plants included Labrador Tea,
laurels, and other small shrubs. Traps were set in the
lichens and often did not penetrate into the soil be-
cause of the thickness of the lichens.

High Boreal Forest

This ecoregion encompasses the Churchill River
valley and the coastal plain surrounding Lake Melville
(Figure 1). Alluvial soils can be found in river terraces,
while the uplands have shallow, well-drained soils.
Summers are cool and winters very cold. The growing
season is 120-140 days, and annual precipitation is
800-1100 mm. Annual snowfall averages approximate-
ly 4.0 m. Forests in the area have closed canopies and
are highly productive. Richer slopes contain Balsam
Fir, Paper Birch, and Trembling Aspen. Black Spruce
is present in most stands and dominates upland areas
and lichen woodlands. Ribbed fens and plateau bogs
occur in upland depressions and coastal plains, respec-
tively.

366

In this ecoregion we chose two trap-sampling sites,
characterized by different types of vegetation. The first
was near the town of Goose Bay, on the branch road
to Muskrat Falls (Site A, Figure 1). The area consists
of large sand hills with intermittent Black Spruce and
large areas of Cladina spp. Traps were placed in open
sandy and lichen-dominated areas. The second site, also
located close to Goose Bay, was on a southeast facing
slope dominated by hardwoods, such as Paper Birch,
Red Maple (Acer rubrum L.), Pin Cherry and Trem-
bling Aspen (Site B, Figure |). The understory con-
tained clubmosses (Lycopodium spp.), broom mosses
(Dicranum spp.), ferns, small Red Alders (Alnus rubra
Bongard), American Mountain-Ash, and a thick layer
of leaf litter.

Mid-Subarctic Forest:

This ecoregion encompasses the upland plateaus of
central and western Labrador and is characterized by
eskers and drumlin ridges. The climate is continental
and subarctic with cool, short summers and long, cold
winters. The growing season is 100-120 days, annual
precipitation is 900-1110 mm, and annual snowfall
averages 4.0 m. White Spruce (Picea glauca [Moench]
Voss) dominates in the north, Black Spruce elsewhere
in this ecoregion. Trembling Aspen, open lichen wood-
lands, and, in areas with flat topography, string bog
complexes surrounded by Black Spruce— sphagnum
forests are also characteristic of the area.

In this ecoregion two sites with different types of
vegetation were chosen for trap sampling. The first,
near the Ossakmanuan Reservoir (Site C, Figure 1),
was dominated by closed-canopy Black Spruce/Kalmia
and Black Spruce/Cladina, with some leaf litter and
mosses also present. The second sampling site was in
a recently severely burned forest near Labrador City
(Site D, Figure 1). Most of the trees were fire killed,
and a thin layer of charred humus remained on the
ground. Some areas were beginning to be colonized by
Blueberry (Vaccinium L. spp.), laurels, and mosses.

Methods
Sampling

Ad hoe collections (random, non-systematic col-
lection of spiders by hand) were carried out between
7 August and 3 September 2003, before construction
of the last phase of the Trans-Labrador Highway, as
preliminary surveys at two sites in each of the String
Bog and Low Subarctic Forest ecoregions and one
site in the Mid-Boreal Forest ecoregion (Figure 1).
They were conducted by two survey teams of four
Newfoundland government conservation officers.
Each person was directed to look for spiders by
exploring shorelines, turning over rocks, and exam-
ining plants and debris during their spare time. When
found, spiders were placed in sample jars and pre-
served with ethyl alcohol.

Trap sampling was conducted in the summers of
2004 and 2005 between early June and early Octo-

THE CANADIAN FIELD-NATURALIST

Vol. 128

ber. In 2004, the Forteau Barrens, Coastal Barrens,
Mid-Boreal Forest and Mid-Sub Arctic Forest ecore-
gions were sampled. In 2005, collections were made
in the High Boreal Forest and Low Subarctic Forest
ecoregions (Figure 1).

Pitfall traps were placed at eight sites in the six eco-
regions (Table 1). Sampling sites were chosen based
on whether they contained vegetation typical for an
ecoregion. At each site, seven plots were established,
each containing 10 pitfall traps (for a total of 70 traps
per site and 560 for the entire study) placed in a circle
with a diameter of about 10 m. Samples were retrieved
from pitfall traps at each site from one to seven times
(average four), depending on weather conditions and
collector schedules, from June through early October.
Collecting was done by regional biologists from the
Wildlife Division, Department of Environment and Con-
servation, and conservation officers from the Depart-
ment of Natural Resources, Government of Newfound-
land and Labrador.

Each pitfall trap consisted of a 10-cm diameter
flower pot (450 mL volume) set in the ground. An ice
cream sundae cup (250 mL) was half filled with propy-
lene glycol and placed in the flower pot. This system
allowed for simple collection of contents and resetting
of traps. Each trap was covered with a white plastic
card held in place with four large nails (Spence and
Niemela 1994) to exclude rainwater. Samples from
each plot were placed in a single jar, labeled by plot
and site number.

Spider sorting and curation

Specimens and other materials were removed from
the collected samples using a sieve. Spiders were then
separated and placed, with data labels, in clean vials
containing 95% ethanol. The spiders were subsequent-
ly sent for identification to Memorial University of
Newfoundland and Labrador.

Checklist

Specimens that could not be identified are shown as
indeterminate in the checklist and housed for future
examination at either The Rooms Provincial Museum
(marked NF in the checklist) or at the Canadian Na-
tional Collection of Insects and Arachnids, Agriculture
and Agri-Food Canada, Ottawa (marked CNC ). If they
have been catalogued, a number is also listed.

Species nomenclature follows Platnick (2014). The
number of species identified in each family is given
in parentheses after each family heading. Collection
locality is expressed as numbers | through 7, corre-
sponding to the ecoregions (Figure 1). Ecoregions 6
and 7 each contained two sampling sites and, there-
fore, location is further divided into A and B or C and
D, respectively. Collection dates are presented as month
and day. The total number of each species is presented,
separated into male () and female (9) specimens.
Comments are primarily limited to general species dis-
tribution (Holarctic or Nearctic). In some instances,

2014

comments also include species abundance and state
whether the record is new for Labrador or the entire
province,

Results

In total, ad hoe and trap-sampling collections pro-
duced 14 964 spider specimens (including indetermi-
nates) representing 161 species in 15 families (see
checklist and Table 2). The five ad hoc collections
produced 136 spiders representing 29 species in 11
families (73 of these were immature and identified only
to family). Nineteen species were represented by only
a single specimen. From the pitfall trap samples, 14
901 specimens were collected, representing 147 species
in 15 families.

Checklist of Labrador Spiders
AGELENIDAE (1 SppP.)

Agelenopsis utahana (Chamberlin & Ivie, 1933)
Ecoregions: 6A

Collection date: Sept. 13

Total samples: 2 = 1, ¢=1

Comment: Nearctic; new record for Labrador

AMAUROBIIDAE (5 SPP.)

Arctobius agelenoides (Emerton, 1919)

Ecoregions: 5, 7C

Collection dates: July 18; Oct. 8

Total samples: 2 = 1, ¢=1

Comment: Nearctic; new record for both the province and
Eastern Canada

Callobius bennetti (Blackwall, 1846)

Ecoregion: 6B

Collection dates: June 17; July 4, 5; Aug. 3, 12, 25; Sept. 13
Total samples: 2 = 17, ¢ =41

Comment: Nearctic

PERRY ET AL.: SPIDERS OF THE SOUTHERN TAIGA BIOME OF LABRADOR

367

Callobius nomeus (Chamberlin, 1919)
Ecoregions: 5, 6A

Collection date: Aug. 25

Total samples: 2 = 3, ¢ = 1
Comment: Nearctic

Cyhaeopsis euopla (Bishop & Crosby, 1935)

Ecoregions: 1, 2, 3, 5, 6A and B, 7C

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 2 = 52, ¢ = 82

Comment: Nearctic; good representation across all sampling
stations

Cybaeopsis tibialis (Emerton, 1888)

Ecoregion: 6A and B

Collection dates: June 17; July 4; Aug. 3, 12; Oct. 27.
Total samples: 2 = 32, ¢ =2

Comment: Nearctic; majority (31) found at site B

ARANEIDAE (8 SPP. AND | INDETERMINATE)
Araneus nordmanni (Thorell, 1870)
Ecoregion: 4

Collection date: Aug. 3

Total samples: 9 = 1, d= 1

Comment: Holarctic

Araneus saevus (L. Koch, 1872)
Ecoregion: 6B

Collection date: Sept. 13

Total samples: 2 = 1, d =0
Comment: Holarctic

Araneus trifolium (Hentz, 1847)

Ecoregion: 2

Collection date: Aug. 7

Total samples: 9 = 1, d =0

Comment: Nearctic

Araneus sp. Clerck, 1757 (indeterminate; NF)
Ecoregion: 3

Collection date: Aug. 25

Total samples: 9 = 0, d = 1

TABLE 2: Orders and composition of the spider fauna sampled by ad hoc (2003) and pitfall trap (2004 and 2005) collection

in southern Labrador.

Holarctic
0

Family Nearctic

Agelenidae
Amaurobiidae
Araneidae
Clubionidae
Dictynidae
Gnaphosidae
Hahniidae
Linyphiidae
Liocranidae
Lycosidae
Philodromidae
Salticidae
Tetragnahidae
Theridiidae
Thomisidae
Total 2

NN DSO

Nn
WwW

WNW WNOSOWANNN NH —

COM ONA & =

lon

Introduced Total species % of total
0 | 0.6
0 5 Sl
0 8 5.0
0 4 pes)
0 4 2.5
0 16 9.9
0 4 PEs)
0 80 49.7
0 | 0.6
0 IS 9.3
() 4 2:5
0 3 1.9
0 2 1.2
0 4 D3
0 10 6.2
0 16] 100

368 THE CANADIAN FIELD-NATURALIST Vol. 128

Araniella displicata (Hentz, 1847)
Ecoregion: 3

Collection date: Aug. 25

Total samples: 9 = 1, d =0
Comment: Holarctic

Araniella proxima (Kulezynski, 1885)
Ecoregion: 6B

Collection date: Aug. 3

Total samples: 9 =0, d= 1

Comment: Holarctic; new record for Labrador

Cyclosa conica (Pallas, 1772)
Ecoregion: 4

Collection date: ees 7

Total samples: 2 = 1, ¢=0
Comment: Faleretie

Hypsosinga rubens (Hentz, 1847)

Ecoregion: 6A

Collection date: July 5

Total samples: 9 =0, d= 1

Comment: Nearctic; new record for Labrador

Larinioides patagiatus (Clerck, 1757)
Ecoregion: 4

Collection dates: Aug. 7; Sept. 3

Total samples: 9 =2, ¢=1

Comment: Holarctic

CLUBIONIDAE (4 spp.)

Clubiona bryantae Gertsch, 1941

Ecoregion: |

Collection dates: Aug. 6, 23; Sept. 7; Oct. 26
Total samples: 9 =9, d =6

Comment: Neatete new record for Labrador

Clubiona canadensis Emerton, 1890

Ecoregions: 1, 2, 3, 6B, 7C and D

Collection dates: June 17; July 4, 11, 22; Aug. 5, 6, 10, 25
Total samples: 9 = 7, 6 = 16

Comment: Nearctic

Clubiona kulczynskii Lessert, 1905

Ecoregions: 1, 2, 5, 6B, 7D

Collection dates: July 4, 5, 11, 22; Aug. 3, 6, 25
Total samples: 9 =6, 6 =5

Comment: Holarctic

Clubiona trivialis C. L. Koch, 1843

Ecoregions: 1, 2, 6A, 7D

Collection dates: June 17; Aug. 1, 6, 25; Sept. 7; Oct. 26
Total samples: 2 = 5, ¢ =3

Comment: Holarctic

DICTYNIDAE (4 spp.)
Dictyna brevitarsa Emerton, 1915

Ecoregion: 2
Collection date: Aug. 7
Total samples: 9 =0, 6 =1

Comment: ee new record for Labrador

Emblyna annulipes (Blackwall, 1846)
Ecoregion: 2

Collection date: Aug. 7

Total samples: 9 = 1, 6 =0
Comment: Biotarete

Emblyna manitoba (Ivie, 1947)

Ecoregions: 4, 5

Collection date: Aug. 7

Total samples: 9 = 1, ¢ =1

Comment: Holarctic; new record for the province

Hackmania prominula (Tullgren, 1948)

Ecoregions: 6A, 7C

Collection dates: July 5, 19; Aug. 13, 29

Total samples: 2 = 0, 4 = 18

Comment: Nestea: new to Eastern Canada and the province

GNAPHOSIDAE (16 spp.)

Drassodes mirus Platnick and Shadab, 1976
Ecoregion: 2

Collection dates: July 22; Aug. 10; Oct. 27

Total samples: 9 = 1, ¢ =3

Comment: Nearctic; new record for the province

Drassodes neglectus (Keyserling, 1887)
Ecoregions: 2, 6A

Collection dates: June 17; July 5, 11, 22

Total samples: 9 = 1, d =4

Comment: Holarctic; new record for Labrador

Gnaphosa borea Kulezynski, 1908

Ecoregions: 1, 2, 3, 4,5, 6A, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 9 = 109, ¢ = 365

Comment: Holarctic; the large majority were found in
ecoregion 3 in July

Gnaphosa brumalis Thorell, 1875

Ecoregions: 2, 7C and D

Collection dates: bes lil, 225 Aug. 1, 29

Total samples: 9 =7, ¢ =31

Comment: ee the majority came from ecoregion 7,
site D

Gnaphosa microps Holm, 1939

Ecoregions: 1, 2, 3, 5, 6A and B, 7C and D

Collection dates: June; July; Aug.; Sept.: Oct.

Total samples: 9 = 49, 4 = 100

Comment: Holarctic; the majority were found in ecoregion
7, sites C and D

Gnaphosa muscorum (L. Koch, 1866)

Ecoregions: 1, 2, 3, 4, 5, 6A and B, 7D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 2 = 70, ¢ = 215

Comment: Holarctic: the majority were found in ecoregion
6, site A and ecoregion 7, site D

Gnaphosa parvula Banks, 1896

Ecoregion: |

Collection dates: oe 6, 7, 235 Sept: 7

Total samples: 9 = 1, ¢=5

Comment: NEKOHC: new record for Labrador

Haplodrassus eunis Chamberlin, 1922

Ecoregions: 5, 6A

Collection dates: June a July 5, 19; Aug. 3, 25

Total samples: 2 = 24, ¢ =70

Comment: feet a one specimen from ecoregion 5;
new record for the province

Haplodrassus hiemalis (Emerton, 1909)

Ecoregion: 2

Collection coke July 22; Aug. 25; Sept. 7; Oct. 4, 27
Total samples: 2 = 0, 4 =5

Comment: aac new record for Labrador

2014

Haplodrassus signifer (C. L. Koch, 1839)

Ecoregions: 1, 2, 3,5, 6A, 7C and D

Collection dates: June 17; July 4, 5, 11, 18, 20, 22; Aug 1, 2,
29; Oct. 8 :
Total samples: 2 = 49, 4 = 66

Comment: Holarctic; the majority came from ecoregion 7,
site D; new record for Labrador

Micaria aenea Thorell, 1871

Ecoregions: 3, 5, 6A and B

Collection dates: June 17; July 4, 5, 19, 22; Sept. 13

Total samples: 2 = 46, 4 = 30

Comment: Holarctic. The majority (70) were found in
ecoregion 6, site A; new record for Labrador

Micaria constricta Emerton, 1894
Ecoregion: 7D

Collection dates: July 11; Aug. 1
Total samples: 2 = 0, ¢ =3
Comment: Holarctic

Micaria pulicaria (Sundevall, 1831)
Ecoregions: 1, 2, 3,5, 6A, 7D

Collection dates: June; July; Aug.; Sept.

Total samples: 2 = 23, ¢ = 19

Comment: Holarctic; new record for Labrador

Orodrassus canadensis Platnick & Shadab, 1975
Ecoregion: 5

Collection date: Aug. 7

Total samples: 2 = 0, ¢ = 1

Comment: Nearctic

Zelotes fratris Chamberlin, 1920

Ecoregions: 2, 3,5, 6A

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 2 = 25, ¢ = 50

Comment: Holarctic; the majority were found in ecoregions
5 and 6

Zelotes sula Lowrie and Gertsch, 1955
Ecoregions: 6A, 7C

Collection dates: July 19; Aug. 3, 25, 29
Total samples: 2 = 0, ¢ =4

Comment: Holarctic

HAHNIIDAE (4 SPP.)

Cryphoeca montana Emerton, 1909

Ecoregions: 5, 6B

Collection dates: June 17; July 4, 5, 19, 22; Aug. |
Total samples: 2 = 5, ¢ = 55

Comment: Nearctic

Hahnia cinerea Emerton, 1890

Ecoregion: 6A

Collection date: June 17

Total samples: 9 = 0, 3 =!

Comment: Nearctic; new record for Labrador
Hahnia glacialis Sorenson, 1898

Ecoregions: 1, 2, 3, 7C and D

Collection dates: July; Aug.; Sept.; Oct.

Total samples: 2 = 78, ¢ = 124

Comment: Holarctic; specimens were plentiful at all of the
listed sites

Neoantistea magna (Keyserling, 1887)
Ecoregions: 2, 3, 5, 6B, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 383, a= sl

Comment: Nearctic; specimens were plentiful at all of the
listed sites

PERRY ET AL.: SPIDERS OF THE SOUTHERN TAIGA BIOME OF LABRADOR

369

LINYPHIDAE (80 spp. AND 15 INDETERMINATE)

Agyneta allosubtilis Loksa, 1965

Ecoregions: 1, 3, 6A

Collection dates: June 17; July 5, 11, 19, 22, Aug. 7, 23;
Sept. 7

Total samples: Y = 8, 4 = 30

Comment: Holarctic

Agyneta dynica Saaristo & Koponen, 1998
Ecoregion: 2

Collection date: July 22

Total samples: 9 = 0, ¢ =2

Comment: Nearctic; new record for Labrador

Agyneta olivacea (Emerton, 1882)

Ecoregions: |, 2; 5, 6A and B, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 9 = 129, ¢ = 139

Comment: Holarctic; the majority were found in ecoregions
5 and 6, site A; new record for Labrador

Agyneta simplex (Emerton, 1926)

Ecoregions: 1, 2, 3,5, 6A, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 9 = 52, ¢ = 172

Comment: Nearctic; only 13 specimens from the combined
sites at ecoregions 1, 2, and 3; new record for Labrador

Allomengea dentisetis (Grube, 1861)
Ecoregion: 6B

Collection dates: Aug. 12, 25; Sept. 13
Total samples: 9 = 1, ¢ =2
Comment: Holarctic

Bathyphantes eumenis (L. Koch, 1879)
Ecoregions: 1, 2, 3, 6B, 7C and D
Collection dates: July; Aug.; Sept.; Oct.
Total samples: 2 = 12, ¢ = 10
Comment: Holarctic

Bathyphantes pallidus (Banks, 1892)

Ecoregions: 3, 6A and B, 7C

Collection dates: June; July; Aug.; Sept.

Total samples: 2 = 91, ¢ = 44

Comment: Nearctic; the majority were found in ecoregion 6,
site A; only two were found at site B

Carorita limnaea (Crosby & Bishop, 1927)
Ecoregion: 6A

Collection dates: June 17; July 5, 19; Aug. 3
Total samples: 2 = 0, ¢ = 13

Comment: Holarctic; new record for Labrador

Centromerus longibulbus (Emerton, 1882)
Ecoregions: 1, 5, 7C

Collection dates: June 17; July 11, 20

Total samples: ° = 0, ¢ =5

Comment: Nearctic; new record for Labrador

Centromerus sylvaticus (Blackwall, 1841)
Ecoregions: 3, 6B

Collection dates: Aug. 25; Sept. 13, 27; Oct. 27
Total samples: 2 = 13, 6 = 23

Comment: Holarctic

Ceraticelus atriceps (O. P.-Cambridge, 1874)
Ecoregions: 1, 2, 7C and D

Collection dates: Aug. 1, 23; Sept. 7

Total samples: 9 =4, ¢ = 1

Comment: Nearctic

370 THE CANADIAN FIELD-NATURALIST Vol. 128

Ceraticelus crassiceps Chamberlin & Ivie, 1939
Ecoregions: 1, 3

Collection dates: UE: 10; 23; Sept. 7; Oct. 27
Total samples: 2 = 5, d =2

Comment: Neaete

Ceraticelus fissiceps (O. P.-Cambridge, 1874)
Ecoregions: 5, 6A and B

Collection dates: June; July; Aug.; Sept.

Total samples: 9 = 17, ¢ =3

Comment: Nesrcttc

Ceratinella brunnea Emerton, 1882
Ecoregions: 1, 2, 3,5, 6A

Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 28, ¢ =4

Comment: Nenpatte

Ceratinella ornatula (Crosby & Bishop, 1925)
Ecoregion: 7D

Collection date: July 11

Total samples: 2 = 0, ¢ = 1

Comment: Nearctic

Cnephalocotes obscurus (Blackwall, 1834)
Ecoregions: 3, 6A, 7D

Collection dates: June; poy, Aug.; Sept.

Total samples: 2 = 27, ¢ =27

Comment: Holarctic; new record for the province

Diplocentria bidentata (Emerton, 1882)

Ecoregions: 1, 3, 5, 6A and B, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 9 = 63, 4 = 295

Comment: Holarctic; good representation from all sites in
listed ecoregions; new record for Labrador

Diplocentria rectangulata (Emerton, 1915)
Ecoregions: 2, 5, 6A and B, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 45, 4 = 205

Comment: Holarctic; large majority sampled from
ecoregion 5; new record for Labrador

Diplocentria retinax (Crosby & Bishop, 1936)
Ecoregion: 6A

Collection dates: June 17; July 5

Total samples: 9 = 1, ¢ =3

Comment: Nearctic; new record for the province

Diplocephalus subrostratus (O. P.-Cambridge, 1873)
Ecoregion: 6B

Collection dates: June 17; July 5, 19; Ags 35 12525
Total samples: 9 = 19, =27

Comment: Hiplavene: new record for Labrador

Erigone blaesa Crosby & Bishop, 1928
Ecoregions: 4, 5

Collection date: Aug. 7

Total samples: 9 = 0, @ =2

Comment: Nearctic

Estrandia grandaeva (Keyserling, 1886)
Ecoregions: 3, 6B

Collection dates: July 19; Aug. 5

Total samples: 9 =1, ¢=1

Comment: Holarctic

Gonatium crassipalpum Bryant, 1933

Ecoregions: 1, 2, 3, 5, 6A, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: ° = 68, 4 = 62

Comment: eases well represented in all sampled ecore-
gions

Grammonota angusta Dondale, 1959

Ecoregions: 2, 5, 6A

Collection dates: June 17; Aug. 7; Sept. 13; Oct. 27
Total samples: 2 = 7, 6 =0

Comment: peere

Helophora insignis (Blackwall, 1841)
Ecoregions: 2, 3, 6B, 7C

Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 15, 6 = 14

Comment: Holarctic

Hilaira herniosa (Thorell, 1875)

Beorecions le 2) 5) Sone ©

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 9 = 103, ¢ =75

Comment: Holarctic; well represented at all sites in listed
ecoregions

Hybauchenidium gibbosum (Sorenson, 1898)
Ecoregions: 3; 5; 6, Sites A and B; 7, Site C

Collection dates: July 5, 18, 19, 22; Aug. 3; Sept. 13, 27;
Och 7/

Total samples: 9 = 8, d =2

Comment: Holarctic

Improphantes complicatus (Emerton ,1882)
Ecoregions: 1, 2, 3,5, 6A and B, 7C and D
Collection dates: June; ee Aug.; Sept.; Oct.
Total samples: 9 = 49, ¢ = 54

Comment: Holentib

Incestophantes washingtoni (Zorsch, 1937)
Ecoregions: 1, 2, 3, 4, 6B, 7C and D
Collection Os June; pee Aug.; Sept.; Oct.
Total samples: 2 = 30,

Comment: Nésretie

Islandiana flaveola (Banks, 1892)

Ecoregion: 6A

Collection date: a —

Total samples: 2 = 0, ¢ =

Comment: nee new record for Labrador

Islandiana sp. Braendegaard, 1932 (indeterminate: NF)
Ecoregions: 1, 5, 7D

Collection dates: June 17; July 11, 20

Total samples: 2 = 2, ¢=5

Lepthyphantes aes (Emerton, 1882)
Ecoregions: 1, 2, 3,5, 6A and B, 7C
Collection dates: June; fees Aug.; Sept.; Oct.
Total samples: 9 = 57, 4 = 24

Comment: Torarsue: new record for Labrador

Lepthyphantes turbatrix (O. P.-C ambridge, 1877)
Ecoregion: 6A

Collection date: Aug. 12

Total samples: 2 = 1, ¢ =0

Comment: Nearctic; new record for Labrador

2014 PERRY ET AL.: SPIDERS OF THE SOUTHERN TAIGA BIOME OF LABRADOR 371

Lepthyphantes sp. Menge, 1866 (indeterminate; CNC #7)
Ecoregions: 5, 6A and B

Collection date: June 17

Total samples: 9 = 0, ¢ =7

Macrargus multesimus (O. P.-Cambridge, 1875)
Ecoregions: 3, 6A

Collection dates: June 17; Aug. 25; Sept. 13; Oct. 27
Total samples: 2 = 8, ¢ =0

Comment: Holarctic; new record for the province

Maro amplus Dondale & Buckle, 2001
Ecoregion: 6B

Collection date: June 17

Total samples: 9 = 0, 4 =8

Comment: Nearctic; new record for Labrador

Maro nearcticus Dondale & Buckle, 2001
Ecoregion: 6B

Collection date: June 17

Total samples: 2 = 0, ¢ =2

Comment: Nearctic; new record for Labrador

Mermessus entomologicus (Emerton, 1911)
Ecoregion: 3

Collection date: July 22

Total samples: 2 = 0, ¢ = 1

Comment: Nearctic; new record for Labrador

Mermessus trilobatus (Emerton, 1882)
Ecoregion: 6A

Collection dates: July 5, 19

Total samples: 2 = 0, d =3

Comment: Holarctic; new record for Labrador

Mermessus undulatus (Emerton, 1914)
Ecoregion: |

Collection dates: July 20; Aug. 23

Total samples: 2 = 0, ¢ =2

Comment: Nearctic; new record for Labrador

Metopobactrus prominulus (O. P.-Cambridge, 1872)
Ecoregion: 6A

Collection date: July 5

Total samples: 2 = 0, d= 1

Comment: Holarctic; new record for the Province
Microlinyphia mandibulata (Emerton, 1882)
Ecoregions: 3, 7C

Collection date: Sept. 27

Total samples: 2 = 4, ¢ = 0

Comment: Nearctic; new record for Labrador
Microneta viaria (Blackwall, 1841)

Ecoregion: 6A

Collection date: June 17

Total samples: 2 (sex not reported)
Comment: Holarctic; new record for the province

Oreoneta brunnea (Emerton, 1882)

Ecoregions: 1, 2

Collection dates: July 20; Aug. 25; Sept. 7; Oct. 26
Total samples: 2 = 26, d = 24

Comment: Nearctic; new record for Labrador
Oreoneta sp. Kulcynski, 1894 (indeterminate; NF 167)
Ecoregion: 7D

Collection date: July 11

Total samples: 2 = 0, 5 =!

Oreonetides flavescens (Crosby, 1937)
Ecoregion: 6A

Collection date: June 17

Total samples: Y = 0, ¢ = 2

Comment: Nearctic; new record for Labrador

Oreonetides vaginatus (Thorell, 1872)
Ecoregions: 1, 2, 3,5, 6B, 7C and D
Collection dates: June; July; Aug.; Sept.; Oct
Total samples: 2 =6, 0 = 11

Comment: Holarctic

Oreophantes recurvatus (Emerton, 1913)
Ecoregion: 3

Collection date: Oct. 27

Total samples: 9 = 0, ¢ =2

Comment: Nearctic

Pelecopsis mengei (Simon, 1884)

Ecoregions: 3, 6B

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 9 = 19, 6 =9

Comment: Holarctic; only one spider collected from ecore-
gion 3; new record for Labrador

Pelecopsis moesta (Banks, 1892)

Ecoregion: 6A

Collection date: July 5

Total samples: 2 =0, ¢ = 1

Comment: Nearctic; new record for the province

Pityohyphantes subarcticus Chamberlin & Ivie, 1943
Ecoregions: 1, 2, 3, 4, 5, 6A, 7C

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 9 = 14, ¢ =3

Comment: Nearctic

Pocadicnemis americana Millidge, 1976

Ecoregions: 1, 2, 3, 5, 6A, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 2 = 392, ¢ = 405

Comment: Nearctic; 11 came from ecoregions 1, 2, and 3

Pocadicnemis pumila (Blackwall, 1841)
Ecoregion: 6B

Collection date: June 17

Total samples: 2 =0, J = 1

Comment: Nearctic; new record for the province

Poeciloneta calcaratus (Emerton, 1909)
Ecoregions: 1, 3, 4

Collection dates: Aug. 7, 23; Oct. 27
Total samples: 2 = 1, ¢ =2

Comment: Nearctic

Satilatlas sp. Keyserling, 1886 (indeterminate; NF)
Ecoregions: 3, 7D

Collection date: July 11

Total samples: 9 = 0, ¢ =3

Satilatlas sp. Keyserling, 1886 (indeterminate; NF)
Ecoregion: 3

Collection dates: July 22; Sept. 7

Total samples: 2 =2, ¢ =0

Sciastes truncatus (Emerton, 1882)

Ecoregions: 3, 5, 6A and B, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 39, d = 26

Comment: Nearctic; new record for Labrador

372 THE CANADIAN FIELD-NATURALIST

Scironis tarsalis (Emerton, 1911)

Ecoregion: 6B

Collection dates: June 17; July 5; Sept. 13
Total samples: 9 = 3, ¢ =8

Comment: Nearctic; new record for Labrador

Scotinotylus alpinus (Banks, 1896)

Ecoregion: 3

Collection dates: Seph Octe2 7
a —_

Total samples: 9 = 0, 4 =2
Comment: Neareite: new record for the province

Scotinotylus sacer (Crosby, 1929)
Ecoregions: 3, 5, 7C and D

Collection dates: June 17; July 11; Aug. 1, 24, 29; Sept. 7;

Oct. 8
Total samples: 9 = 6, 4 = 11
Comment: Holseto: new record for Labrador

Semljicola obtusus (Emerton, 1915)
Ecoregion: |

Collection date: July 20

Total samples: 2 = 1, 4 =0

Comment: Nearctte: new record for Labrador

Sisicottus montanus (Emerton, 1882)

Ecoregions: 1, 6B, 7D

Collection dates: June 17; July 11, 20; Aug. 6; Oct. 26, 27
Total samples: 9 =5, d =

Comment: Nearctic; new record for Labrador

Sisicus penifusifer Bishop & Crosby, 1938
Ecoregion: 6A

Collection date: July S8

Total samples: 9 =2, ¢=0

Comment: Nearetie: new record for Labrador

Sisis rotundus (Emerton, 1925)

Ecoregions: 5, 6A, 7C

Collection dates: June 17; July 11, 18; Aug. 1, 29
Total samples: 9 = 12, ¢=7

Comment: Nearctio

Stemonyphantes blauveltae Gertsch, 1951
Ecoregion: 2

Collection dates: Aug. 2, 10, 25: Oct. 27
Total samples: 9 =3, 6 =6

Comment: Nearctic; new record for Labrador

Styloctetor stativus (Simon, 1881)

Ecoregions: 5, 6A and B, 7C and D

Collection dates: June 17; July 4, 5, 11; Aug. 1, 2, 3, 29
Total samples: 2 = 9, ¢ = 47

Comment: Holarctic; new record for Labrador

Tapinocyba bicarinata (Emerton, 1913)
Ecoregions: 1, 7C

Collection dates: July 11, 20; Aug. 1, 29
Total samples: 9 = 1, 6 =7

Comment: Nearotes new record for Labrador

Tapinocyba prima Dupérré & Paquin, 2005
Ecoregions: 5, 6A, 7C and D

Collection dates: June; July; Aug.

Total samples: 2 = 3, 4 = 61

Comment: Nearatio: new record for Labrador

Tapinocyba simplex (Emerton, 1882)
Ecoregions: 1, 5, 6A and B

Collection dates: June; pal Aug.

Total samples: 2 = 17,

Comment: Nearetio: new record for Labrador

Vol.

128

Tapinocyha sp. Simon, 1884 (indeterminate; CNC)

Ecoregion: 6A
Collection date: Aug. 3
Total samples: 9 = 1, 3 =0

Tunagyna debilis (Banks, 1892)

Ecoregions: 1, 2, 5, 6A and B, 7C and D
Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: = 14, ¢ =31

Comment: Holarctic; new record for Labrador

Wabasso cacuminatus Millidge, 1984
Ecoregions: 1, 2, 7C and D
Collection dates: July 11,
Total samples: 9 = 22, d= 18

Comment: Holarctic; new record for Labrador

Walckenaeria arctica Millidge, 1983
Ecoregions: 1, 2, 3,5, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 39, ¢ = 22

Comment: Nearctic; new record for Labrador

20, 22; Aug. 1, 6, 29; Oct. 8, 27

Walckenaeria atrotibialis (O. P.-Cambridge, 1878)

Ecoregions: 5, 6A and B, 7C

Collection dates: June; July; Aug.; Sept.

Total samples: 2 = 51, ¢ = 46

Comment: Holarctic; new record for Labrador

Walckenaeria castanea (Emerton, 1882)
Ecoregions: 1, 2, 3,5, 6A, 7C and D
Collection dates: June; July; Aug.; Oct.
Total samples: 9 = 21, ¢ =27
Comment: Nearctic

Walckenaeria clavipalpis Millidge, 1983
Ecoregion: |

Collection date: Aug. 6

Total samples: 9 = 1, ¢=0

Comment: Nearstia new record for Labrador

Walckenaeria communis (Emerton, 1882)
Ecoregions: 1, 2, 3, 5, 6A and B, 7C and D
Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 160, ¢ = 33

Comment: Nearctic; new record for Labrador

Walckenaeria cuspidata brevicula (Crosby & Bishop, 1931)

Ecoregion: |

Collection date: Sept. 7

Total samples: 9 = 1, ¢ =0

Comment: Nearctic; new record for Labrador

Walckenaeria directa (O. P.-Cambridge, 1874)
Ecoregions: 1, 3, 5, 6A and B, 7D

Collection dates: Tune: July; Aug.; Sept.; Oct.
Total samples: 9 = 43, ¢ =9

Comment: Nearctic

Walckenaeria exigua Millidge, 1983
Ecoregions: 2, 3, 5, 6A and B, 7C and D
Collection dates: June; ge Aug.; Sept.; Oct.
Total samples: 2 = 15, 4 = 200

Comment: Nearctic; new record for Labrador

Walckenaeria karpinskii (O. P.- -Cambridge, 1873)

Ecoregions: 5, 6A, 7C and D
Collection dates: June 17; July 4, 11, 18; Aug. 2
Total samples: 2 = 6, 6 =5
Comment: Pioteeade: new record for Labrador

8. 4,

Sept. 12

2014

Walckenaeria lepida (Kulezynski, 1885)
Ecoregion: 4

Collection date: Aug. 7

Total samples: ° = 0, 4 = 1

Comment: Holarctic; new record for Labrador

Walckenaeria spiralis (Emerton, 1882)
Ecoregions: 1, 2,5, 6A, 7C and D

Collection dates: June; July; Aug.; Oct.

Total samples: 2 = 1, 4 = 50

Comment: Holarctic; new record for Labrador

Walckenaeria tricornis (Emerton, 1882)

Ecoregions: 1, 3,5, 6A and B, 7C and D

Collection dates: June; July; Aug.; Sept.

Total samples: ° = 50, ¢ = 179

Comment: Nearctic; one found in September, but the vast
majority were collected in June, July, and at the beginning
of August

Wubana pacifica (Banks, 1896)

Ecoregion: 3

Collection date: Sept. 13

Total samples: 2 = 0, ¢ = 1

Comment: Nearctic; new record for Labrador

Zornella armata (Banks, 1906)

Ecoregions: 1, 3, 6A, 7C

Collection dates: June 17; Sept. 27; Oct. 26, 27
Total samples: 2 = 22, d =7

Comment: Nearctic; new record for Labrador
Indeterminate; NF 158

Ecoregion: |

Collection date: July 20
Total samples: 2 = 0, ¢ = 1
Indeterminate; NF 159

Ecoregion: |

Collection date: July 20
Total samples: 2 = 1, d =0
Indeterminate; NF 160

Ecoregion: 3

Collection date: Oct. 27
Total samples: 2 = 1, ¢ =0
Indeterminate; NF 161

Ecoregion: 3

Collection date: Oct. 27
Total samples: 9 = 1, ¢ =0
Indeterminate; NF 162

Ecoregion: 6A

Collection date: July 19
Total samples: 2 = 1, d =90
Indeterminate; NF 163

Ecoregion: |

Collection date: July 20
Total samples: 2 = 1, d =9
Indeterminate; NF 164

Ecoregion: 3

Collection date: July 22
Total samples: 2 = 1, d =9
Indeterminate; NF 165

Ecoregion: 7D

Collection date: July 11
Total samples: 2 = 1, d =0
Indeterminate; NF 166

PERRY ET AL.: SPIDERS OF THE SOUTHERN TAIGA BIOME OF LABRADOR 373

Ecoregion: 6A
Collection date: June 17
Total samples: Y = 1, ¢ =0

LIOCRANIDAE (1 SPP.)

Agroeca ornata Banks, 1892

Ecoregions: |, 2, 3, 5, 6A and B, 7C and D
Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 71, ¢ = 56

Comment: Holaretic; new record for Labrador

LYCOSIDAE (15 SppP.)

Alopecosa aculeata (Clerck, 1757)
Ecoregions: |, 2, 3, 5, 6A and B, 7C and D
Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 9 = 95, 4 = 272

Comment: Holarctic

Arctosa alpigena (Doleschall, 1852)

Ecoregions: 1, 2,5, 7C and D

Collection dates: July 11, 20; Aug. 1, 29, 23; Sept. 7; Oct. 8,
PHI

Total samples: 2 = 82, 4 = 48

Comment: Holarctic

Arctosa raptor (Kulezynski, 1885)

Ecoregion: |

Collection dates: July 20; Aug. 6

Total samples: 2 = 4, ¢ = 18

Comment: Holarctic; new record for Labrador

Arctosa rubicunda (Keyserling, 1877)
Ecoregion: 3

Collection dates: Aug. 25; Sept. 13

Total samples: 9 = 2, d =0

Comment: Nearctic; new record for Labrador

Hogna frondicola (Emerton, 1885)

Ecoregions: 3, 6A

Collection dates: June17; July 5, 19, 22; Aug. 25; Sept. 13
Total samples: 2 = 8, d =3

Comment: Nearctic; new record for Labrador

Pardosa concinna (Thorell, 1877)
Ecoregions: 1, 2, 3, 5, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 124, ¢ = 256

Comment: Nearctic

Pardosa furcifera (Thorell, 1875)

Ecoregions: 1, 2

Collection dates: July 20, 22; Aug. 6, 23; Sept. 7, 17; Oct. 26,
27

Total samples: 2 = 297, 3 = 326

Comment: Nearctic

Pardosa fuscula (Thorell, 1875)

Ecoregions: 1, 3, 7C

Collection dates: July 20, 22; Aug. 6, 23, 25; Sept. 27; Oct. 26
Total samples: 9 = 12, 4 =7

Comment: Nearctic

Pardosa hyperborea (Thorell, 1872)

Ecoregions: 1, 2, 3, 5, 6A, 7C and D

Collection dates: June; July; Aug.; Sept.; Oct.

Total samples: 2 = 1143, ¢ = 2401

Comment: Holarctic

374 THE CANADIAN FIELD-NATURALIST Vol.

Pardosa mackenziana (Keyserling, 1877)
Ecoregions: 1, 3,5, 6A and B

Collection dates: June 17; July 5, 20, 22; Aug. 3, 5
Total samples: 2 = 26, d = 72

Comment: Nearctic; new record for Labrador

Pardosa uintana Gertsch, 1933

Ecoregions: 1, 2, 3, 4,5, 6A and B, 7C and D
Collection dates: June; July; Aug.; Sept.; Oct.
Total samples: 2 = 439, ¢ = 572

Comment: Nearctic

Pardosa xerampelina (Keyserling, 1877)
Ecoregions: 5, 6B, 7D

Collection dates: June 17; July 11; Aug. 1; Oct. 8
Total samples: 2 = 7, ¢ = 26

Comment: Nearctic

Pirata bryantae Kurata, 1944
Ecoregions: 1, 3, 5, 6A and B, 7C and D
Collection dates: June; July; Aug.; Oct.
Total samples: 2 = 181, 4 = 560
Comment: Nearctic

Pirata piraticus (Clerck, 1757)

Ecoregion: 3

Collection date: Sept. 13

Total samples: 9 = 1, ¢=0

Comment: Holarctic; new record for Labrador

Trochosa terricola Thorell, 1856
Ecoregions: 3, 5, 6A and B, 7C and D
Collection dates: June; July; Aug.; Sept.
Total samples: 2 = 73, 4 = 106
Comment: Holarctic

PHILODROMIDAE (4 spp.)

Philodromus alascensis Keyserling, 1884
Ecoregion: 7D

Collection date: July 11

Total samples: 9 = 1, =0

Comment: Holarctic

Philodromus placidus Banks, 1892
Ecoregion: 6A

Collection date: July 5

Total samples: 9 =0, d =1
Comment: Nearctic

Philodromus rufus quartus Dondale & Redner, 1968
Ecoregions: 5, 6B

Collection dates: June 17; July 4

Total samples: 2 = 2; ¢=0

Comment: Nearctic

Thanatus formicinus (Clerck, 1757)
Ecoregions: 1, 2, 3, 7D

Collection dates: July 11, 22; Aug. 1, 10, 25, 29; Sept. 7,

Io Ove, 27
Total samples: 9 = 17, 4 = 30
Comment: Holarctic; new record for Labrador

SALTICIDAE (3 SPP.)

Evarcha hoyi (Peckham & Peckham, 1883)
Ecoregions: 5, 6B

Collection dates: July 18, 19

Total samples: 9 = 1, 5 =1

Comment: Nearctic; new record for Labrador

Neon nelli Peckham & Peckham, 1888

Ecoregion: 6A and B

Collection dates: June 17; July 4, 5, 19; Aug. 3, 12, 25
Total samples: 9 = 25, ¢ = 29

Comment: Nearctic

Talavera minuta (Banks, 1895)

Ecoregion: 6A

Collection date: July 5

Total samples: 2 =0, ¢ = 1

Comment: Nearctic; new record for the province

TETRAGNAHIDAE (2 SPP.)

Tetragnatha elongata Walckenaer, 1841

Ecoregion: 5

Collection date: Aug. 7

Total samples: 2 =1, d=0

Comment: Nearctic and Neotropical; new record for
Labrador

Tetragnatha extensa (Linnaeus, 1758)
Ecoregion: 5

Collection date: Aug. 7

Total samples: 2 = 0, d =1
Comment: Holarctic

THERIDIIDAE (4 SpP.)

Enoplognatha intrepida (Serenson, 1898)
Ecoregion: 2

Collection dates: Aug. 10, 25; Sept. 7; Oct. 27
Total samples: 9 = 1, ¢ = 16

Comment: Nearctic; new record for Labrador

Robertus fuscus (Emerton, 1894)
Ecoregions: 2, 3, 5, 6B, 7C and D

Collection dates: June 17; July 5; Aug. 25; Sept. 7, 17; Oct. 27

Total samples: 9 =7, ¢ = 15
Comment: Nearctic

Rugathodes sexpunctatus (Emerton, 1882)
Ecoregion: 7D

Collection date: Aug. |

Total samples: 9 = 0, ¢ =1

Comment: Holarctic; new record for Labrador

Theonoe stridula Crosby, 1906
Ecoregions: 5, 6A, 7C and D

Collection dates: June 17; July 4, 5, 11, 19: Aug. 3, 29; Oct. 8

Total samples: 9 = 7, ¢ = 42
Comment: Nearctic; new record for Labrador

THOMISIDAE (10 spp.)
Misumena vatia (Clerck, 1757)
Ecoregion: |

Collection date: July 20

Total samples: 9 = 0, ¢ =
Comment: Holarctic

Ozyptila sincera canadensis Dondale & Redner, 1975
Ecoregions: 2, 3

Collection dates: Aug. 5, 25; Sept. 13, 27

Total samples: 9 =9, 4 =

Comment: Nearctic; new record for Labrador

Xysticus canadensis Gertsch, 1934

Ecoregions: 3, 6A and B, 7C

Collection dates: June 17; July 5, | 1; Aug. 29; Oct. 27
Total samples: 9 =2, 6 =12

Comment: Holarctic

2014

Xysticus durus (Sorenson, 1898)

Ecoregion: 7

Collection date: Aug. |

Total samples: 9 = 1, 4 =0

Comment: Nearctic; new record for the province
Xysticus ellipticus Turnbull, Dondale & Redner, 1965
Ecoregions: 2, 3

Collection dates: July 22; Oct. 27

Total samples: 9 = 0, 4 = 15

Comment: Nearctic; new record for the Province
Xysticus emertoni Keyserling, 1880

Ecoregions: 1, 2, 3, 5, 6A and B, 7D

Collection dates: June 17; July 4, 5, 11, 22; Aug. 1, 5, 10;
Octk27

Total samples: 2 = 6, ¢ = 50

Comment: Holarctic; new record for Labrador
Xysticus keyserlingi Bryant, 1930

Ecoregions: 1, 2, 3, 5, 7D

Collection dates: July; Aug.; Sept.; Oct.

Total samples: 2 = 30, ¢ = 90

Comment: Nearctic; new record for Labrador
Xysticus luctuosus (Blackwall, 1836)
Ecoregions: 1, 2, 3, 7D

Collection dates: July; Aug.; Sept.; Oct.

Total samples: 2 = 5, ¢ = 40

Comment: Holarctic; new record for Labrador
Xysticus obscurus Collett, 1877

Ecoregion: 3

Collection date: Aug. 7

Total samples: 3 (sex unknown)

Comment: Holarctic

Xysticus triguttatus Keyserling, 1880
Ecoregions: 1, 2, 3

Collection dates: July 22; Aug. 5; Oct. 26, 27
Total samples: 2 = 2, ¢ = 14

Comment: Nearctic; new record for Labrador

Discussion
Origins of the Labrador spider fauna

Most (58.4%) of the species identified in this study
have Nearctic distributions; however, a significant Hol-
arctic component was also present (41.6%) (Table 2).
Noticeably absent from the collections were introduced
species or ones previously known only from Palearctic
or other regions.

The frequency of Holarctic species increases as one
moves further north in the Nearctic region (Pickavance
and Dondale 2005) and our data appear to support this.
Of spider species reported for the island of Newfound-
land (primarily south of our study area), 33% have Hol-
arctic distributions (Pickavance and Dondale 2005). In
a more northerly locality, subarctic and arctic Quebec,
the percentage of spider species with Holarctic distribu-
tion is nearly 50% (Koponen 1994). Still further north,
on Belcher Island, the proportion rises to 58% (Kopo-

nen 1992).

Introduced species ae
The absence of introduced species in this study may

be attributed to a combination of factors. First, most of

PERRY ET AL.: SPIDERS OF THE SOUTHERN TAIGA BIOME OF LABRADOR 2)

wn

the sample sites were in relatively pristine old-growth
forests in remote locations. Collection sites near com-
munities were still well outside town boundaries. Also,
the population of Labrador is approximately 29 000,
and species introductions may be less likely to occur
there than in more heavily populated areas elsewhere.
The severe cold of Labrador winters may also limit the
colonization and spread of more southerly exotics. With
the creation of the new Trans-Labrador Highway, spe-
cies introductions may increase and it will be interest-
ing to continue to monitor spider diversity in the study
area to examine influences of the new highway on
species introductions.

Noteworthy occurrences

In total, 161 species were identified in this collec-
tion, 89 of which are new records for Labrador. Of the
89 new species, 16 species are new records for the
province. The former species complement reported for
Labrador was 124 (Paquin ef a/. 2010); our addition
of 89 species raises the new species total to 213. For
the province, the total number of spider species has
been raised to 395: 361 (Newfoundland total) + 18
(reported only in Labrador) + 16 (new records).

Of the 16 species records new to the province, almost
all can be found as far east as Quebec or New Bruns-
wick and, therefore, it is not surprising that they can
be found in Newfoundland and Labrador. However, a
few stand out as noteworthy. The collection of Arcto-
bius agelenoides (Amaurobiidae) is of interest because,
in the Nearctic region, this has been considered a west-
ern species (Marusik and Koponen 2005) recorded in
Canada from Yukon, Northwest Territories, Nunavut,
British Columbia, Alberta, Saskatchewan, and Mani-
toba (Paquin et al. 2010). Arctobius agelenoides may
have a continuous distribution across Canada or per-
haps the Labrador population is disjunct.

Hackmania prominula (Dictynidae) is relatively
rarely encountered. It is a northern Holarctic species
previously reported in western North America from
Alaska, Yukon, British Columbia, Alberta, Saskatche-
wan, and Manitoba (Paquin et a/. 2010). Our Labrador
records are the first for eastern North America.

Haplodrassus eunis (Gnaphosidae) is primarily a
western species with records from Alaska to California
and eastward to the Great Lakes (Platnick and Dondale
1992). Our report indicates that its distribution extends
throughout the North of the Neararctic region Our
record of Gnaphosa parvula also extends this species
range across the north of the Nearctic region.

Agyneta dynica (Linyphiidae) is a rarely collected
Nearctic endemic reported in Canada only from Yukon,
subarctic Quebec, and on parts of the subarctic barrens
of the Northern Peninsula of Newfoundland (Picka-
vance and Dondale 2005; Paquin ef al. 2010; Dupérre
2013). Our Labrador record helps define the true dis-
tribution of this species.

With the addition of 89 species, the total known
species complement for Labrador stands at 213. It is

376

likely that this total is not yet complete; much of Lab-
rador remains to be surveyed. In addition, this survey
was largely confined to the ground and, therefore, spi-
ders that make their homes in trees and shrubs are most
likely underrepresented. Further, this collection was
confined to lower elevations in the southerly latitudes
of Labrador. To reveal the full species complement of
the northern fauna, further sampling is still required
north of 54° latitude and at higher elevations.

Acknowledgements

We are indebted to Dr. Jaime Pinzon, Dr. Robert
Bennett and an anonymous reviewer whose editorial
comments and shared knowledge were invaluable in
the completion of this work. We thank conservation
officer Chuck Porter, senior biologist Rebecca Jeffery,
and field technician Ted Pardy for routinely collecting
our pitfall trap samples. We also thank Dara Walsh for
sorting and collating the collection and Carl Marks for
his global information system contributions. Finally, we
would like to thank the many natural resource officers
who participated in the initial ad hoc survey; their ef-
forts ultimately inspired the completion of the larger
survey.

Literature Cited

Anderson, T. C. 1985. The Rivers of Labrador. Canadian Spe-
cial Publication of Fisheries and Aquatic. Sciences 81. Fish-
eries and Oceans, Ottawa, Ontario, Canada. 389 pages.

Dupérré, N. 2013. Taxonomic revision of the spider genera
Agyneta and Tennesseellum (Araneae, Linyphiidae) of North
America north of Mexico with a study of the embolic divi-
sion within Micronetinae sensu Saaristo & Tanasevitch
1996. Zootaxa 3674: 1-189.

Dyke, A. S., J. T. Andrews, P. U. Clark, J. H. England,
G. H. Miller, J. Shaw, J. J. Veillette. 2002. The Lauren-

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tide and Innuitian ice sheets during the last glacial maxi-
mum. Quaternary Science Reviews 21: 9-31.

Hackman, W. 1954. The Spiders of Newfoundland. Acta
Zoologica Fennica 79: 1-99.

Koponen, S. 1992. Spider fauna (Araneae) of the low Arc-
tic Belcher Islands, Hudson Bay. Arctic 45(4):358-362.
Koponen, S. 1994. Ground-living spiders, opilionids, and
pseudoscorpions of peatlands in Quebec. Memoirs of the
Entomological Society of Canada 126 (suppl. 169): 41-60.

Marusik, Y. M. and S. Koponen. 2005. A survey of spiders
(Araneae) with Holarctic distribution. Journal of Archnolo-
gy 33 (2): 300-305.

Meades, S. J. 1990. Natural regions of Newfoundland and
Labrador. Technical report. Protected Areas Association, St.
John’s, Newfoundland and Labrador, Canada. 373 pages.

Paquin, P., D. J. Buckle, N. Dupérré, and C. D. Dondale.
2010. Checklist of the spiders (Araneae) of Canada and
Alaska. Zootaxa 2461: 1-170.

Pickavance, J. R., and C. D. Dondale. 2005. An annotated
checklist of the spiders of Newfoundland. Canadian Field-
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.canadianfieldnaturalist.ca/index.php/cfn/article/viewFile
/114/114.

Platnick, N. I. 2014. The world spider catalog, version
14.5. American Museum of Natural History, New York,
New York, USA. Accessed March 2014. http://research
.amnh.org/iz/spiders/catalog/.

Platnick, N. I., and C. D. Dondale. 1992. The Insects and
Arachnids of Canada Part 19: The Ground Spiders of Cana-
da and Alaska (Araneae: Gnaphosidae). Research Branch,
Agriculture Canada, Ottawa, Ontario, Canada. Publication
1875. 297 pages.

Spence, J. R., and J. K. Niemela. 1994. Sampling carabid
assemblages with pitfall traps: the madness and the method.
The Canadian Entomologist 126(3): 881-894.

Received 30 January 2014
Accepted 28 March 2014

Morphological and Ecological Variation among Populations and

Subspecies of Burbot (Lota lota [L, 1758]) from the Mackenzie River
Delta, Canada

HANS RECKNAGEL!, AMy AMos2:3, and KATHRYN R. ELMER!

Institute of Biodiversity, Animal Health & Comparative Medicine, College of Medical, Veterinary & Life Sciences, University
ee. of Glasgow, Glasgow G12 8QQ United Kingdom

wich in Renewable Resources Board, PO Box 2240, Inuvik, Northwest Territories XOE 0T0 Canada

Corresponding author: aamos@grrb.nt.ca

Recknagel, Hans, Amy Amos, and Kathryn R. Elmer. 2014. Morphological and ecological variation among populations and sub-
eon Burbot (Lota lota [L, 1758]) from the Mackenzie River Delta, Canada. Canadian Field-Naturalist 128(4):
377-384.

The Burbot (Lota /ota [{L, 1758]) is a holarctic distributed freshwater fish in the Gadidae family. In northwestern Canada, it has
an important value for local and traditional fisheries. We describe the morphology and ecology of Burbot from four populations
in the Mackenzie River Delta. Two subspecies come into contact in this area, which is the western edge of Lota lota maculosa
distribution (one population in our study) and the eastern edge of Lota /ota lota distribution (three populations in our study).
We found the combined length—weight relationship in these four populations to be log,, (weight) = —3.986 + 2.617 * log
(length). There was no difference in mean body length (overall mean and standard deviation 73.4 + 8.7 cm), although the L.
lL. /ota populations were heavier than the L. /. maculosa population. All populations differed in their carbon and nitrogen stable
isotope signatures (6!°N overall mean 12.1 + 1.59; 6'°C overall mean —25.54 + 1.11). Main prey items were four fish species:
Ninespine Stickleback (Pungitius pungitius [L, 1758]), Broad Whitefish (Coregonus nasus [Pallas, 1776]), Burbot, and Northern
Pike (Esox lucius [L, 1758]). Ecological and morphological differences may be due to local conditions and population variability
or to the different Lota lota subspecies. More ecological and morphological information from this region is needed to resolve
possible phenotypic differences between the subspecies.

Key Words: Burbot; Lota lota; Mackenzie River; weight—length relationship; sexual dimorphism; ecological variation; parapatric

subspecies; freshwater fish; loche

Introduction

The Burbot (Lota Jota [L, 1758]) is a northern fish
species with a circumpolar distribution in freshwater
rivers, creeks, and lakes. Burbot live in cold lakes and
rivers, where they tend to frequent deep water near the
bottom (McPhail and Paragamian 2000). They can at-
tain a length of | mand a weight of 8 kg, although most
fish are 0.3—0.6 m in length and weigh 1—3 kg (McPhail
and Paragamian 2000). The Mackenzie River Delta
(Northwest Territories, Canada) is one of the northern-
most ranges of Burbot distribution. In this area, they are
fished for their liver and eggs, which are their most val-
ued parts (Gwich’in Elders 1999). Recently, there have
been increased reports of Burbot with malformations,
poor muscle tone, unpleasant tasting flesh, and diseased
livers from local and traditional fishers in the Gwich’in
Settlement Area (Thompson 2008). This prompted re-
search by the Gwich’in Renewable Resources Board,
the data from which are the basis of this study.

Burbot are great dispersers and individuals may cov-
er extensive geographic areas, perhaps driven by spawn-
ing behaviour and the search for food (McPhail and
Paragamian 2000; Slavik e¢ al. 2005). Their spawn-
ing period ranges from winter (December) until early
spring (March), when water is often still covered by ice
(Gwich’in Elders 1999; McPhail and Paragamian 2000).

Little is known about spawning behaviour in nature,
but it seems to be polygynous broadcast spawning, as
Burbot form spawning aggregations where a few fe-
males are surrounded by many males (McCrimmon
1959). There is no evidence of sex-biased dispersal
based on population genetic analyses (Elmer er al.
2012). Whether Burbot have defined home ranges de-
pends on their habitat. In lakes, Burbot do not keep
stable positions as food availability is low and dispersed,
whereas, in rivers, higher food availability permits them
to occupy defined home ranges (Slavik et a/. 2005).
Burbot home ranges are much bigger than those of other
temperate freshwater fish, with measured movements
of up to 255 km (McPhail and Paragamian 2000).
There are two subspecies of Burbot: Lota lota mac-
ulosa is restricted to North America and Lota lota lota
is distributed across Eurasia and Beringia. Based on
population genetic analyses, Elmer ev al. (2012) locat-
ed a precise contact zone between Burbot subspecies in
the Mackenzie River Delta. At Tsiigehtchic (included
in the current study), most individuals collected were
L. l. maculosa and upriver of that locality only L. /.
maculosa were found. Downstream from Tsiigehtchic,
all populations in the Mackenzie River Delta were
exclusively L. /. Jota. Therefore, the lower Mackenzie
River is where the discrete parapatric distributions of

377

378

the Burbot subspecies meet at a narrow contact zone
and there is very little hybridization.

Some biologists consider the two subspecies to be
morphologically variable. The subspecies may differ in
caudal—peduncle ratio, the size of the pectoral fins,
and the number of pyloric caeca (e.g., Pivnicka 1970;
McPhail and Paragamian 2000), but some authors (e.g.,
McPhail and Lindsey 1970; Scott and Crossman 1973)
have declined to distinguish subspecies because sup-
posedly diagnostic traits are highly variable. Weight
is thought to differ between fish from lentic and lotic
environments and possibly between the two subspecies
as well (Fisher et a/. 1996). The causes for this mor-
phological variation are not clear (McPhail 1997). To
some extent, the variation might be a result of repeat-
ed isolation in refugia during multiple glacial cycles,
which led to different genetic lineages and even sub-
species (Van Houdt ef al. 2005; Elmer et al. 2008).
Morphological differences between fluvial and lacus-
trine Burbot have also been observed and led to the
conclusion that lotic and lentic morphologies may be
adapted to alternative contemporary niches (McPhail
1997). To date, there is a dearth of information about
the morphological, meristic, and ecological variability
of this wide-ranging species and, unfortunately, much
of it is only available in grey literature (e.g., McPhail
1997; although see recent research advances by Cott
et al. 2013a,b).

In this study, we describe and compare aspects of
ecological differentiation and morphological variability
among Burbot populations in the Mackenzie River
Delta. As the two subspecies abut in this zone (Elmer
et al. 2008, 2012) and there is little or no information
available about possible ecological and reproductive
barriers between subspecies (McPhail 1997), our aim
was to assess variability in body size and weight, gona-
dosomatic index, and ecology (trophic niche and diet)
and highlight possible areas for future research.

Study Area

Burbot were caught during winter 2007 and 2008
by line hooking at four traditional Burbot fishing loca-
tions in the Mackenzie River Delta by Gwich’in com-
munity fish monitors (Table 1, Figure 1). In Inuvik,
two localities were fished: at Bombadeer Creek down-
stream from a sewage lagoon and at Sam Arey’s Creek

THE CANADIAN FIELD-NATURALIST

Vol. 128

just upstream. In Aklavik, all fish were collected from
Jackfish Creek, across from town. In Tsiigehtchic, Bur-
bot were mostly collected from the Arctic Red River
but some were captured from the Mackenzie River
within 10 km of Tsiigehtchic. For Fort McPherson, all
locations were Peel River tributary creeks (Woody Elias
Creek, Basook Creek, Nelson Creek, Husky River
Creek, Hudson Bay Creek, Rotten Eye Creek) within
50 km of Fort McPherson.

Methods

Total length was measured to the nearest millimetre.
Wet weight of each fish was taken to the nearest gram.
Stomachs were weighed and examined for species of
macrofauna by a Gwich’in Renewable Resources Board
technician. Gonads were extracted and weighed for
fecundity analysis using the gonadosomatic index (GSI
= [gonad weight/body weight] < 100). Fish ages were
inferred from otoliths by A.A. in 2007 and under con-
tract by North/South Consultants Inc. in 2008. A mus-
cle tissue sample was collected and stable isotope meas-
urements were taken by Environment Canada using
mass spectrometry.

Statistical analyses

We tested whether morphological (weight and
length) and ecological characteristics (stable isotopes,
stomach content) differed among populations using
analysis of covariance (ANCOVA) with age and sex as
covariates to correct for age-specific differences and
sexual dimorphism. Stomach content data were first
transformed into a contingency table and then analyzed
using a generalized linear model and a 7? test with prey
type as a covariate. In case a characteristic differed sig-
nificantly among populations, all populations were con-
trasted using generalized linear models, setting sex and
age as fixed effects as they are likely to influence mor-
phological and ecological characteristics (i.e., older fish
are usually larger, diet may shift during ontogeny, etc.)
to determine which populations differed from each oth-
er. In these pairwise population comparisons, we cor-
rected for multiple testing by using the Bonferroni cor-
rection. We conducted statistical analyses in R version
3.0.2 open-source programming language. We pooled
individual data by population to compare means and
standard deviations. We calculated weight—length rela-
tionships using weight in grams and length in milli-

TABLE |. Sampling location and dates for Burbot subspecies (Lota /ota lota and L. 1. maculosa) collected from four popula-
tions in the Mackenzie River Delta, Northwest Territories, Canada.

Population* A Latitude, °N

Longitude, °W Collection dates

Lota lota lota

Inuvik 94 68.4
Aklavik ies 68.3
Fort McPherson 137 67.7
Lota lota maculosa

Tsiigehtchic 28 67.4

133.9 Feb. and May 2007, Jan. and Feb. 2008
134.9 Novy. 2007, Jan. 2008

134.7 Nov. 2607, Feb. 2008

133.8 Nov. 2007

*Subspecies inferred from Elmer ef al. (2012)

2014

NX

-

Te

Aklavik

|
(SE

a %
Fort McPherson

JOAIY [B98d
VW

RECKNAGEL ET. 4L: ECOLOGICAL VARIATION IN MACKENZIE DELTA BURBOT

379

RAL Eos, |
¥ Norhwest-Territories .

& c sd / xX
NA

FicurE |. Sampling locations (black dots) for Burbot subspecies (Lota lota lota and L. |. maculosa) in the Mackenzie River

Delta, Northwest Territories, Canada.

metres across all populations (following Fisher eg al.
1996). We compared GSI for the two sexes using
ANOVA.

Results
Age

The individuals in our samples were mostly older
adults (mean age 13.6 years) (Table 2). Mean ages were
approximately equal across populations, except for the
Tsiigehtchic population, which was older (mean 14.9
years, range 9-25 years, F = 3.559, P =0.0176; Table
2). The age difference was significant only between
the Fort McPherson and Tsiigehtchic populations (¢ =
3.221. P=0.0018). Therefore, our ecological and mor-
phological analyses are unlikely to be influenced by
varying age effects across populations; however, we
performed subsequent statistical analyses with age as
a covariate.

Sexual dimorphism s
Across all populations, female and male Burbot dif-

fered significantly in weight and length, females being
longer (mean [+ standard deviation] 757 + 82 mm ver-
sus 683.4 + 72 mm, F = 10.15, P= 0.002) and heavier
(mean 3699 + 1240 g versus 2785 + 891 g, F = 13.16,
P = 0.0005) than males. Diet did not differ significant-

ly between the sexes, as indicated by stomach contents
(F = 3.82, P = 0.35) and stable isotope (6'°N: F = 1.65,
P=0.20; 63C: F=0.24, P = 0.63) analyses. Mean GSI
differed significantly between the sexes (F = 303.69,
P <0.0001), and was on average 5.66 + 1.58 (n = 245)
for mature females and 11.26 + 4.26 (n = 84) for
mature males (Table 2).

Morphological variation across populations

Body length differed significantly across popula-
tions (F = 10.51, P< 0.0001). Mean length was less in
the Tsiigehtchic population (707.5 + 73.8 mm) com-
pared with the other three populations (Aklavik 731.3
+ 78.0 mm, Fort McPherson 726.5 + 97.3 mm, Inuvik
748.7 + 86.3 mm), but only Fort McPherson and Ak-
lavik (¢ = —3.762, P = 0.0003) and Inuvik and Fort
McPherson (t = —3.428, P = 0.0009) differed signifi-
cantly from each other after Bonferroni correction
(Tables 2 and 3).

Weight differed significantly between populations
(F = 20.37, P < 0.0001) and ranged from 2471.0 +
829.2 g in Tsiigehtchic up to 3523.9 + 1304.0 g in Inu-
vik (Table 2). The Tsiigehtchic population had the low-
est mean weight and, overall, three of the six pairwise
comparisons of weight were significantly different

(Table 3).

THE CANADIAN FIELD-NATURALIST Vol. 128

12.73 (1.34)
MESS (eZ )
WAZ ((eréSy)
12.10 (1.59)

8'5N, mean (SD)
13.43 (1.32)

Son GROG)

83C, mean (SD)
—24.49 (0.87)
—26.03 (1.05)
—26.02 (0.86)
= Sosa lle ils)

Male
GSI
12.38 (2.58)
12.32 (4.22)
NOS on(Sa7.9))
7.91 (5.99)
11.26 (4.26)

Female
GSI
Opn (e28))
5.65 (1.44)
SE (le)
4.30 (3.13)
5.66 (1.58)

years (SD)
1373723)
13-7 (2.0)
13.2 (2.4)
14.9 (4.0)
IB AG1(@e5))

Age, mean

Weight,
mean g (SD)
3419.5 (1269.3)
2471.0 (829.2)
3391.0 (1229.3)

3523.9 (1304.0)
3473.0 (1112.8)

Length,
mean mm (SD)
748.7 (86.3)
731.3 (78.0)
726.5 (97.3)
707.5 (73.8)
732.4 (8.8)

TABLE 2. Characteristics of Burbot subspecies (Lota /ota lota and L. 1. maculosa) collected from four populations in the Mackenzie River Delta, Northwest Territories, Canada.

Note: 6'°C and 6'°N = stable isotopes of carbon and nitrogen, GSI = gonadosomatic index, SD = standard deviation.

Lota lota maculosa

Tsiigehtchic

Fort McPherson
All

Population
Lota lota lota
Inuvik
Aklavik

Across populations and sexes combined, the length-
weight relationship was log, (weight) = 3.986 + 2.617
* log, (length). For subspecies L. /. /ota (1.¢., Burbot
from Inuvik, Aklavik, and Fort McPherson populations)
the length—weight relationship was log, ,(weight) =
—3.882 + 2.583 * log, (length) (n = 311). For subspecies
L. 1. maculosa (1.e., Burbot from the Tsiigehtchic pop-
ulation) the length—weight relationship was log,,
(weight) = —5.458 + 3.102 * log, (length) (” = 27).

Ecological variation across populations

The mean 6!°N in the four Burbot populations
ranged from 11.33 + 1.21 to 13.43 + 1.32. Values of
63C ranged from —24.49 + 0.87 to —26.03 + 1.05
(Table 2). Populations differed significantly in stable
isotope composition, for both 6!°C and 6'°N (Table 3,
Figure 2). Populations from Inuvik and Fort McPher-
son differed most, both in 6!°N (P < 0.001) and 6C
values (P < 0.001) (Table 3). Northern populations
tended to differ significantly in their 6'°N isotope com-
position when compared to southern populations (Fort
McPherson versus Aklavik: P = 0.013, Inuvik versus
Fort McPherson: P < 0.0001, Tsiigehtchic versus Inu-
vik: P = 0.002) except for the populations Tsiigehtchic
versus Aklavik (P = 0.176), although Fort McPherson
versus Aklavik was not statistically significant after
Bonferroni correction at a = 0.05 (P > 0.0083). Com-
parisons of 6!°N isotope composition within the two
northern populations (Inuvik—Aklavik) and the two
southern populations (Tsiigehtchic-Fort McPherson)
were not significantly different. The opposite was the
case for 6'°C isotope composition, where most popula-
tions differed significantly, except for two comparisons
between north and south populations (Fort McPher-
son—Aklavik and Tsiigehtchic—Inuvik, the latter not sig-
nificantly after Bonferroni correction),

Stomach contents of Burbot consisted mainly of four
fish species: Ninespine Stickleback (Pungitius pungi-
tius [L, 1758]), Broad Whitefish (Coregonus nasus
[Pallas, 1776]), Burbot, and Northern Pike (Esox lucius
[L, 1758]). Aside from fish, some stomachs contained
pebbles or vegetation and one fish’s stomach (from Tsi-
igehtchic) contained a mouse; however, overall, non-
fish contents were extremely rare (data not shown). The
absolute number of prey items in a stomach varied quite
widely (data not shown). Overall Northern Pike was
the most frequent prey item (53-66%). Populations dif-
fered significantly in which prey items they had con-
sumed (x7 = 2054.8, df= 9, P< 0.001, Table 3), Tsiige-
htchic differed significantly from Aklavik (P = 0.003)
and Inuvik (P = 0.003) populations and from the Fort
McPherson (P = 0.028) population, although this lat-
ter difference was not significant after Bonferroni
correction (Table 3). The most prominent differences
in consumed prey items between Tsiigehtchic and the
other populations were that the Tsiigehtchic populations
lacked Burbot, had few Ninespine Sticklebacks. and
had a high proportion of Broad Whitefish (Figure 3).

2014 RECKNAGEL ET. AL: ECOLOGICAL VARIATION IN MACKENZIE DELTA BURBOT 38]

TABLE 3. Statistical analyses of morphological and ecological differences between populations of Burbot subspecies (Lota
lota lota and L. |. maculosa) collected from four populations in the Mackenzie River Delta, Northwest Territories, Canada,
using analysis of covariance (ANCOVA) with generalized linear model and F or 72 tests. A. Interactions between population
and age, length, weight, and trophic parameters. A significant interaction indicates that populations differed in that parameter.
B. Uncorrected probability values from further population pairwise comparisons indicate which populations differed from
each other in the parameters identified in A as differing significantly (P < 0.001).

| eee SaaS ass
—_—_—_—<—_—_—COEEEEEOE——EEEE ne

A Parameter n df F value P value
Age 88 3 3.70 0.0176
Length 338 3 10.51 <().001
Weight 338 3 20.37 <().001
5PN 103 3 7.16 < 0.001
SC 103 3 12.34 < 0.001
Parameter n df we P value
Stomach contents 322 9 2054.8 < 0.001

Probability values
B Populations Weight Length 55N 6BC Stomach contents
FM-AK 0.009 0.0003** 0.013 0.864 0.109
IN-AK 0.129 0.758 0.113 =) 0001"*= 0.905
TS-AK. < 0.0001*** 0.022 0.176 0.004* 0.003*
IN-FM = 0 00015s* 0.0009** =, 00014** = 00001+** 0.143
TS-FM 0.031 0.14 0.217 0.0008** 0.028
TS-IN <'0:0001*** 0.047 0.002* 0.043 0.003*

Note: df = degrees of freedom, IN = Inuvik, AK = Aklavik, FM = Fort McPherson, and TS = Tsiigehtchic.
Note: Bonferroni corrected values at a of 0.05 (* = P < 0.0083), 0.01 (** = P < 0.0017) or 0.001 (*** = P < 0.0002).

re oS SSS

5°C

-29 ———"_ I I I 1 I I I 1
t 8 9 10 11 12 13 14 15 16

5 '°N

FIGURE 2. Individual carbon and nitrogen stable isotope composition of four populations of Burbot subspecies (Lota lota lota
and L. L maculosa) from the Mackenzie River Delta, Northwest Territories, C anada. Populations are circumscribed by
75% confidence ellipses: IN = Inuvik (asterisks and dotted line), AK= Aklavik (black circles and solid line), FM = Fort
McPherson (grey circles and grey line), and TS = Tsiigehtchic (open circles and dashed line).

Proportion

TS

AK FM

Population
Burbot (Broad Whitefish

Northern Pike E Ninespine Stickleback

FiGuRE 3. Average proportion of four main prey items found in
the stomachs of four populations of Burbot subspecies
(Lota lota lota and L. |. maculosa) from the Mackenzie
River Delta, Northwest Territories, Canada: IN = Inu-
vik (n = 87), AK = Aklavik, (n = 120), FM = Fort
McPherson (7 = 105), and TS = Tsiigehtchic (n = 10).

Discussion
Sexual dimorphism

We found that, consistently across populations, fe-
males had greater total length and were heavier than
males. In contrast, a study by Cott et al. (2013a) found
a low level of sexual dimorphism in Burbot and no
differences in body size between males and females,
and, in general, it has been suggested that there is lit-
tle sexual dimorphism in terms of size of Burbot. In
addition, we found GSI to be significantly different
between females and males (means 5.66 and 11.26.
respectively). In a study spanning lakes in the L. /.
maculosa range across Canada, Cott et al. (2013b)
also found a higher GSI among males, with GSI in
both sexes at spawning time similar to our findings.
Compared with other fish species, Burbot has a rela-
tively high value of GSI, which suggests that their
reproductive behaviour includes external fertilization
and high communal spawning (Stockley ef al. 1997;
Cott et al. 2013a). This is in agreement with observa-
tions from nature that Burbot spawn in aggregations
with many males surrounding one or two females and
that Burbot are broadcast spawners (McPhail and Para-
gamian 2000). The high GSI also indicates that sperm
competition may be present in Burbot males (Stockley
et al. 1997; Bekkevold et al. 2002; Cott et al. 2013a).
Other research has suggested that GSI in Burbot can
be highly variable at spawning time (Wiggs 1974);
thus, this may warrant further investigation.

Morphology

Populations varied significantly in weight and length,
The sampled population from Tsiigehtchic weighed less
than other populations and, on average, was also small-
er in total length, although the differences were not al-

THE CANADIAN FIELD-NATURALIST

Vol.

ways statistically significant when controlled for age,
sex, and multiple testing. Variation in Burbot morphol-
ogy has been proposed to be a result of different life
histories, as lotic Burbot are exposed to a different envi-
ronment than lentic Burbot, and from their evolutionary
history, as isolation during several cycles of glaciation
has led to different genetic lineages (Fisher ef al. 1996;
McPhail and Paragamian 2000; Elmer et a/. 2008).
Therefore, the variation we found in weight and length
may result from a different evolutionary history, as fish
from Tsiigehtchic genetically belong to the L. /. macu-
losa lineage (Elmer et a/. 2012) in contrast to the oth-
er populations, which belong to the L. /. /ota lineage.
Because our sampling included only one L. /. maculosa
population, Tsiigehtchic, we cannot conclude that ob-
served differences in length, weight, and diet were a
result of ecological differences between the two sub-
species rather than simply local interpopulation vari-
ation.

The weight—length relationship of Burbot in the
Mackenzie River Delta had an intercept of —3.986 and
a slope of +2.617 when all populations were combined.
This relationship differed between L. 1. maculosa and
L. 1. lota populations, consistent with differences found
for each variable separately. Fisher et al. (1996) found
that Burbot have a slightly different weight-length
relationship, on average, across 79 populations, with
an intercept of —4.868 and a slope of +2.898, and
recommended only including individuals longer than
20 cm because weight measurements tend to be impre-
cise for small fish. All individuals in our analysis were
considerably longer than 20 cm; therefore, breadth of
initial sampling is not likely to explain differences
between studies. Although Fisher ef a/. (1996) found
considerable local variation in the Burbot weight—
length relationship across its range, other research has
found relatively little variation in size across lakes that
differed in climate, productivity, or ecosystem size (Cott
et al. 2013b). The extent of morphological variation
may depend somewhat on environmental features, such
as lentic versus lotic water bodies as well as post-glacial
patterns, and deserves further research effort.
Ecological variation

Heavier isotopes are concentrated with ascent up
the trophic chain, such that 5!°N composition is higher
in carnivores than herbivores (Post 2002). We found
Burbot are situated in a high trophic position as indi-
cated by their relatively high 8'5N. In a study con-
ducted by Hesslein et al. (1991) in the Mackenzie Riv-
er Delta, the highest trophic level was achieved by
Northern Pike and Lake Trout (Salvelinus namaycush
[Walbaum in Artedi, 1792]), with values of 85N of
12.3—-15 in Travaillant Lake and 12-14 in Kukjuktuk
Creek. Burbot were not included in that study. We
found Burbot to have a mean 8!5N of 12.1, which is
Close to the level of the top predators, Lake Trout and
Northern Pike, in Hesslein et al. (1991). A study inves-
tigating stable isotopes of Burbot and other piscivo-

2014

rous fish from water bodies close to Great Slave Lake
in Canada (approximately 1400 km upstream from the
Mackenzie River Delta) found that Burbot occupy a
high trophic position, similar to Lake Trout and higher
than Northern Pike and Lake Whitefish (Coregonus
clupeaformis {Mitchill, 1818]) (Cott e¢ al. 2011 ). Vari-
ation in 6'°C could result from differences in abun-
dance, interactions, and composition of organisms at
lower trophic levels (primary producers and herbivores)
and, therefore, provides information about the ultimate
energy source of higher consumers, as the ratio changes
little moving up the food chain (Zanden and Rasmus-
sen 1999; Post 2002). Our findings place Burbot with a
carbon signature similar to that found for Lake White-
fish and Broad Whitefish in previous studies (e.g
Hesslein ef al. 1991).

The trophic niche inferred from stable isotope lev-
els is also supported by our analysis of stomach con-
tents, which consisted almost exclusively of fish prey.
Other authors have noted that adult Burbot are pis-
civorous, with usually more than 80% of their diet con-
sisting of fish and the remainder of macroinvertebrates
and insects (Bailey 1972; McPhail and Paragamian
2000; Amundsen e¢ a/. 2003). Burbot in the Great
Lakes may eat a greater variety of fish prey species
(Bailey 1972; Fratt et a/. 1997). In our study, stomach
contents consisted almost entirely of fish and of only
four species. Altogether, the stable isotope data and the
stomach content analyses underline the piscivorous
lifestyle of Burbot and their high trophic position in
aquatic food webs.

Burbot populations differed significantly in 5'°N
and 63C. We found differences in 6!°N between popu-
lations from Fort McPherson and Inuvik and between
those from Tsiigehtchic and Inuvik, but no other pop-
ulations differed. In contrast all populations differed in
5'3C composition except for Aklavik—Fort McPherson
and Tsiigehtchic—Inuvik. Values of 5'°N and 6'°C are
known to differ within species between different water
bodies, at least partly due to variable baselines (e.g.,
Cott et al. 2011). However, our stomach content analy-
ses confirm that there are at least subtle differences in
ecology between populations. As in morphological
analyses, Tsiigehtchic was the most divergent popu-
lation and significantly differed from two of the other
three populations. In contrast to other populations,
Burbot from Tsiigehtchic showed no cannibalism and
their stomachs contained a high proportion of Lake
Whitefish. .

The trophic analysis indicates complex interactions
between Burbot and their resources, which differ local-
ly, leading to variation in consumed prey and the signals
of stable isotopes. It is important to note that stable
isotope composition reflects a long-term (weeks or
months) integration of diet, whereas stomach contents
are a snapshot of the day of collection and are inher-
ently more variable. As observed in morphological
analysis, Burbot caught in Tsiigehtchic were most dif-

a |

RECKNAGEL ET. AL: ECOLOGICAL VARIATION IN MACKENZIE DELTA BURBOT

ferent ecologically from Burbot caught in other loca-
tions, but it remains to be tested whether this difference
is a result of local interspecific variation or fixed be-
tween subspecies.

Other research found that Burbot from Tsiigehtchic
had fewer liver parasites than fish from Aklavik, Inu-
vik, or Fort McPherson (Goater 2010*). In other fresh-
water fish, parasite load has been found to be tightly
correlated with trophic niche; pelagic fish have a high-
er parasite load, even within the same lake and the
same species (Knudsen e¢ a/. 2013). More intensive
sampling near the contact zone would be necessary
to address the interesting possibility that either sub-
species or localities differ in ecology or parasite sus-
ceptibility.

Implications for future research

Future research efforts should target the contact
zone and its surrounding areas more intensively. Our
study suggests that the Tsiigehtchic population (1.¢., the
only L. 1. maculosa population) may differ in body
length compared with all other sampled populations,
which were L. /. ota. The extent to which this corre-
lates with other morphological and ecological traits,
such as trophic niche and dietary preferences, is less
clear. Although we were unable to distinguish between
population and subspecies differences because we sam-
pled only one L. /. maculosa population, our study
illuminates several lines of future research. From an
ecological and morphological aspect, it would be in-
teresting to identify the differences — if any exist —
between Burbot subspecies. From a genetic perspec-
tive, previously identified low levels of admixture at
the contact zone (Elmer et a/. 2012) may provide an
opportunity to hone in on the genetic basis of traits that
differ between subspecies through high-resolution
genomic analyses (sensu Buerkle and Lexer 2008).
Now that the contact zone has been rather precisely
located and some key parameters of variability char-
acterized, we suggest that this geographic, ecological,
and genetic suture zone would be an excellent launch
pad for evolutionary and ecological research on Burbot.

Acknowledgements

This research was funded by the Gwich’in Renew-
able Resource Board, Aurora Research Institute, Envi-
ronment Canada’s Northern Ecosystem Initiative, and
Fisheries and Oceans Canada (A.A.). H.R. was funded
by a Leonardo da Vinci exchange to the University of
Glasgow. K.R.E. thanks R. Eckmann for advice with
analyses. Thanks to Marlene Evans (Environment
Canada) for organizing the stable isotope analyses, and
to Dr. Cameron Goater (University of Lethbridge), Dr.
Terry Dick (University of Manitoba), and Nathan
Millar (Yukon Government) for advice and support
throughout the project. Thanks to all the Gwich’in par-
ticipants who assisted with the project (sample collec-
tion, traditional knowledge, and macro stomach analy-
Sis).

384

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Received 11 February 2014
Accepted 28 March 2014

Observations of the Summer Birds of Tukarak Island (Belcher Islands,

Nunavut), Nastapoka Islands (Nunavut), and Lac Guillaume-Delisle
(Northern Quebec)

CHRISTOPHER W. SWARTH!* and WOUTER BLEEKER2

Natural Reserve System, University of California, Merced, California 95343 USA
“Geological Survey of Canada, Ottawa, Ontario Canada
*Corresponding author: cswarth@ucmerced.edu

Swarth, Christopher W., and Wouter Bleeker. 2014. Observations of the summer birds of Tukarak Island (Belcher Islands,

Slates ), Nastapoka Islands (Nunavut), and Lac Guillaume-Delisle (northern Quebec). Canadian Field-Naturalist 128(4):
S85-972.

In a survey of land and water birds of Tukarak and Nastapoka Islands and Lac Guillaume-Delisle in summer 2011, we recorded
22 of the 30 species known to breed on the Belcher Islands, as well as five non-breeding species, and we observed 32 species in
Lac Guillaume-Delisle. In all areas surveyed, we observed a total of 43 species: 10 waterfowl, 2 gallinaceous birds, 3 loons,
2 hawks, 3 shorebirds, 1 auk, 3 gulls, 2 falcons, and 17 songbirds. In this area of Hudson Bay, a number of species reach the
southern or northern limit of their breeding distribution in eastern Canada. In light of the impact that climate change may have
on bird distribution in northerly latitudes, the Belcher Islands and adjacent mainland areas could be particularly useful loca-

tions for monitoring changes in the breeding range of birds.

Key Words: Breeding birds; Hudson Bay; Belcher Islands; Tukarak Island; Lac Guillaume-Delisle; Nastapoka Islands

Introduction

The breeding avifauna of the Belcher Islands, Nuna-
vut, has been documented by Twomey (1942), Todd
(1963), Freeman (1970), and Manning (1976). Twom-
ey was the first to make systematic bird observations
on the Belcher Islands, which he described in general
terms in Needle to the North (Twomey 1942). On
Tukarak and Flaherty Islands, Twomey collected 19
bird species (273 specimens and 59 clutches of eggs);
his collection is held at the Carnegie Museum of Natu-
ral History, Pittsburgh, Pennsylvania. Freeman (1970)
spent the summers of 1959 and 1960, and March and
April 1961 in the area. Most of his observations were
made in the southwest region of Flaherty Island near
the mouth of the Kasegalik River, although he also
visited Tukarak Island. Manning (1976) visited the is-
lands from May to September 1971, principally to study
polar bears, but he also collected birds and estimated the
size of breeding populations. More recently, information
on Common Eiders was summarized by Abraham and
Finney (1986) and by Robertson and Gilchrist (1998).
Arctic Tern and gull population trends on the Belcher
Islands were described by Gilchrist and Robertson
(1999). A checklist of the bird life of Nunavut (Richards
and White 2008) is based on recent sightings from
throughout this vast territory. A report by the Kativik
Regional Government (KRG 2007) on plans to create
national parks in the western Ungava Peninsula includes
a list of birds of Lac Guillaume-Delisle. The Quebec
Breeding Bird Atlas (www.atlas-oiseaux.qe.ca/index_en
jsp), now in its fifth year, is mapping recent sightings
by birdwatchers to Lac Guillaume-Delisle.

This report summarizes bird observations made from
17 to 30 July 2011 in eastern Hudson Bay, Canada,
specifically on Tukarak Island (Belcher Island group,
Nunavut), on the Nastapoka Islands (Nunavut), and in
Lac Guillaume-Delisle (northern Quebec). Considering
the long periods between visits by ornithologists to these
remote areas, our observations augment the earlier more
lengthy surveys and contribute new knowledge of the
bird life of this region.

Study Area

The Belcher Islands (56°00' to 57°30'N, 79°30" to
80°00’W) comprise 1500 islands and islets covering
almost 3000 km? in eastern Hudson Bay (Figure 1a).
They are located about 120 km off the eastern shore of
Hudson Bay. This region is well north of the tree line.
The only trees we observed on the Belcher Islands were
low-growing willows (Salix spp.) and Paper Birch
(Betula papyrifera Marshall). Also, the area around the
town of Umiujaq on the mainland is mostly treeless.
In contrast, Lac Guillaume-Delisle (Figure 1b) has
extensive groves of spruce (Picea spp.), willow, and
Green Alder (Alnus viridis [Chaix] de Candolle), espe-
cially in ravines and along the lower sections of streams.

Methods

In 2011, we visited Tukarak Island in the Belcher
Islands group (17-24 July), Lac Guillaume-Delisle (25—
30 July), and several of the Nastapoka Islands near
Umiujaq (25 and 30 July). Our primary purpose was to
investigate geological formations in this region; how-
ever, each day we made a concerted effort to identify,
count, and record all the birds in every area we visited.

385

386

Pe!

‘Islands

\)

Belcher
Islands

Hudson Bay

x

5, Laddie °
Split S28 \ isl Y
Isl. 4 i s
0

Isl.
: :
' Wiegand
et Isl.

« Bakers
Dozen Isl.

‘ ot
Px\ :
13)

i- oF 2
SANIKILUAQ*B..”
NIK

- Flat Tukarak Is].
o aN - ian
gf

j S
yas Z
i , 7 x

9S 7 Up

7*Mavor Isl.
OTE

th
4

Innetalling
IsI. ;

THE CANADIAN FIELD-NATURALIST

Vol. 128

58°N

55°N
74°W

:. —Z. Riviere Nastapoka

B | an aa .

spuejs| E40de}SEN

Gillies Isl. |
)

~

Clarke Isl.
Ku

Copper ee

Anderson if }

Riviére a

!'Eau Claire

ki ( &
Le Goulet » 3} o

FiGURE 1. Locations of 2011 fieldwork in eastern Hudson Bay: Tukarak Island (Belcher Island group), Nunavut (a); Lac
Guillaume-Delisle, Quebec; and the Nastapoka Islands, Nunavut (b). HBO = Hudson Bay outpost.

We traveled to and between the islands on the fish-
ing boat, Kakivak, skippered by Inuit fishermen from
Inukjuak, Quebec. While traveling across open water
between shore-based sites, we maintained a constant
watch and tallied all waterbirds visible from the boat.
At night we anchored near shore, which provided excel-
lent views of the rocky shorelines and beaches. Each
day, we covered about 5-10 km on foot across a variety
of habitats: dry and moist tundra, exposed rock forma-
tions, small streams, willow-covered ravines, and lakes,
which we circumnavigated. We observed birds with 8x
binoculars and a 20-60x spotting scope. The figures in
the tables represent the sum total of independent daily
surveys that were often, but not always, made sepa-
rately by the two observers. We took care to avoid com-

bining duplicate counts of the same birds. On the boat,
the observers scanned the open waters from opposite
sides of the boat. C.W.S. devoted about 12 hours each
day to bird observations and W.B. about 6 hours.

Observations on Tukarak Island were concentrated
in three areas: the southeast region; around the old
Hudson Bay outpost along Omarolluk Sound on the
west side; and at Laddie Harbour on the east-central
side of the island (Figure 1a). We did not visit any of
the small coastal islets.

In the Nastapoka Island group, we visited Clarke.
Anderson, and Gillies Islands. In Lac Guillaume-
Delisle, fieldwork took place in Baie Sikutaaluk near
the Gulf (“Le Goulet”), south of the mouth of the

2014

Riviere 4 Eau Claire, and at Copper Cove on the
west-central shore (Figure |b).

Results

On the Belcher Islands, we observed 22 of the 30
species that are regular or occasional breeders there
(Table 1). Breeding species not observed were King
Eider (Somateria spectabilis), Long-tailed Duck (Clan-
gula hyemalis), Semipalmated Sandpiper (Calidris
pusilla), Red-necked Phalarope (Phalaropus lobatus),
Parasitic Jaeger (Stercorarius parasiticus), Arctic Tern
(Sterna paradisaea), Snowy Owl (Bubo scandiacus),
and Common Redpoll (Carduelis flammea). We also
observed five migrant, non-breeding species. The sum-
mer avifauna of the Lac Guillaume-Delisle region, in
contrast, is richer than that of the Belcher Islands. In
the former, we observed 22 species that breed in this
region (including Umiujaq and the Nastapoka Islands)
and 10 migrants (Table 2). In all areas surveyed, we
observed a total of 43 species: 10 waterfowl, 2 galli-
naceous birds, 3 loons, 2 hawks, 3 shorebirds, | auk,
3 gulls, 2 falcons, and 17 songbirds. Several sightings
represent extra-limital records of birds well north of
their regular breeding range. The notes that follow sum-
marize details about these observations.

On Tukarak Island, we observed flocks of 164 and
200 flightless Canada Geese (Branta canadensis),
adults and goslings, and a flock of about 200 on Clarke
Island (Nastapoka Island group). We observed adult
Tundra Swans (Cygnus columbianus) swimming on
lakes on Tukarak Island, but we did not see any cygnets.

We saw Common Eiders (Somateria mollisima seden-
taria) daily in the coastal waters around Tukarak Island
and in Lac Guillaume-Delisle. No eiders were seen on
any inland lakes and we saw no nests. In the waters
around Tukarak Island, over 90% of males were in
breeding plumage, and we observed many females
there with broods of various ages. The first male seen
in eclipse plumage near Tukarak Island was on 21 July,
whereas all the males we identified in Lac Guillaume-
Delisle were in eclipse plumage.

Three male Harlequin Ducks (Histrionicus histrioni-
cus) were seen at Copper Cove, Lac Guillaume-Delisle,
on two days; presumably the same group was seen
each day.

Scoters were seen almost daily. Of those we were able
to identify to species, Surf Scoters (Melanitta perspicil-
lata) were the most abundant. The largest single flock
of 1200 unidentified scoters was seen on Lac Guil-
laume-Delisle on 27 July. Black Scoters (M. ameri-
cana) were seen on Lac Guillaume-Delisle, but not on
Hudson Bay proper. The 30 Red-breasted Mergansers
(Mergus serrator) in Lac Guillaume-Delisle on 27 July
were flightless and in a single flock.

Two female Northern Harriers (Circus cyaneus) were
seen on Lac Guillaume-Delisle on 27 July, and one
female flying with prey was in Umiujaq on 30 July.

Spotted Sandpipers (Actitis maculartus) were seen on
four days along the shores of Lac Guillaume-Delisle.

SWARTH AND BLEEKER: OBSERVATIONS OF SUMMER BIRDS IN NUNAVUT AND QUEBEC

387

An adult was attending three, one-third-grown chicks
on 26 July.

Two Purple Sandpipers (Calidris maritima), the only
ones seen during our trip, flew in and briefly mobbed
us on Tukarak Island on 17 July.

Black Guillemots (Cepphus grylle) were observed
around Tukarak Island and on Lac Guillaume-Delisle
on 10 of 15 days for a mean of 27 per day. The high-
est daily counts were 88 near Tukarak Island and 61
on Lac Guillaume-Delisle. The guillemots were often
in pairs, or in groups of 10 or fewer, and all were in
breeding plumage. No birds were perched on rocks or
cliffs, nor were any carrying food, and we saw no juve-
niles. These observations suggest that they were not
nesting in these areas at this time.

Glaucous Gulls (Larus hyperboreus) were seen on
eight days on Hudson Bay. The largest group was 10
at Laddie Harbour on 22 July, where several half-
grown chicks were seen on rocks near the water’s edge.
No Glaucous Gulls were seen on Lac Guillaume-
Delisle. Great Black-backed Gulls (Larus marinus)
were observed on five days. An adult pair was seen at
Tukarak Island on 18 July and five (four adults and one
third-year bird) were seen near the Nastapoka Islands
on 25 July. We saw nothing to suggest that they had
nests, and we saw no chicks.

Peregrine Falcons (Falco peregrinus) were seen at
two locations on Tukarak Island. On 19 July one flew
along the cliffs at the south end of the island. On 23
and 24 July, we observed a pair closely for several
hours and photographed them at their nest at Laddie
Harbour. The nest site was on a sparsely vegetated
ledge on a vertical cliff, 7-10 m above the water at the
head of a small cove. The nest held two downy young,
which appeared to be several weeks old based on their
size and white downy plumage, and a single egg. At
one point, the male arrived clutching an adult Horned
Lark (Eremophila alpestris).

Single Hermit Thrushes (Catharus guttatus) were
seen and heard singing in a wooded area along the
south shore of Lac Guillaume-Delisle on 25 and 27
July.

Discussion

Many bird species currently reach the northern or
southern limit of their breeding range in the eastern
Hudson Bay region (Godfrey 1986), making this area
especially interesting from a zoogeographic perspec-
tive. However, range boundaries are rarely static and
arctic regions are changing rapidly in response to cli-
mate change (Serrize et al. 2000; Comiso 2003). Bird
populations, especially those at high latitudes, could
undergo significant range shifts in the near future (Jetz
et al. 2007).

Few ornithologists have visited the Belcher Islands,
in part because of their remote location and the asso-
ciated logistics and costs that make access difficult.
Lac Guillaume-Delisle is also remote, although plans
to create a national park there could increase visits in

128

Vol.

THE CANADIAN FIELD-NATURALIST

388

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390)

the near future. Knowledge of the summer bird life of
the Belcher Islands and Lac Guillaume-Delisle is, there-
fore, based on just a few reports and publications that
span many decades. Twomey’s observations, summa-
rized in a popular book (Twomey 1942), provide fas-
cinating details about the bird life, but unfortunately
the results of his extensive study were not published in
the scientific literature. The reports of Thomas Manning
(1946, 1976) were especially thorough, and the sight-
ings by Freeman (1970) are also useful. Based on these
publications, the regular breeding avifauna is consid-
ered to consist of about 30 species. The Quebec Breed-
ing Bird Atlas project (www.atlas-oiseaux.qc.ca/index
_en.jsp) is compiling recent bird records from Lac
Guillaume-Delisle, although there are few mapped
sightings to date from this region. Our observational
records add to the limited number of reports from east-
ern Hudson Bay and, especially, augment knowledge
of the birds that breed or spend the summer around the
Belcher Islands. Conversely, the lack of sightings of
some probable breeding species such as Red-necked
Phalaropes and Common Redpolls does not mean those
species were not present during our visit.

We were not able to determine the age of the flight-
less Canada Geese in the flocks we observed on Tuka-
rak Island and the Nastapoka Islands. Populations in
late spring and summer are likely made up of both local
breeders and of migrants from southern populations.
Based on satellite radio telemetry, it is known that some
geese that breed on the Atlantic flyway in eastern North
America migrate in June to northern Quebec and the
east coast of Hudson Bay (Sheaffer et a/. 2007) where
they undergo a complete annual molt.

Tundra Swans are irregular or uncommon breeders on
the Belcher Islands, which are near the southern limit of
their breeding range (Godfrey 1986). Freeman (1970)
saw only one pair on the Belcher Islands in mid-June
1960, and he cites a report by Burwash (1927) that sug-
gests that swans were scarce in this region by the mid-
1920s. Tundra Swans were common breeders along
southern Hudson Bay in the 1800s, but heavy hunting
greatly reduced their numbers thereafter. Swans did not
begin to re-inhabit these areas until the 1960s (Lumsden
1975).

The sighting of three male Harlequin Ducks was for-
tuitous, but not unexpected as they are known to nest
around Lac Guillaume-Delisle and north to the Nasta-
poka River (Robertson and Goudie 1999; KRG 2007).

The breeding ranges of the Surf Scoter and the Black
Scoter in North America are imperfectly known (Savard
et al. 1998); however, in eastern Canada the centre of
abundance of both species is the Ungava Peninsula.
White-winged Scoters breed to the south and west.
There are no records of scoters breeding on the Belcher
Islands and we saw nothing to suggest that they might
have been nesting during our visit.

The three Northern Harriers observed represent extra-
limital records, well north of their published breeding

THE CANADIAN FIELD-NATURALIST

Vol. 128

range (Godfrey 1986; Smith ef a/ 2011). In the Lac
Guillaume-Delisle region, there is at least one recent
nesting record (KRG 2007) and a single “possible
nesting” record in the Quebec Breeding Bird Atlas. The
female we observed in Umiujaq was carrying prey,
which suggests that a nest may have been nearby.

Purple Sandpipers were first documented as breed-
ing on the Belcher Islands by Twomey (1942). In 1938,
he collected 44 adults and 13 complete clutches (35
specimens and 7 clutches were from Tukarak Island).
Clutch collection dates ranged from 9 June to 25 July
1938 (data courtesy of the Carnegie Museum of Nat-
ural History). Freeman (1970) discovered a nest con-
taining four eggs on 8 July 1959 on an island in the
Kasegalik River. Manning (1976) estimated that there
were 4000 Purple Sandpipers on Kugong Island, the
westernmost of the Belcher Islands. Freeman (1970)
wrote (restated by Jehl 2004) that the Purple Sand-
piper, along with the Semipalmated Plover, is “proba-
bly the commonest [breeding] shorebird” on the Belch-
er Islands. The fact that we saw just two birds suggests
that large numbers may not breed regularly in the areas
of Tukarak Island that we visited. Tukarak is a large is-
land, and the areas we visited may not have been ideal
breeding habitats for this and other shorebird species.
Alternatively, the dates when we visited (mid to late
July) are near the end of the breeding season for many
sandpipers, and birds may have already departed the
island. However, Twomey collected a full clutch on 25
July 1938, and Manning (1976) saw flocks of up to
150 Purple Sandpipers on Split Island (about 70 km to
the northwest of Tukarak Island) in early August 1973,
and he collected a male with four downy young there
on 3 August. A thorough survey of all the Belcher Is-
lands would be required to develop a clear understand-
ing of their current breeding status. The Semipalmated
Plover, Semipalmated Sandpiper, and Red-necked Pha-
larope also nest on the Belcher Islands (Todd 1963:
Godfrey 1986); however, we observed only the plovers.

The single summer record for Least Sandpiper (Cali-
dris minutilla) on the Belcher Islands is an observation
of one nest by Twomey (1942) on Tukarak Island. Even
though we did not observe any Least Sandpipers and
Godfrey (1986) indicates that they do not nest on the
Belcher Islands, further fieldwork could reveal that
they are occasional breeders, as they are known to
nest around Lac Guillaume-Delisle (Todd 1963: KRG
2007).

The sightings of adult Great Black-backed Gulls
were noteworthy because known breeding colonies are
quite distant: at Akimiski Strait, James Bay, 350 km
south of the Belcher Islands (Eckert 2007: Dunn and
Alderfer 2011) and 1400 km (by the coastline) north-
east at the mouth of the Riviére aux Feuilles in Ungava
Bay (Good 1998). The Belcher Islands could be col-
onized in the near future as nesting at Akimiski Strait
was documented for the first time in 2007 (Eckert
2007). The Great Black-backed Gull is undergoing rap-

2014

id range expansion throughout eastern North Ameri-
ca (Good 1998),

We did not observe Arctic Terns. Arctic Tern popula-
tions declined between the 1980s and 1997 in the three
northern regions (Sleeper, Split, and Laddie Islands) of
the Belcher Islands (Gilchrist and Robertson 1999),
Only 19 of the 431 islets in that region had nesting terns
in 1997. This decline is attributed either to winter mor-
tality or to emigration out of the Belcher Islands as a
response to egging or disturbance by residents of Sani-
kiluaq, the only village on the islands. The tern surveys,
summarized by Gilchrist and Robertson (1999), were
not made on Tukarak Island, presumably because Arc-
tic Terns are not known to nest on this island.

The nesting Peregrine Falcons we observed at Lad-
die Harbour belonged to the tundra subspecies, P. f-
tundrius, based on the thin malar stripe, white upper
breast, and extensive white in the auricular area. Free-
man (1970) assigned the Peregrine Falcons he observed
on Tukarak Island to Pf. anatum, although he pro-
vided no evidence to support this determination. The
Nastapoka River represents a dividing line between
Pf. tundrius on the north and Pf anatum to the south
(Murphy 1990).

The Hermit Thrushes seen in the Lac Guillaume-
Delisle were extra-limital records, over 450 km north
of the northern limit of their breeding range in south-
western Quebec (Godfrey 1986; Dellinger et al. 2012).
Yves Aubrey (in KRG 2007) reports that they do nest
in this area.

The Belcher Islands are the southernmost breeding
site in North America for Snow Buntings (Plectro-
phenax nivalis), whereas the Yellow Warbler (Setopha-
ga petechia) and White-throated Sparrow (Zonotrichia
albicollis) are at the northern limits of their range at
Lac Guillaume-Delisle. Sightings of these three species
in the regions we visited are consistent with their known
occurrence in eastern Canada.

In summary, the Belcher Islands are large, we only
visited parts of Tukarak Island, and our fieldwork was
brief — factors that make our survey less than com-
plete. Clearly fieldwork in more areas and for longer
periods is required to determine the current status of
all breeding and summering birds there. The strategic
location of the Belcher Islands and Lac Guillaume-
Delisle — at the tree line and in a climatic zone where
arctic conditions exist, as well as the fact that they sup-
port a number of species at the limits of their geograph-
ic range — make them useful sites for studying shifts
in species’ breeding ranges that could result from glob-
al climate change.

Acknowledgements

Our expedition was made possible by grants to Dom-
inic Papineau and Marilyn Fogel from the Carnegie
Institution of Canada, the National Aeronautical and
Space Administration Astrobiology Institute, the W. M.
Keck Foundation, and the Geophysical Laboratory of

SWARTH AND BLEEKER: OBSERVATIONS OF SUMMER BIRDS IN NUNAVUT AND QUEBEC

39]

the Carnegie Institution of Washington. The authors and
scientific team thank the municipality of Umiujaq, the
Sakkuq Landholding Corporation, the Makivik Cor-
poration, the Nunavut Research Institute, the Govern-
ment of Nunavut (Department of Culture, Language,
Elders and Youth), and the Qikigtani Inuit Association
for permission to conduct fieldwork on their territory.
We are grateful to the captain and crew of the Kakivak
for expert seamanship and, especially, to Arthur Eli-

jassiapik for cheerful assistance in the field. Stephen

Rogers, bird collection manager at the Carnegie Muse-
um of Natural History, provided data on bird and egg
specimens from the 1938 Twomey expedition. Jason
Chartrand (GSC, Ottawa) provided GIS assistance in
drafting Figure 1. David Shuford, Dominic Papineau
and two anonymous reviewers made constructive com-
ments on an early draft.

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Scribner. 2007. Management implications of molt migra-
tion by the Atlantic Flyway resident population of Cana-
da Geese, Branta canadensis. Canadian Field-Naturalist
121(3): 313-320.

Smith, K. G., S. R. Wittenberg, R. B. Macwhirter, and K.
L. Bildstein. 2011. Northern Harrier (Circus cyaneus). In
The Birds of North America Online. Edited by A. Poole.
Cornell Lab of Ornithology, Ithaca, New York, USA.
Accessed 20 September 2014. http://bna.birds.cornell.edu
/bna/species/2 1 0/

Todd, W. E. C. 1963. Birds of Labrador and Adjacent Areas.
University of Toronto Press (with the Carnegie Museum),
Toronto, Ontario, Canada. 819 pages.

Twomey, A. C. 1942. Needle to the North: The Story of an
Expedition to Ungava and the Belcher Islands. Houghton
Mifflin, Boston, Massachusetts, USA.

Received 15 March 2014
Accepted 1 May 2014

Comparing the Diet of Great Horned Owls (Bubo virginianus) in Rural
and Urban Areas of Southwestern British Columbia

SOFI HINDMARCH!:3 and JOHN E. ELLIOTT?

Langley, British Columbia, Canada
Environment Canada, Science and Technology Branch, Delta, British Columbia V4K 3N2 Canada
Corresponding author: sofi.hindmarch@gmail.com

Hindmarch, Sofi, and John E. Elhott. 2014. Comparing the diet of Great Horned Owls (Bubo virginianus) in rural and urban areas
of southwestern British Columbia. Canadian Field-Naturalist 128(4): 393-399.

We investigated the diet of Great Horned Owls (Bubo virginianus) in southwestern British Columbia. Our objective was to
compare the diets of owls in urban and rural areas and determine whether urban owls consume a higher proportion of com-
mensal rodents to understand possible pathways of secondary rodenticide poisoning of Great Horned Owls. Among 546 prey
items identified at seven sites, Townsend’s Vole (Microtus townsendii [Bachman, 1839]) and rats (Rattus G. Fischer, 1803)
were the two main prey items, making up 65.9% and 13.1% of the diet, respectively. The proportion of rats in the diet was
positively correlated with the degree of urban development in the owls’ home range Vs 0.83, P< 0.05, df=5).

Key Words: Great Horned Owl; Bubo virginianus; diet; British Columbia; voles; Townsend’s Vole; Microtus townsendii, rats;

Rattus; rodenticide

Introduction

The Great Horned Owl (Bubo virginianus) is found
throughout North America, except in the Arctic, mak-
ing it the most common and widespread owl on the
continent (Houston et al. 1998*). The large and diverse
range of the Great Horned Owl is also reflected in its
diet. As a generalist and opportunistic feeder, it con-
sumes a wide range of prey species, including lago-
morphs, rodents, waterfowl, game birds, raptors, insects,
and even larger birds, such as herons (Houston et al.
1998*; Johnsgard 2002). Although a diverse array of
prey species have been identified, over most of the
Great Horned Owl’s range, its diet consists of 90% mam-
mals, predominantly rodents, and 10% birds (Houston
et al. 1998*; Johnsgard 2002).

Despite several studies and reviews investigating
the food habits of the Great Horned Owl in a variety
of regions of North America (Houston et al. 1998*;
Johnsgard 2001), no Great Horned Owl diet studies
have been conducted in southwestern British Columbia,
and only one such study has been reported from the
province (Van Damme 2005). Most previous studies
have been conducted in forest, grassland, and agricul-
tural settings; few have investigated the diet of the Great
Horned Owl in more urban landscapes. Like other preda-
tory birds, such as the Sharp-shinned Hawk (Accipiter
striatus) and the Northern Goshawk (Accipiter gentilis),
Great Horned Owls are increasingly finding niches in
urban ecosystems across North America (Lambert 1981;
Powers 1996; Rutz 2008) where their exposure to envi-
ronmental contaminants may be enhanced, especially
through consuming rodents that have previously ingest-
ed anticoagulant rodenticides. The risk of secondary
exposure to these rodenticides in raptors is currently

receiving increased attention in Canada (Albert et al.
2010; Thomas et al. 2011; Elliott et a/. 2014).

Our study had two objectives: to obtain dietary infor-
mation for Great Horned Owls in a region where no
previous data had been collected and to compare the
diet of owls that inhabit agricultural versus suburban
landscapes in southwestern British Columbia. Such data
are of particular interest in rapidly urbanizing regions,
such as southwestern British Columbia, where the loss
of forests and agricultural lands surrounding urban cen-
tres forces owls and other wildlife into the remaining
patches of green space, such as parks, suburban wood-
lots, and fragments of undeveloped land. We were
specifically interested in evaluating whether the amount
of urban development within Great Horned Owl home
ranges influenced the consumption of commensal ro-
dents such as Norway Rats (Rattus norwegicus |Berken-
hout 1769]), Black Rats (Rattus rattus [L., 1758]), and
House Mice (Mus musculus L., 1758), which are the
species commonly targeted for control with anticoag-
ulant rodenticides.

Study Area

Surveys for Great Horned Owl nest and roost sites
were conducted from December 2010 to December
2013 in the municipalities of Richmond, Vancouver,
Burnaby, Delta, Surrey, and New Westminster (a total
area of 847 km?) in southwestern British Columbia,
Canada (49°8'N, 122°18'W). The area includes some
of the main stopover sites for birds migrating on the
Pacific flyway and encompasses important wildlife
areas such as the Alaksen National Wildlife Area, Burns
Bog, Stanley Park, Pacific Spirit Regional Park and
Boundary Bay Regional Park. Before European settle-
ment, the low-lying floodplains were dominated by
grassland, low shrub vegetation, and extensive stands of

395

mixed woodlands dominated by cottonwood, alder, and
cedar, while higher elevations were covered primari-
ly by mixed coniferous forest (North and Teversham
1984). Today, the landscape ranges from agricultural
to suburban to highly urban, and the remaining lower
grassland and forested habitats face ongoing develop-
ment pressure as the projected human population in the
region is expected to increase from the current 2.4 mil-
lion to 3.4 million in 2040 (Metro Vancouver 2009*).

Methods
Owl surveys, pellet collection and analyses

Suitable areas to survey for nest and roost sites and,
thus, pellet and prey remains were identified with the
collaboration of local natural history clubs and by sur-
veying parks and green spaces for signs of owl pres-
ence, such as whitewash, feathers, and pellets. To con-
firm the presence of Great Horned Owls and to guide
pellet searching in larger parks, call play-back was used
at dusk following the North American nocturnal owl
survey guidelines (Takats et al. 2001*). When we found
evidence of an owl roosting or nesting, we revisited the
site every 2~3 months to search for additional pellets
and prey remains. Although the potential for occasional
loss of larger prey remains to scavenging existed (Mar-
ti et al. 2007), we believe there would likely be suffi-
cient remains of such large prey items to allow iden-
tification. Incidental observations of foraging and
prey type were also recorded.

Pellet analysis

We dissected pellets carefully to ensure that prey
items could be identified from bone remnants, fur and
other body parts using British Columbia small mam-
mal field guides (Nagorsen 1996, 2005). We deter-
mined the number of individuals of any species in each
pellet by pairing each skull with the correct number
of ischia, left and right mandibles, and tibiae/fibulae
or, in the case of birds, each skull with sternum, giz-
zard sac, and feet. We assembled the remaining bones
in the pellet to determine the minimum number of addi-
tional individuals whose skull may have been crushed.
For smaller prey items (< 100 g), we assumed that the
remains of each were contained in a single pellet, as
finding bones from one prey item in two successive pel-
lets is rare (Raczynski and Ruprecht 1974). We estimat-
ed the weights of rats from intact jaw bones in the prey
remains following Morris (1973).

Very few rat (Norway Rat or Black Rat) and shrew
(Vagrant Shrew (Sorex vagrans) and Montane shrew
(Sorex monticolus) prey remains were sufficiently intact
to determine species) and, hence, we pooled all rats
and shrews into one category. For the same reason, we
allocated songbird (Passeriformes) prey remains to
two categories: small songbirds (< 30 g) and medium
songbirds (30-80 g). We considered all insect exoskele-
ton remains to belong to the order Coleoptera.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Evaluation of land use within home ranges

In examining differences in the diet of Great Horned
Owls between agricultural and suburban sites, we quan-
tified the amount of urban land (residential, industri-
al, and transportation) within a 1-km radius (3 km? or
300 ha) of each nest or roost site from digitized data
layers using geographic information system software
(ArcMap 10, Esri, Redlands, California, USA). We
used a 1-km radius, as the average home range of the
Great Horned Owl is approximately 3 km* (Petersen
1979*: Houston et al. 1998*). Data on land use within
the home ranges were obtained from a 2006 Vancouver
Regional District land-use layer map that categorized
land parcels based on zoning (Metro Vancouver 2008*).
We compared the 2006 land-use layer map with 2010
Bing Ortho photos (Bing Maps, Microsoft, Redmond,
Washington, USA) to control for any recent changes
in land use or discrepancies between current land use
and zoning.

We conducted a Pearson correlation analysis to deter-
mine if there was any relationship between the amount
of urban development within home ranges and the
proportion of rats in the diet of Great Horned Owls.
The Pearson correlation analysis was carried out using
IBM SPSS 22 (IBM Inc. Armonk, New York, USA).

Results

We found seven sites (five nests and two roosts) oc-
cupied by Great Horned Owls in our study area and
monitored them regularly. Three sites were located in
predominantly agricultural landscapes (Alaksen Nation-
al Wildlife Area, Forest Richmond, and Arthur Drive)
and four in urban parks and green spaces (Terra Nova
Park, Beach Grove Park, Crescent Park, and Central
Surrey). The proportion of urban development within
Great Horned Owl home ranges (3 km?) varied consid-
erably, from 0%-—7.6% at rural sites to 33.8—93.3% at
urban ones (Table 1).

Pellets and prey remains were found predominant-
ly under nest or roost trees. Some pellets were weath-
ered considerably, and it was impossible to get an exact
count of the number of pellets collected, but, in total,
546 prey items of 21 species were identified. Overall.
Townsend’s Vole (Microtus townsendii [Bachman,
1839]) was the most common (65.9%), and was the
dominant prey species at six of the seven sites. fol-
lowed by Rattus spp. (13.1%). Other species were only
marginally represented in the diet, each contributing
less than 5% of the total number of individuals overall
(Table 1). On average 7.1 prey species were identified
at each site (range 2—9), and the intact prey ranged in
weight from about 1 g (e.g., beetle) to over 2 kg (Great
Blue Heron, Ardea herodias).

In the late summer and fall of 2012, one Great
Horned Owl pair residing in Terra Nova Park. Rich-
mond, was observed by park employees preying on
Barn Owls (Zvto alba) (Figure 1). In total, eight Barn

395

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396

THE CANADIAN FIELD-NATURALIST

Vol. 128

www.sharingfarm.ca

FIGURE |. Great Horned Owl (Bubo virginianus) sitting on shed roof with a captured Barn Owl (Tyto alba). The Great Horned
Owl pair at Terra Nova Park, Richmond, caught a total of eight Barn Owls in fall 2012. Photographed 10 October 2012

at Terra Nova Park Richmond; photo by Sharing Farm.

Owls were preyed on, most probably young of the year
as band recovery confirmed that two were recently
fledged chicks from nest sites 14 and 34 km away.
Eastern Cottontails (Sy/vilagus floridanus [J. A. Allen,
1890]) were not an important component of the diet
and were recorded at only three urban sites. However,
at one nest site located in a park, they were the second
most consumed prey along with rats at 23.5%. Further,

12

10

Number
Lo>)

20 40 60 80 100

only one House Mouse (Mus musculus L., 1766) and
one Eastern Gray Squirrel (Sciurus carolinensis Gmelin,
1788) were found among all prey remains.

Following Morris’ (1973) model for predicting the
body weight of rat prey items, the average mass of rats
consumed by Great Horned Owls was 118 + 63.3 g
(range 20-280 g, n = 39). Based on the mode (Figure
2), Great Horned Owls most frequently consumed rats

140 160 180 220 260 280

Rattus Sp. Mass (g)

FIGURE 2. Number of rats (Rattus spp.) by weight class consumed by Great Horned Owls (Bubo virginianus )
in southwestern British Columbia, Canada (average = 118 + 63.3 g)

at seven sites

2014 HINDMARCH AND ELLIOTT: THE Diet OF GREAT HORNED OWLS IN URBAN AND RURAL BC

weighing 80 g. The difference between the average and
the mode is explained by the Great Horned Owls’ capa-
bility of capturing rats of all sizes, including the occa-
sional larger rat (> 200 g). The proportion of rats in the
diet was significantly correlated with the amount of
iy land within home ranges (7, = 0.83, P < 0.05,
df= 5).

Discussion

Despite the highly flexible foraging behaviour of

Great Horned Owls, they tend to focus on only one or
two profitable prey species within a geographic region
(Houston et al. 1998*; Marchesi et al. 2002). The food
habits of the Great Horned Owls within our study area
showed no exception to this trend. Overall, Townsend’s
Voles dominated the diet followed by rats, and a diverse
range of other species each accounted for less than 5%
of the prey consumed at all sites combined. Land use
surrounding individual nest and roost sites likely influ-
enced the diversity and abundance of available prey
species and, ultimately, the diet of Great Horned Owl
pairs. This was evident from the consumption of rats,
which increased as home ranges became gradually
more urban.

The move toward increased rat consumption in more
urban environments was previously documented in
Great Horned Owls nesting in city parks in Seattle,
Washington (Lambert 1981). Similarly, the larger cousin
of the Great Horned Owl, the Eurasian Eagle Owl (Bubo
bubo) consumed more rats when nesting in European
urbanized landscapes (Marchesi et al. 2002; Sandor
and Ionesco 2009), while the diet of a pair of Desert
Eagle Owls (Bubo ascalaphus) in the city of Hurghada,
Egypt, was made up of 71.8% House Mice and Norway
Rats (Sandor and Moldovan 2010). In South Korea,
Eurasian Eagle Owls consumed rats predominantly in
both urban and agricultural landscapes (Shin ef al.
2013). In all cases, the increased consumption of rats
was attributed to their status as an abundant, stable,
year-round food source. In our study, Great Horned
Owls consumed predominantly smaller rats, which are
likely to be more abundant, less risky to handle, and
faster to process than larger rats.

Among other commensal rodents, only one House
Mouse was found. Similarly, only one Eastern Gray
Squirrel was recorded, at a nest in an urban park, des-
pite the abundance of squirrels in all urban and rural
parks, at least partly a result of public feeding (S. Hind-
march personal observation).

Second-generation anticoagulant rodenticides
(SGARs) used to suppress commensal rodent popu-
lations worldwide, have been shown to be persistent,
highly bioaccumulative, and very toxic to non-target
species (Parmar et al. 1987; US EPA 2004*). Among
raptors, Great Horned Owls have one of the highest
rates of exposure to, and toxicity from these rodenti-
cides (Stone et al. 1999; 2003; Albert et al. 2010;
Murray 2011, Thomas et al. 2011). Further, predators

397

may be at a greater risk of SGAR exposure in urban
settings where larger quantities of these compounds
are used to suppress commensal rodent populations
(Stone et al. 1999, 2003; Riley et al. 2007; McMillin
et al. 2008).

In southwestern British Columbia, 70% (1 = 61) of
Great Horned Owl carcasses tested between 1988 and
2003 contained one or more SGARs (Albert e7 al. 2010).
More recently, all of the Great Horned Owls collected
in southwestern British Columbia between 2005 and
2011 (n = 29) tested positive for one or more SGARs
(J.E.E. unpublished data). There is some evidence to
suggest that non-target species including native mice
and voles, squirrels, and passerines enter SGAR bait
stations and feed (US EPA 2004*; Brakes and Smith
2005; Tosh et al. 2012), although rats are likely one
of the main vectors responsible for secondary expo-
sure of non-target predators (Cox and Smith 1990;
Birks 1998; Elliott et a/. 2014). Our data showing in-
creased consumption of rats by urban Great Horned
Owls are consistent with the idea that rats are the main
source of exposure to SGARs. However, the dominance
of Townsend’s Voles in the diet of Great Horned Owls
suggests the need for SGAR residue sampling in non-
target small mammals in addition to rats in urban envi-
ronments.

Our land-use analysis may not have been able to
identify fine-scale landscape differences between sites.
For example, one Great Horned Owl pair nesting in an
urban nature park (26 ha) consumed predominantly
Townsend’s Voles (70.3%), and only 2 rats were found
in the pellets. Although that park is surrounded by res-
idential development, over recent years an old field
restoration project has resulted in the removal of inva-
sive plant species and an increase in the abundance of
Townsend’s Voles, with the expressed goal of encour-
aging the nesting of raptors. This was also the site
where the Great Horned Owl pair consumed eight Barn
Owls and one Great Blue Heron in the fall of 2012.
Great Horned Ow] predation on Barn Owls has been
documented previously (Rudolph 1978, Knight and
Jackman 1984, Millsap and Millsap 1987, and Van
Damme 2005). In this case, on several occasions we
observed a Great Horned Owl entering a Barn Owl
nest box to feed. Such behaviour can be prevented by
reducing the size of the entry hole on Barn Owl nest
boxes and installing a vertical predator guard on the
inside. The size of the entry hole was reduced on all
the boxes in the park shortly after these observations
were made.

Our diet data revealed that Great Horned Owls in
southwestern British Columbia feed primarily on voles
and rats. Despite our small sample size, the consump-
tion of rats was significantly higher among Great
Horned Owls with a higher proportion of urban land
within their home range. The increased consumption
of rats and the negligible number of House Mice and
squirrels in the diet indicate that rats could be a major

398

pathway for secondary SGAR exposure in Great Horned
Owls.

Acknowledgements

We thank Paul Levesque and Alexa and Jamie Coote
for their great help in looking for owl pellets. In many
instances, their efforts included crawling around in the
rough understory of urban parks. We appreciate and
thank Sean McCann’s efforts in proofreading and pro-
viding constructive comments on an earlier draft of
this manuscript. Also, we wish to thank the two inde-
pendent reviewers for their knowledgeable insights
and efforts in reviewing our manuscript. Funding for
this research was provided to John Elliott by the Pes-
ticide Science Fund of Environment Canada.

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Received 17 March 2014
Accepted 28 June 2014

Grouseberry (Vaccinium scoparium Leiberg ex Coville) Fruit Produc-
tion in Forest Openings in Banff National Park, Alberta

IAN PENGELLY! and DAviID HAMER”?

'235 Lady MacDonald Drive, Canmore, Alberta T1W 1H2 Canada
2300-379 Spring Creek Drive, Canmore, Alberta T1 W 0G8 Canada
3Corresponding author: j.david.hamer@gmail.com

Pengelly, lan, and David Hamer. 2014. Grouseberry (Vaccinium scoparium Leiberg ex Coville) fruit production in forest openings
in Banff National Park, Alberta. Canadian Field-Naturalist 128(4): 400-407.

Grouseberry (Vaccinium scoparium Leiberg ex Coville) is an abundant dwarf shrub in the understory of many areas of sub-
alpine forest in Banff National Park and has the potential to produce fruit important for wildlife. However, the suppression
and prevention of wildfires, which began in the early 1900s in the park, have reduced fire-dependent shrubland and open forest
and increased the extent of closed, mature forest. Because canopy closure is typically associated with decreased fruit production
by understory shrubs, the decline in fire disturbance may be reducing Grouseberry fruit production. To quantify this effect, we
measured Grouseberry fruit production under various forest canopies at 10 sites in Banff during 2004-2012. We measured site
openness by modeling photosynthetically active direct solar radiation (dPAR) adjusted for overshadowing by topography and
coniferous foliage. We found a positive relation between Grouseberry fruit production and dPAR in 2006 and 2010-2012,
but not in 2008 or 2009, the 2 years of lowest fruit production; data were lacking for 2004, 2005, and 2007. We also recorded
high Grouseberry fruit densities beginning 5 years after fire removed the forest canopy in four prescribed burns conducted
during 2001: fruit production was 3.3 to more than 20 times that in adjacent mature forests in 5 of the 6 years analyzed. This
study shows the potential ecological benefits of both prescribed burns and wildfire in upper subalpine forests where Grouseberry
is widespread, but fruit production is low under the forest canopy.

Key Words: Banff National Park; fruit production; Grouseberry; prescribed burn; photosynthetically active radiation; solar

radiation; Vaccinium scoparium: wildfire

Introduction

Grouseberry (Vaccinium scoparium Leiberg ex Cov-
ille) is a dwarf shrub 10-30 cm in height that produces
a small, reddish fruit 4-6 mm in diameter (Szczawinski
1962). These fruits are eaten by a variety of birds and
mammals (Hamer and Herrero 1987; De Franceschi and
Boag 1991; Mattson 1997). Grouseberry is abundant in
the understory of many forest communities of Banff
National Park, where Grouseberry forest community
types cover 28% of a 123-km? mapped area in the Front
Ranges (Hamer 1985).

Fruit production in many Vaccinium species is re-
duced, sometimes dramatically, under a forest canopy
(Hamer and Herrero 1987; Noyce and Coy 1990; Jor-
dano 2000; Greenberg et a/. 2007). Interception of solar
radiation by the forest canopy decreases the amount of
photosynthetic energy available to understory plants for
production of seed and fruit (Kudo et al. 2005). Martin
(1983) recorded the production of less than 134 L/ha
of Mountain Huckleberry (V. membranaceum Douglas
ex Torrey) fruit in Montana where canopy cover was
greater than 30%; in more open sites, up to 1400 L/ha
of fruit were noted. Weaver et al. (1990) also found that
the abundance of Huckleberry fruit had a strong neg-
ative relation to canopy cover (7? = 0.96). The impor-
tance of light was inferred in a Huckleberry study in
northwest British Columbia, where fruit production was
roughly 50% less in transects where the forest canopy
blocked more than 40% of incoming solar radiation

compared to transects in sites with greater solar radi-
ation (Burton 1998*).

Before the 20th century, fire was the dominant eco-
logical process affecting forests in Banff National Park.
However, since the early 1900s, the incidence of wild-
fires has declined to a small fraction of the previous
rate because of fire prevention and suppression (White
1985). We hypothesized that without wildfires, the ex-
tent of post-fire shrubland and open-canopy forest is
reduced, causing a decline in Grouseberry fruit produc-
tion. During 2004-2012, we studied the relation be-
tween site openness and Grouseberry fruit production
in Banff National Park.

Study Area

Our study area was in the upper subalpine zone of
Banff National Park (Figure 1). Annual precipitation
was estimated at 50-125 cm (Janz and Storr LOFT):
Warm, drying Chinook winds descend the east slopes
of the Rocky Mountains and reduce moisture and snow
pack, particularly on slopes with a south or west aspect.

Elevations in the park range from 1330 to 3610 m
with the treeline at roughly 2300 m. The subalpine zone
is at approximately 1500-2350 m. The upper subalpine
area, which is generally cooler and wetter, with deep-
er and longer-lasting snow, begins at roughly 2000 m
(Achuff 1982*). Our study sites were at an elevation
of 2080-2380 m in the upper subalpine zone where
Grouseberry is abundant.

400

2014

7 y / \
_ | Banffl °
‘National

\\ - Park (

‘he ra
rasa:
By National \

é ~~ \ Park
“ .) ( :
ite

Grouseberry Sample Sites
a Front Range
* Main Range
A Prescribed Burn

oa |

PENGELLY AND HAMER: GROUSEBERRY FRUIT PRODUCTION IN BANFI

4()|

10 Kilometres

FicuRE 1. Location of Grouseberry study sites in Banff National Park, Alberta, 2004-2012.

The forests in our study area were dominated by
Engelmann Spruce (Picea engelmannii Parry ex Engel-
mann). In some sites, Lodgepole Pine (Pinus contorta
Douglas ex Loudon), Subalpine Fir (Abies lasiocarpa
[Hooker] Nuttall), and Subalpine Larch (Larix lyallti
Parlatore) were also abundant. The upper subalpine
forest is opened by a variety of factors including fire,
avalanches, semi-xeric conditions on exposed south-
west-facing slopes, which can lead to grassland and
shrubland, and cold air pooling in depressions at the
toe of slopes (Hamer 1996).

Because of our sampling design, the understory of
most sites was dominated by Grouseberry. Other fre-

quent understory species included Heart-leaved Arni-
ca (Arnica cordifolia Hooker), Fireweed (Epilobium
angustifolium L.), Bracted Lousewort (Pedicularis
bracteosa Bentham), Pink Mountain Heather (Phyi-
lodoce empetriformis [Smith] D. Don), Arctic Willow
(Salix arctica Pallas), Low Blueberry (Vaccinium myr-
tillus L.), Sitka Valerian (Valeriana sitchensis Bon-
gard), and grasses. Regenerating Engelmann Spruce
and Subalpine Fir were frequent tall shrubs; willows
(Salix spp.) occurred in fewer than 10% of the tran-
sects.

Methods
Site selection

We established 10 study sites in total: eight in the
Front Ranges and two in the Main Ranges (Figure 1).
We located one Front Range site in an 1889 burn and
a Main Range site in a 1920 burn. These large wild-
fires had burned into the upper subalpine and created
shrubland and regenerating forest with varying degrees
of canopy closure, including communities where
Grouseberry was a dominant shrub (Hamer and Her-
rero 1987). Four sites were 6-10 km from the 1889 and
1920 burn sites, where we found relatively accessible
terrain, communities with Grouseberry dominant in
the shrub layer, and forest openings in stands originat-
ing from wildfires that had occurred between 1850
and 1900. Another four sites were located where pre-
scribed burns had been conducted from mid-Septem-
ber through mid-October 2001. These fires removed the
forest canopy and scorched the ground layer; Vaccini-
um shrubs had subsequently resprouted from under-
ground rootstocks. The forests in this area were multi-
aged, originating from fires that occurred from about
1800 to 1868 (Hamer and Herrero 1987).

Transect selection

We established 11 pairs of transects in the four pre-
scribed burn sites and adjacent forests. Because the
fires had run upslope from ignition points, we were
able to establish pairs of burned and unburned transects
where ignition, rather than habitat characteristics, was
responsible for which areas burned.

Transects in the burned areas were at 60—125 m inter-
vals (mean 90 m). Because Vaccinium cover was patchy
following fire, we adjusted the transect locations by up
to a few tens of metres from the systematically located
points to ensure relatively high Vaccinium cover (mean
41%, standard deviation [SD] 18%). We located con-
trol transects in the unburned adjacent forest along
the same elevational contour as the burned transects,
with similar adjustment to locate transects in commu-
nities where Vaccinium was dominant in the shrub layer
(mean 62%, SD 16%). We established five transect
pairs in the largest burn, two pairs in a burn 1.2 km
north, and two pairs in a burn 1.1 km south of the lar-
gest burn, all on westerly slopes of 215—271°. Two pairs
of transects were located in a fourth burn, on an east-
erly slope of 84-124°.

The six additional sites in the study area were also
established where Vaccinium was dominant in the shrub
layer (mean cover 57%, SD 15%). At each site, we lo-
cated one or more transects under a forest canopy cover
of less than 40%. When the terrain and Grouseberry
cover were suitable, we established transects with no-
tably different slope aspects and forest cover.

Transect analysis

The ends of each transect were marked with per-
manent metal bars. We recorded transect location and
elevation using a handheld geographic positioning sys-

THE CANADIAN FIELD-NATURALIST

Vol. 128

tem unit and slope steepness and aspect using a com-
pass with built-in clinometer. In the year when transects
were established, we estimated Vaccinium cover and
height at 1-m intervals on the 20-m transects (1.e., 20
measurements/transect). We estimated cover visually
to the nearest 5% using a 10 cm by 10 cm frame and
measured height as the length of the longest Vaccinium
shoot rooted within 5 cm of the Vaccinium stem rooted
nearest to a preselected corner of the frame.

We estimated Grouseberry fruit density by counting
all fruits within a 20 cm by 20 cm frame placed at 2—m
intervals along the 20—m transect (10 quadrats/transect).
At each 2—m point, a 180° forward-facing selection
zone (Hamer 1996) was scanned and the frame was
placed in the first available location with more than
70% Vaccinium cover (lower cover was accepted if the
alternative meant moving the frame more than 2 m
from the original 2-m point). Because of these sub-
jective criteria, quadrats were not in exactly the same
location each year. Fruits were picked to ensure that
each was counted only once. Vaccinium cover in the
quadrat was estimated to the nearest 5%. Counts were
then scaled to 100% Vaccinium cover (e.g., a quadrat
with 35 fruits and 50% Vaccinium cover scored 70
fruits) to allow comparison among locations with vary-
ing Vaccinium cover. In other words, we calculated
fruits/m* of Vaccinium shrub cover, not fruits/m2 of
habitat. Because fewer than 1% of the fruits we count-
ed in 2004-2012 were Low Blueberry, we refer to fruits
as Grouseberry. Because Grouseberry and Low Blue-
berry can be difficult to separate based on vegetative
characteristics (Szczawinski 1962, Vander Kloet and
Dickinson 1999), we report cover and height for Vac-
cinium species.

Estimating solar radiation

Although forest canopy is a primary cause of shad-
ing, nearby or tall mountains also block solar radiation.
Thus, we measured site openness by modeling direct
photosynthetically active radiation (dPAR) adjusted
for overshadowing by both topography and coniferous
canopy.

We calculated the relative amount of dPAR (band
1, wavelength 290-700 nm) received at our transects
under cloudless conditions by applying the REST2
model (Gueymard 2008; Gueymard and Myers 2008),
using site-specific latitude, slope aspect, slope steep-
ness, and elevation. We calculated radiation at 1-minute
intervals from sunrise to sunset and summed these val-
ues for 1 June to 31 August. We chose this 92-day peri-
od because we did not have an a priori prediction of the
critical time for Grouseberry flower and fruit devel-
opment and because the length of the growing season
varies from year to year depending on weather and snow
melt and among sites depending on microclimate.

We imposed an overshadowing function that set
dPAR to zero when the height of obstacles on the solar
azimuth blocked the sun by exceeding the solar alti-
tude (Quaschning and Hanitsch 1998: Yard et al. 2005).

2014

We used a clinometer (Suunto, Vantaa, Finland) and
a Ranger sighting compass (Silva, Bromma, Sweden)
to record the height (degrees of elevation above hori-
zon) and horizontal sweep (compass bearings) of obsta-
cles, whether geological features or fully closed canopy
of coniferous foliage, for azimuths 50° through 310°
(1.e., sunrise to sunset at the summer solstice). We
recorded these data for each relatively homogeneous
block of foliage or terrain; blocks ranged from 1° to
several tens of degrees of horizontal sweep (block
average 7.2°).

We imposed a second overshadowing function on
the model to partly restrict dPAR because of the par-
tial shade resulting from open coniferous canopy. We
recorded the angular height and horizontal sweep, plus
the average canopy openness in 5% increments from
5% for essentially closed canopy (sky almost com-
pletely obstructed; 5% of dPAR transmitted) to 95%
for essentially no foliage (sky almost fully visible; 95%
of dPAR transmitted), for each relatively homogeneous
block of coniferous foliage (block average 18.2°), from
azimuths 50° through 310°. Because these readings
required subjective estimation of coniferous canopy
density, all data were recorded by the same observer
to avoid inter-observer variability.

These two procedures for estimating overshadowing
often captured coniferous foliage up to 200 m away
and, hence, were not equivalent to the “fish-eye” lens
photographic method often employed in forest-gap
studies (e.g., Englund et al. 2000).

We calculated dPAR at the 7-m and 13-m marks on
our 20-m transects and averaged these to obtain one
value per transect. Because comparisons among tran-
sects were relative in our analyses, we did not require
absolute PAR values. Thus, we did not require continu-
ous integration over the entire day, nor did we require
locally corrected values for atmospheric parameters
used by REST2 (we used REST2 default values).

We calculated only direct radiation (roughly 77-81%
of total PAR, calculated for our transects using REST2
without overshadowing). Indirect PAR radiates from
the entire sky, but not isotropically. Complex modeling
of overshadowed, indirect, anisotropic PAR (Gueymard
1987) was beyond the scope of this study, as was meas-
urement of the relatively minor shade created by shrubs

and herbs.

Data analysis é

Because of logistics associated with our remote sites,
not all transects were established until 2009, and not all
transects were monitored annually. In addition, we ex-
cluded 2007 data from our analyses, because that year
many grouseberries on warm, south and west aspects
ripened early and then became desiccated. When we
began our counts, many fruits at warm sites had fallen,
and other fruits fell to the ground when we touched
the shrubs, becoming unrecognizable in the litter.

We scaled dPAR values relative to that from the tran-
sect with the largest dPAR value, which was set at 100.

PENGELLY AND HAMER: GROUSEBERRY FRUIT PRODUCTION IN BANFF

403

To avoid pseudoreplication, we combined the 22 tran-
sects at the prescribed burn sites into eight sampling
units, i.e., the four burn sites and the four adjacent un-
burned forests, and we combined three transects at
another site because they had essentially the same slope
steepness, aspect, and forest cover. This produced 15
39 sampling units (transects or collapsed transects) for
2006-2012. Because the transects at the prescribed burn
sites were reduced to four pairs of sampling units, we
present differences in fruit densities between burned
and unburned habitat using descriptive statistics.

Fruit densities were characterized by large variabil-
ity, with numerous outliers. Therefore, we used robust
regression analysis (R open-source software, version
3.0.2, WRS package) to assess the relation between
fruit production and dPAR. This robust Theil-Sen based
regression method uses bootstrap to analyze unconven-
tionally distributed data and more accurately reflects
trends in the data compared with ordinary least squares
methods or nonparametric data transformations, which
can give biased results (Wright and Field 2009).

Results
Annual fruit abundance

Based on 28 sampling units for which we had con-
tinuous records during 2008-2012, and scaling results
relative to the year of highest fruit density (2010), we
recorded relative Grouseberry fruit abundance of 22%
in 2008, 35% in 2009, 100% (reference year) in 2010,
32% in 2011, and 48% in 2012. We also estimated rel-
ative abundance of 103% in 2006 based on 15 sampling
units for which we had continuous records for 2006—
2012.

Prescribed burns

In 2004, 3 years post-fire, Grouseberry fruit density
in the burned transects averaged 0.17 times that in the
adjacent, forested transects at the two sites we measured
(Figure 2). The 2001 fires had burned the Grouseberry
plants to ground level, and the plants were immature in
2004 (average Grouseberry height in the burns, 5.6 cm

@ Forest
2500
®@ Burn
i:
2 1250
2004 2005 2006 2008

FIGURE 2. Grouseberry (Vaccinium scoparium Leiberg ex
Coville) fruit production (fruits/m? of Vaccinium cov-
er) during 2004-2008 at three prescribed burns con-
ducted in Banff National Park, Alberta, during autumn
2001. Matched transects were located in adjacent, un-
burned forests. Data are missing for the third burn in
2004, the second burn in 2005, and all burns in 2007
when early fruit-fall invalidated counts.

404

[SD 0.9 cm] versus 12.7 cm [SD 1.9 cm] in the forests).
In 2005, 4 years post-fire, results were mixed: one burn
had 0.64 times the fruit density of the forested tran-
sects, whereas the second burn we measured had 2.3
times the fruit density of the forested transects.

In 2006 and 2008, 5 and 7 years post-fire, fruit den-
sities in the three burns averaged 5.7 times and 4.8
times the densities recorded for the forested transects,
respectively. There are no data for 2007, when early
fruit-fall invalidated our counts.

In 2009, 8 years post-fire, fruit densities in the three
west-facing burns averaged 0.41 times those in the
forested transects, but at the east-facing site, fruit den-
sity in the burned transects was 10.1 times that in the
adjacent, forested transects (Figure 3). During 2010—
2012, fruit densities in the three west-facing burned

2500

1250

Fruits/m2

2009

2010

THE CANADIAN FIELD-NATURALIST

Vol. 128

transects were 3.3 times to more than 30 times those in
the adjacent forested transects; those in the east-facing
burned transects were 6.2 to more than 20 times those
in the forested transects. Not all ratios are defined
because the scarcity of fruits in some forested tran-
sects resulted in very small denominators and, thus,
misleading ratios.
Relation between Grouseberry fruit density and site
openness

Grouseberry fruit densities were positively associat-
ed with dPAR in 2006 and 2010-2012 (P < 0.02), but
not in 2008 or 2009 (P > 0.35, Table 1). In 2010-2012,
this positive relationship held whether the four sam-
pling units in the prescribed burns were included or
excluded (P < 0.02, Table 1). Figure 4 illustrates results
for 2010, the year of highest fruit density.

m Forest - West
® Burn - West
Z% Forest - East
S Burn - East

2011 2012

FIGURE 3. Grouseberry (Vaccinium scoparium Leiberg ex Coville) fruit production (fruits/m2 of Vaccinium cover) during
2009-2012 at three west-facing prescribed burns and one east-facing prescribed burn conducted in Banff National
Park, Alberta, during autumn 2001. Matched transects were located in adjacent, unburned forests.

TABLE 1. Association between Grouseberry (Vaccinium scoparium Leiberg ex Coville) fruit production and incoming direct pho-
tosynthetically active radiation including and excluding the four prescribed burn sites, Banff National Park, Alberta. 2006-2012.

Toons SEE

Regression
coefficient Explanatory
Year Intercept b (95% CI) power n I
Including prescribed burns
2006 —412 11.25 (0.39 to 25.16) 0.45 it) 0.017
2008 50 0.03 (—0.57 to 2.04) 0.00 34 0.354
2009 142 = OLG5i(G2187 toe) 0.04 33 0.993
2010 oll 7.70 (4.65 to 14.04) 0.38 3M) < 0.001
2011 ele 2, 3.50 (0.33 to 8.29) 0.47 34 < 0.001
2012 = 1195 4.72 (2.16 to 9.01) 0.49 34 <0.001
Excluding prescribed burns .
2006 579 14.93 (—7.72 to 15.72) 0.38 10 0.541
2008 61 —0.34 (-1.84 to 0.756) 0.05 27 0.5 14
2009 153 ~0.90 (—4.18 to 1.83) 0.04 28 0.541
2010 30 5.84 (2.51 to 12.99) 0.21 31 < 0.001
2011 =a) 1.24 (0.15 to 5.69) 0.21 28 0.015
2012 1) he! 3.16 (1.09 to 6.54) 0.34 28 0.007

RI I I IEEE EEE
Note: CI = confidence interval.

2014

2000

1000

Fruits/m?

2000

Fruits/m?
8

0) 25 50 75
dPAR

100

Ficure 4. Relation between Grouseberry (Vaccinium scopari-
um Leiberg ex Coville) fruit production (fruits/m* of
Vaccinium cover) in 2010 and direct photosynthetically
active radiation adjusted for overshadowing by topog-
raphy and conifer foliage and summed for the 92-day
period from | June through 31 August in Banff Na-
tional Park, Alberta, including the four prescribed
burns (a) and excluding the four prescribed burns (b).
The best-fit lines were derived from Theil-Sen robust
regression.

Discussion

We recorded high Grouseberry fruit densities in the
open sites created when the 2001 prescribed fires re-
moved the forest canopy. During 2006-2012, fruit den-
sities at the burn sites were 3.3 to more than 30 times
those in the adjacent forest except in 2009 when den-
sity was high only in the east-facing burn site (Fig-
ures 2, 3). Similarly, Weaver ef al. (1990) reported
that Grouseberry fruit production at a site where the
Whitebark Pine (Pinus albicaulis Engelmann) canopy
had been removed was 6 times that in two adjacent
forests. High fruit production at sites where fire, log-
ging, avalanching, or other factors have removed the
forest overstory is common for many fruiting species
(Lindzey et al. 1986; Hamer 1996; Greenberg ef al.
2007; McCord et al. 2014).

PENGELLY AND HAMER: GROUSEBERRY FRUIT PRODUCTION IN BANFI

405

We attribute the fact that, 3 and 4 years following
fire, fruit production was lower in the west-facing pre-
scribed burns than the adjacent forests in three of the
four cases (Figure 2) to the time required for Grouse-
berry shrubs to re-establish following fire. The fires re-
moved the above-ground portions of the Grouseberry
shrubs and left a black scorched surface layer. Five
years post-fire, however, the burned transects produced
5.7 times more fruits than the transects in the adjacent
forests. A Buffaloberry (Shepherdia canadensis |L|
Nuttall) study in the same valley of Banff National Park
also found a 5-year lag in fruit production following
prescribed fire (Hamer 1996).

Changes in soil nutrient status can occur following
fire. However, in our prescribed burns, by 2012 (11
years post-fire), the fires’ effects on nutrient cycling
may have been minimal. The effect on nutrients can be
greatest immediately following fire, but nitrogen can
return to pre-fire levels in a few years and phosphorus
in a few months; effects on other nutrients can be even
more ephemeral (Certini 2005). We did not conduct soil
or nutrient analyses for our study area.

We found a positive relation between Grouseberry
fruit density and dPAR in 4 of the 6 years analyzed
(Table 1). There was no significant relation in the 2
years of lowest fruit production. For 2010-2012, these
positive relations held, with or without the prescribed
burn sites included in the analysis, and, hence, were not
simply driven by the higher fruit production we found
in the prescribed burns following the 5-year lag in post-
fire recovery. In 2006, the relation between dPAR and
fruit density did not hold when the burns were exclud-
ed, but this analysis is based on only 10 sampling units
because several study sites had not yet been established.

When measuring fruit density, we subjectively repo-
sitioned our quadrats (normally by a few decimetres or
less) from the 2-m marks along transects when nec-
essary to ensure high cover of Vaccinium inside the
frame. We also converted these counts to a 100% Vac-
cinium cover basis to standardize fruit densities among
sites and observers. Thus we measured relative or eco-
logical fruit density per square metre of Vaccinium
cover, not absolute fruit density per hectare of habitat.
This approach emulates the behaviour of frugivores,
which can forage with high efficiency by moving from
shrub to shrub (patch to patch).

Although we recorded high fruit densities at the four
burn sites we studied, we searched seven other pre-
scribed burns in Banff National Park, but did not find
sufficient Vaccinium cover for sampling. These other
burns were typically in warmer habitat at lower ele-
vations where there was a high cover of grasses and
other herbs. We also recorded low Grouseberry fruit
densities in some open but xeric, south-facing habitat.
This included an open, xeric, south southwest-facing
transect with the highest dPAR value of our study.
During 2008-2012, this transect averaged 0.29 times
the fruit density of a second transect located 20 m away

406

under more mesic conditions associated with 40% for-
est canopy cover. Keefer ef a/. (2010*) reviewed stud-
ies on Huckleberry that similarly found reduced fruit
production in fully open sites. We also recorded fruit
densities that were 5 or more times and 3 or more times
higher in the more mesic east-facing prescribed burn
site than in west-facing burn sites in 2009 and 2010,
and equal or greater densities in 2011 and 2012 (Figure
3). Although this observation is based on only one east-
facing burn site, it is consistent with observations from
Huckleberry studies. For example, Martin (1983) found
that the most productive Huckleberry sites were in
north- and east-facing burn sites. Similarly, the high-
est fruit densities recorded in a Buffaloberry study in
Banff National Park were on north northeast-aspect
slopes (Hamer 1996). A positive influence of mesic site
conditions on fruit production was also suggested by
Burton (1998*), who reported a stronger relation be-
tween Huckleberry fruit production and site moisture
conditions, than between fruit production and solar
radiation.

Our study did not specifically address the effects of
mesic site conditions given that it was limited in extent
(to 39 or fewer sampling units), focused on dPAR, and
did not permit a more comprehensive multivariate
analysis. Nevertheless, we have shown that Grouse-
berry fruit production increases with increasing site
openness and that moderate to high fruit production
can be restored when prescribed fire is applied in ap-
propriate habitat types.

Many of the prescribed fires in Banff National Park
have been on warmer slopes (e.g., south and west
aspect) and more often in the lower subalpine than in
the upper subalpine or near the treeline. When pre-
scribed fire is used on these warmer, drier slopes, the
post-fire community is often dominated by grasses. In
contrast, historic wildfires that burned through mesic,
upper subalpine habitat near the treeline have, in some
cases, led to extensive, open Grouseberry communities.
Our study documents the ecological benefits of both
high-elevation prescribed burns and wildfire in forests
where Grouseberry shrubs are widespread but fruit
production is low under the forest canopy.

Acknowledgements

This project was supported by Parks Canada, the
Banff National Park Resource Conservation Service,
and Northern Lights College. Chris Gueymard made his
solar radiation computer programs available to us and
provided suggestions; and Mark Roberts developed our
computer program with overshadowing functions. We
also thank B. Chruszez, A. Forshner, H. Galt, S. Her-
rero, A. Kortello, B. Low, S. Michel, R. Osiowy, J.
Park, L. Pengelly, S. Rasheed, A. Schultz, L. Wiggins,
M. Woode, N. Woode, D. Zell, and anonymous review-
ers for assistance.

THE CANADIAN FIELD-NATURALIST

Vol. 128

Documents Cited (marked * in text)

Achuff, P. L. 1982. Vegetation: Introduction and methods.
Pages 71-75 in Ecological (Biophysical) Land Classifi-
cation of Banff and Jasper National Parks. Volume II: Soil
and Vegetation Resources. Publication SS-82-44. Edited by
W. D. Holland and G. M. Coen. Alberta Institute of Pedol-
ogy, Edmonton, Alberta, Canada.

Burton, P. J. 1998. Inferring the response of berry-produc-
ing shrubs to partial cutting in the ICHme. Project No.
FR—96/97-118, final report. Science Council of British
Columbia, Burnaby, British Columbia, Canada. Accessed
March 2011. http://www.for.gov.be.ca/hfd/library/frbc 1998
/frbc1998mr156.pdf.

Janz, B., and D. Storr. 1977. The climate of the contiguous
mountain parks. Project report no. 30. Atmospheric Envi-
ronment Service, Parks Canada, Toronto, Ontario, Canada.

Keefer, M., W. Cocksedge, R. Munro, J. Meuleman, and
N. MacPherson. 2010. What about the berries? Managing
for understory species. The Centre for Livelihoods and
Ecology, Royal Roads University, Victoria, British Colum-
bia, Canada. Accessed March 2011. http://www. for.gov
-be.ca/hfd/library/FIA/2009/FSP_Y093329¢.pdf.

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Received | April 2014
Accepted 2 July 2014

Song Discrimination by Male Mourning Warblers (Geothlypis
philadelphia) and Implications for Population Divergence across the
Breeding Range

JAY PITOCCHELLI
Biology Department, Saint Anselm College, Manchester, New Hampshire 03102 USA; email: jpitocch@anselm.edu

Pitocchelli, Jay. 2014. Song discrimination by male Mourning Warblers (Geothlypis philadelphia) and implications for population
divergence across the breeding range. Canadian Field-Naturalist 128(4): 408-415.

Geographic variation in song may reduce or eliminate the ability of some populations to recognize each other as conspecifics,
possibly leading to assortative mating, reproductive isolation, and speciation. Song playback experiments, used to evaluate the
significance of geographic variation in song, have been particularly useful in discovering divergence among previously unknown
populations of sibling species. In this study, I report the results of song playback to male Mourning Warblers (Geothlypis
philadelphia) from populations throughout the breeding range and discuss the implications for population divergence. Four
regions in the breeding range contain unique song types or regiolects: western, eastern, Nova Scotia, and Newfoundland. Results
of reciprocal song playback experiments showed that males from the western and Newfoundland regiolects respond more aggres-
sively to songs in their own regiolect than those in the other regiolects. Interior populations, i.e., eastern and Nova Scotia regions,
showed little or no difference in aggressive response toward their own versus other regiolects. This pattern may be due to a
combination of geographic proximity of populations belonging to different regiolects, song learning, experience, and contact
during migration. Song discrimination by populations from the western Prairie Provinces and Newfoundland is consistent

with the existence of at least partial reproductive isolation at the geographic extremes of the breeding range.

Key Words: Mourning Warbler; Geothlypis philadelphia; song; playback experiment; song discrimination

Introduction

Birdsong is a reproductive display recognized by a
network of males and females of the same species
(Paterson 1985; McGregor and Dabelsteen 1996). It is
regarded as one of the most compelling examples of a
prezygotic isolating mechanism (Remsen 2005; Hall
and Hallgrimsson 2008). Because birdsong plays such
an important role in maintaining reproductive cohesion
among populations of the same species, instances of
gradual divergence, such as geographic variation, may
challenge the ability of birds to recognize songs from
different parts of the breeding range leading to repro-
ductive isolation and speciation (Edwards et al. 2005;
Price 2008). Geographic variation in oscine songbirds
is well known and has been described at two spatial
scales (see reviews of Mundinger 1982; Podos and
Warren 2007). Song differences between contiguous
populations separated by short distances and capable
of interbreeding is referred to as microgeographic or
dialect variation. Macrogeographic or regiolect varia-
tion occurs where large regions of the breeding range
contain unique song types.

In an article on the role that song variation plays in
species-level classification, Remsen (2005) asked how
much variation in song is required to exceed the limits
of species recognition, create barriers to gene flow, and
result in speciation. Studies of geographic variation in
song coupled with playback experiments present an op-
portunity to address this issue. Playback experiments
with territorial males have been used to test hypotheses
about song discrimination in many species at the dialect
and regiolect levels (Baker et al. 1981; Searcy et al.

1997; Price 2008). Similarly, song playback experiments
have been performed on captive females, implanted
with estradiol, to test female discrimination of song
types from different parts of the breeding range (Bal-
aban 1988; Searcy 1992; Searcy et al. 2002; Ander-
son 2009; Danner et al. 2011).

A general result of playback studies at the dialect
and regiolect levels is evidence that members of both
sexes are capable of song discrimination. Males react
more aggressively to homotypic or local songs shared
with their neighbours compared with foreign or het-
erotypic songs from different parts of the breeding
range. Females respond more favourably to homotyp-
ic songs (Price 2008). Research on the effects of song
variation and discrimination on gene flow has been
mixed. Studies at the microgeographic level have shown
that dialects do not act as barriers to gene flow in the
Yellow-naped Parrot (Amazona auropalliata; Wright
and Wilkinson 2001), the White-crowned Sparrow
(Zonotrichia leucophrys; Soha et al. 2004), and the
Brown-headed Cowbird (Molothrus ater; Fleischer
and Rothstein 1988). However, genetic and morpho-
logic divergence have been correlated with song dif-
ferences among subspecies of the Swamp Sparrow
(Melospiza georgiana; Liu et al. 2008) and regiolects
in several Palearctic species belonging to the genera
Parus and Phylloscopus (Helbig et al. 1996; Martens
1996; Irwin et al. 2001).

In this paper, I report the results of song playback
experiments with territorial male Mourning Warblers
(Geothlypis philadelphia). This species is monotypic,
with a large breeding range extending over much of

408

2014

Canada and parts of the eastern United States (Mayr
and Short 1970; Pitocchelli 201 1a). There is a well-
defined pattern of geographic variation in songs, with
four regiolects: western, eastern, Nova Scotia, and
Newfoundland. Songs of males within each regiolect
are very similar; variation within a regiolect is usually
minor and involves the omission or substitution of a
single syllable. A zone of admixture exists in western
Ontario, Minnesota, and Wisconsin where males from
both the western and eastern regiolects breed. Hybrid
songs containing a mix of western and eastern sylla-
ble types have been recorded in this zone (Pitocchelli
2011b). Although geographic variation in song exists,
it is unknown to what degree these song differences
are recognized by populations of each regiolect.

I conducted a series of reciprocal playback experi-
ments within the geographic boundaries of each regi-
olect to test three hypotheses. First, populations within
each regiolect discriminate between songs from all of
the other regiolects. The prediction from this hypothesis
is that the responses of males will be more aggressive
toward playback of homotypic songs from the same re-
giolect versus heterotypic songs of each of the other
regiolects. Second, populations within each regiolect
are partly capable of song discrimination. The predic-
tion here is that there will be higher levels of aggres-
sion by males to homotypic songs versus heterotypic
songs of some but not all of the other regiolects. Third,
populations within each regiolect do not discriminate
between homotypic songs and heterotypic songs of any

—— - we
y

FIGURE 1. Distribution of the four regiolects of the Mourning

experiments, and sample songs from: A,

PITOCCHELLI: SONG DISCRIMINATION BY MOURNING WARBLERS

Ba

409

of the regiolects. The prediction from this hypothesis
is that males within each regiolect show equal levels of
aggression toward homotypic and heterotypic songs.
The results of this study show that response differences
and song discrimination do occur, but are not consistent
throughout the breeding range. I discuss the effect of
geographic proximity on the pattern of song discrimi-
nation and the implications of song discrimination for
reproductive isolation among breeding populations.

Study Area

I selected study sites for playback experiments with-
in the geographic boundaries defining each regiolect:
western, eastern, Nova Scotia, and Newfoundland
(Figure 1). Latitude and longitude of study sites were
Hudson Bay, Saskatchewan, 53.96°N, 102.35°W;
Duck Mountain Provincial Park, Manitoba, 51.89°N,
100.85°W; Marathon, Ontario, 48.68°N, 86.10°W;
Twin Mountain, New Hampshire, 44.20°N, 71.70°W;
Oxbow, Maine, 46.33°N, 68.42°W; Wreck Cove, Nova
Scotia, 46.08°N, 60.80°W; and Stephenville Crossing,
Newfoundland, 48.60°N, 58.35°W.

Methods

Approximately 30 focal males from each regiolect
were evenly divided into three experimental groups
(Table 1). Group I was presented with homotypic songs
from their own regiolect and heterotypic songs from
one of the foreign regiolects (e.g., western focal males
presented with western versus eastern songs). Group II

ee

SSS
Z Ppt
1S i i ap ai

Newfoundland
Regiolect

(ox) P Si a
wy.
el OS

Nova Scotia

: Regiolect
oe gg Page wos Chee

0 500

1000

kilometers

Warbler (Geothlypis philadelphia), locations of the playback
the western regiolect; B, the eastern regiolect; C, the Nova Scotia regiolect:

and D. the Newfoundland regiolect Western study sites were Hudson Bay, Saskatchewan (HB) and Duck Mountain
¢ > o 4) J

Provincial Park, Manitoba

(DM); eastern study sites were Marathon, Ontario (MA), Twin Mountain, New Hampshire

(TM), and Oxbow, Maine (OX); the Nova Scotia study site was Wreck Cove (WC), and the Newfoundland study

site was Stephenville Crossing

(ST). Shaded region indicates distribution of breeding range.

410

THE CANADIAN FIELD-NATURALIST

Vol. 128

TABLE |. Design and number of playback experiments to test song discrimination by Mourning Warblers (Geothlypis

philadelphia) throughout their breeding range.

ees

Mean difference

Focal in Principal Level of
Playback male’s Component | significance
locality song* Regiolect of playback songs (n) Scores (95% Cl) Student’s f testt
Western regiolect (WR): WR Group I: WR vs. ER (9) 0.46 (0.39) P=0.026f
Hudson Bay, Group II: WR vs. NSR (10) 1.72 (0.67) P<(0.001
Saskatchewan; Duck Group III: WR vs. NFR (10) 0.99 (0.58) P=0.004
Mountain Provincial
Park, Manitoba
Eastern regiolect (ER): ER Group I: ER vs. WR (11) 0.38 (0.67) P=0.238§
Marathon, Ontario; Twin Group II: ER vs. NSR (10) 1.16 (0.68) P=0.004
Mountain, New Hampshire; Group III: ER vs. NFR (9) 0.99 (1.16) P=0.085§
Oxbow, Maine
Nova Scotia regiolect NSR Group I: NSR vs. WR (10) 0.00 (1.18) P=0.9978§
(NSR): Wreck Cove Group II: NSR vs. ER (12) 0.48 (0.58) P= 0.095§

Group III: NSR vs. NFR (10) —0.04 (0.79) P=0.913§

Newfoundland regiolect NFR Group I: NFR vs. WR (10) 1.67 (0.64) P<0.001
(NFR): Stephenville Group II: NFR vs. ER (10) 1.08 (0.62) P=0.003
Crossing Group III: NFR vs. NSR (10) 1.27 (1.06) P=0.024t

Note: CI = confidence interval.
*Confirmed by digital recordings.

+Two-tailed comparison; null hypothesis is that the mean difference is equal to zero.

t Not significant after sequential Bonferroni correction.
$Not significant.

was challenged with homotypic songs and heterotypic
songs from a different regiolect (e.g., western focal
males presented with western versus Nova Scotia
songs). Group III was challenged with homotypic songs
and heterotypic songs from the remaining regiolect (e.g.,
western focal males presented with western versus
Newfoundland songs). To ensure that multiple stimuli
represented a class of stimuli and to minimize pseu-
doreplication (Kroodsma 1989; Kroodsma et al. 2001),
each focal male was presented with a unique combi-
nation of different, randomly selected homotypic and
heterotypic songs. I selected 30 western, 30 eastern, 30
Nova Scotia, and 30 Newfoundland exemplar songs
for this study. Each song was from a different male. I
made the selections based on the best recordings from
earlier work on this species (Pitocchelli 1988, 1990,
2011b).

I used the simultaneous, two-speaker presentation
of auditory stimuli model for playback experiments
(Lanyon 1978). Simultaneous presentation of different
song types tests the comparative prioritization of re-
sponses and reveals which song is perceived to be a
greater threat to a territorial male (Darren Irwin, Bio-
diversity Research Centre and Department of Zoology,
University of British Columbia, 7 April 2014, personal
communication). I placed two PureFi Anywhere speak-
ers (Logitech, Lausanne, Switzerland) approximately
30 m apart (positions | and 2), on either side of the first
song post that I observed for each male. I assumed that
this song post was within the territory of the focal male.

The speakers were each connected to an iPod (Apple
Inc., Cupertino, California, USA). For the first four
minutes of the experiment (Period I), regiolect A was
played at position | while regiolect B was played at
position 2 (A and B representing any randomly select-
ed pair of homotypic and heterotypic songs; Figure
2). After two minutes of silence, I conducted another
four minute presentation (Period II) with the same
songs but with positions reversed, i.e., regiolect B was
played at position | while regiolect A was played at
position 2). Approximately 19 homotypic and 19 het-
erotypic songs were presented during each four-minute
period, which simulated the average natural rate of one
song per 13 seconds by territorial males.

The songs were digitized and assembled using Raven
Pro version 1.4 for Mac OSX (Cornell Lab of Ornithol-
ogy, Bioacoustics Research Program, Ithaca, New York,
USA) and iTunes version 8.2.1 (Apple Inc.). Homo-
typic and heterotypic songs were arranged so that they
did not overlap and males could clearly hear all songs
during the experiment. When necessary, I manipulated
the songs using the amplify settings in Raven so that
they were all similar in amplitude. All experiments were
performed from 6 to 11 a.m. in June 2008, 2009. and
2010.

I used four response variables to measure aggressive
behaviour of the focal males toward either homotypic
or heterotypic songs: 1) number of flights toward and/or
over a speaker, 2) number of chip notes directed toward
a speaker, 3) number of songs directed toward a speak-

* = ee ~ - y .
2014 PITOCCHELLI: SONG DISCRIMINATION BY MOURNING WARBLERS 41]

Gap of
Silence
Period | Period II
iPod 1 | iPod 1
Focal Male Eastern Regiolect Western Regiolect

Eastern Regiolect

ee | 2

iPod 2
= a Western Regiolect . Eastern Regiolect

FIGURE 2. Experimental setup for an eastern focal male Mourning Warbler (Geothlypis philadelphia) and his song type challenged
by playback of homotypic eastern regiolect and heterotypic western regiolect songs. Periods I and II were four minutes

long and separated by two minutes of silence.

er, and 4) time spent within 3 m of a speaker. For each
focal male, the responses to the homotypic song for
Period I and Period I] were combined, as were the
responses to the heterotypic song for Periods I and II.

Before statistical analysis, I transformed the response
data following guidelines in Zar (2010) to avoid devi-
ations from normality and to account for the large num-
ber of zero data entries often encountered in playback
experiments (Balakrishnan and Sorenson 2006). Num-
ber of flights, number of chip notes, and number of
songs were square root transformed. The time spent
within 3 m of a speaker was log,, transformed, log,,
(x + 1). I subjected the transformed data to a principal
components analysis to reduce the correlated variables
to a new set of uncorrelated principal component (PC)
scores (Martin and Martin 2001; Balakrishnan and
Sorenson 2006). Each male received a PC score based
on its combined responses to a homotypic song dur-
ing Periods I and II. A separate PC score was calcu-
lated from the combined responses to the heterotypic
song during Periods I and II. I chose scores for the
first PC (PC1) for subsequent statistical tests because
PC1 explained 71.4% of the variation and it was also
the best index of aggression. All response variables
were positive for PC1: number of flights (0.67), num-
ber of chips (0.98), number of songs (0.02), and time
spent within 3 m of the speaker (0.24). Higher PCI
scores indicated higher levels of aggression toward a
song type.

I chose a paired samples Student's ¢ test to analyze
these data because each focal male was exposed to two
treatments: playback of homotypic songs and playback
of heterotypic songs. This test determines whether the
average difference between PCI scores for homotypic

and heterotypic songs is significantly different from
zero for each group in a regiolect. Differences signif-
icantly greater than zero indicated a stronger response
to the homotypic song, whereas differences significant-
ly less than zero indicated higher levels of aggression
toward heterotypic songs. Significant differences in
either direction were considered to be evidence of
song discrimination. The absence of a significant dif-
ference from zero indicated equal levels of aggression
toward homotypic and heterotypic songs and was inter-
preted as an inability to discriminate between these
songs.

I performed three paired samples Student’s f tests,
one for each group (I, I, HI) in each regiolect (Table
1). The design produced a total of 12 statistical tests.
The differences for PC1 scores did not deviate from
a normal distribution for 10 of the 12 statistical tests,
which meets a critical assumption of the paired sam-
ples Student’s ¢ test (Zar 2010). In cases where multiple
statistical tests are used, Rice (1989) recommended
using a sequential Bonferroni adjustment to correct for
possible inflation of probability levels. | began the
sequential Bonferroni application with an alpha level
indicating a significant difference at P < 0.0042 (P =
0.05/12 Student’s ¢ tests). The use of this adjustment
has been controversial (Moran 2003), so I have pre-
sented the results with and without the correction in
Table |. I also discuss these specific instances in the
Results and Discussion sections below.

Results

The results of the paired samples Student’s 7 tests
were mixed. Aggressive responses toward homotypic
songs and heterotypic songs were significantly differ-

412

2.20

g

S 1.20

=

a

ww

& 0.20

: aed

= East NS NF West NS F
0.80

Western Males Eastern Males

-1.80 ~

THE CANADIAN FIELD-NATURALIST

Vol. 128

West East F

Nova Scotia Males Newfoundland Males

FIGURE 3. Mean differences in first principal component (PC1) scores between responses to homotypic and heterotypic songs
by focal male Mourning Warblers (Geothlypis philadelphia) within each regiolect. Error bars are confidence intervals
that reflect sequential Bonferroni-corrected alpha values. West = western regiolect, East = eastern regiolect, NS = Nova

Scotia regiolect, NF = Newfoundland regiolect.

ent among males from extreme western and eastern
parts of the breeding range. But interior populations
from the eastern and Nova Scotia regiolects showed
little or no differences in level of aggression toward
homotypic and heterotypic songs (Table 1, Figure 3).

For males singing the western regiolect in the Prairie
Provinces, results from Student’s ¢ tests suggest com-
plete song discrimination. Western males responded
more strongly to homotypic songs compared with het-
erotypic songs from Nova Scotia (¢ = 5.79, two-tailed,
df= 9, P < 0.001) and Newfoundland (t = 3.87, two-
tailed, df = 9, P = 0.004) regiolects. The results sug-
gest that western males also responded more aggres-
sively to homotypic songs versus the eastern regiolect,
but the comparison was not significant after employing
the sequential Bonferroni correction (t = 2.74, two-
tailed, df= 8, P= 0.026).

The Student’s paired ¢ test results for eastern males
(Table 1, Figure 3) supported the partial song discrim-
ination hypothesis. Eastern males responded more ag-
gressively to homotypic songs versus heterotypic songs
from Nova Scotia (t = 3.85, two-tailed, df = 9, P =
0.004). But there were no significant differences in
responses to homotypic and heterotypic songs from the
western (¢ = 1.26, two-tailed, df = 10, P = 0.238) or
Newfoundland (t= 1.99, two-tailed, df = 8, P = 0.085)
regiolects.

The results from experiments with Nova Scotia
males supported the third hypothesis of no song dis-
crimination (Table |, Figure 3). There were no signif-

icant differences in response to their own songs versus
songs from the western (f = 0.004, two-tailed, df= 9,
P =0.997), eastern (t = 1.83, two-tailed, df = 11, P=
0.095), and Newfoundland (¢ = —0.11, two-tailed, df=
9, P=0.913) regiolects.

In contrast, the results for Newfoundland males sug-
gested complete discrimination (Table 1, Figure 3),
similar to that found in western males. The differences
in responses by Newfoundland males toward homo-
typic songs compared with heterotypic songs from
the western (t = 5.85, two-tailed, df = 9, P < 0.001)
and eastern (¢ = 3.93, two-tailed, df = 9, P = 0.003) regi-
olects were significant. In both cases, males responded
more aggressively toward homotypic songs. Although
there also was a difference between responses to homo-
typic versus Nova Scotia regiolects by these males (1
= 2.70, two-tailed, df = 9, P = 0.024), it was not sig-
nificant after the sequential Bonferroni correction.

Discussion

These results for song discrimination by the Mourn-
ing Warbler are in general agreement with previous
studies of song playback experiments in other species
where males exhibited higher levels of aggression
toward local, homotypic songs than heterotypic songs
from distant parts of the breeding range (Regelski and
Moldenhauer 1996; Searcy et al. 1997; Dufty and
Hanson 1999; Nelson and Soha 2004; Price 2008).
However, not all populations discriminated between
homotypic and heterotypic songs (Table 2). Mourning

TABLE 2. Ability of focal males to discriminate among songs of other Mourning Warblers (Geothlypis philadelphia) from
four regiolects within the species’ breeding range. No discrimination = no significant difference in responses to homotypic
and heterotypic songs; Discrimination = significant difference in responses to homotypic and heterotypic songs; Equivocal
= P <0.05 based on Student’s ¢ tests, but not significant after sequential Bonferroni correction.

Regiolect of focal male

Response to regiolect of playback song

Western Eastern Nova Scotia Newfoundland
Western =F Equivocal Discrimination Discrimination
Eastern No discrimination _ Discrimination No discrimination
Nova Scotia No discrimination No discrimination — No discrimination

Newfoundland Discrimination

Discrimination

Equivocal —

.———s—s—S—SSsSsSMS9090”3_—="—M$”@—”—™@M”MS SSS SS

2014

Warbler males at the extremes of the breeding range
showed the highest levels of discrimination by respond-
Ing more aggressively to homotypic than heterotypic
songs. Males from the western and Newfoundland regi-
olects could discriminate between their own songs and
at least two of the other regiolects. They may also rec-
ognize differences for the remaining regiolects but the
Student’s ¢ test results were not significant after the
sequential Bonferroni correction. These findings sup-
port the hypothesis of complete discrimination by pop-
ulations at the extreme eastern and western parts of
the breeding range.

Song discrimination by interior populations of the
eastern regiolect was weaker and supported the partial
discrimination hypothesis. Eastern males discriminated
between eastern songs and the Nova Scotia regiolect.
However, they did not discriminate behaviourally be-
tween homotypic songs and the neighbouring Western
regiolect or between homotypic songs and the New-
foundland regiolect. The results for the Nova Scotia
experiments supported the complete lack of discrimi-
nation hypothesis. These males showed equal levels of
aggression toward homotypic and heterotypic songs
when challenged with songs from all of the other regi-
olects.

One explanation for the spatial pattern of discrimi-
nation by Mourning Warbler males is a combination of
the effects of geographic proximity and song learn-
ing. Repeated exposure and song learning have been
shown to modify song recognition among and within
species (Irwin and Price 1999; MacDougall-Shackle-
ton et al. 2001; Price 2008). If geographic proximity
and experience influence song discrimination in
Mourning Warblers, then we should observe two dif-
ferent results from these playback experiments. Allo-
patric populations separated by large distances should
exhibit song discrimination because there is no contact
or opportunity to learn each other’s songs. In contrast,
sympatric populations should show little or no discrim-
ination because of geographic proximity and previous
experience with different song types that they may later
recognize as equally potent threats to territorial own-
ership.

Males from the western and Newfoundland popu-
lations were mutually capable of song discrimination
between their regiolects. Results for these allopatric
populations support the idea that a lack of contact and
experience with songs leads to song discrimination.
The inconsistent patterns of discrimination by popu-
lations from the interior of the breeding range appear
to support the contention that populations in close prox-
imity may not discriminate among different songs. For
instance, eastern males did not recognize differences in
western songs and these populations overlap in a large
admixture zone in the western Great Lakes region.
However, the role of geographic proximity is still un-
clear for these interior populations.

PITOCCHELLI: SONG DISCRIMINATION BY MOURNING WARBLERS

413

The playback experiments within the western and
eastern regiolects were outside the admixture zone (Fig-
ure |), so it is unclear why males from New Hampshire
and Maine could recognize the western regiolect as a
threat equal to eastern songs from other locations.
Another problem arises from results where allopatric
populations were not able to discriminate between
homotypic and heterotypic songs. Eastern and New-
foundland populations are separated by the Gulf of
St. Lawrence but eastern males responded equally to
both song types. Nova Scotia and western populations
are also allopatric, but Nova Scotia males reacted
strongly to western songs. An alternative explanation
for some of these inconsistencies in song discrimina-
tion is that the divergence among song types from the
eastern and Nova Scotia regiolects is too recent and
does not exceed the limits of recognition by these
males. Although Pitocchelli (2011b) found statistical
differences in syllables and physical parameters of
these songs, males still recognize them as a threat to
territorial ownership.

Playback experiments that show song discrimina-
tion among different populations have been cited as
evidence of reproductive isolation (Balakrishnan and
Sorenson 2006; Danner et al. 2011). Irwin and col-
league’s (2001) study of mitochondrial DNA differ-
ences among populations of Old World Leaf Warblers
(Phylloscopus spp.) confirmed reproductive isolation
that paralleled evidence from playback experiments.
Studies of mitochondrial DNA and playback experi-
ments on Chiffchaff superspecies (Phylloscopus) pro-
duced similar results (Helbig et a/. 1996; Martens
1996). If song discrimination by Mourning Warblers
also indicates prezygotic isolation, then populations
from the Prairie Provinces may be reproductively iso-
lated from allopatric populations in Nova Scotia and
Newfoundland. Newfoundland populations may also
be reproductively isolated from continental populations
based on successful discrimination of their songs from
songs of the western and eastern regiolects.

Although these results suggest prezygotic isolation
between some populations, additional playback exper-
iments with females (if possible) would be informa-
tive. Relying on male responses alone may not
always indicate isolation between different song pop-
ulations. Danner ef al. (2011) found gender differences
in Rufous-collared Sparrow’s (Zonotrichia capensis)
responses to homotypic and heterotypic songs. Males
did not differ in their responses to homotypic or het-
erotypic dialects, while females reacted more strongly
to the local, homotypic songs. The pattern of female
responses to different dialects was correlated with
genetic differences between populations while male
responses were not. Relying on evidence from male
responses alone would not have revealed this diver-
gence. Analyses of mitochondrial DNA and nuclear
DNA from Mourning Warblers in each regiolect would
ultimately be necessary to clarify actual levels of pop-

414

ulation divergence and whether genetic divergence is
correlated with song differences in this species.

Acknowledgements

This work was funded in large part by a summer
research grant from Saint Anselm College. I thank
the staff at the Geisel Library, especially Jeff Waller,
Sue Gagnon, and Sam Urtz, for their help with procur-
ing books and journal titles through inter-library loan.
The manuscript benefited from insightful comments
provided by Darren Irwin, Robert Rockwell, and two
anonymous reviewers. | also thank Joe Troisi, Kathy
Flannery, and Lori LaPlante for discussions and sug-
gestions regarding experimental design and appropri-
ate statistical analyses.

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Received 8 May 2014
Accepted 20 July 2014

Book Reviews

Book Review Editor’s Note: We are continuing to use the current currency codes. Thus Canadian dollars are CAD, US.
dollars are USD, Euros are EUR, China Yuan Remimbi are CNY, Australian dollars are AUD and so on.

ZOOLOGY

Amphibian Conservation — Global Evidence for the Effects of Interventions

By Rebecca K. Smith, and William J. Sutherland. 2014. Pelagic Publishing, PO Box 725, Exeter, UK, EX1 9QU. 279 pages,

29.99 GBP, Paper.

This comprehensive British synopsis was prepared
with funding by Synchronicity Earth and Aria. It is the
fourth volume in the Synopses of Conservation Evi-
dence series and is linked to the online www.conser
vation evidence.com where the full text is available
without charge as an alternative to paying for it in print.
Its dense text summarizes and discusses selected world
studies concerned with the results, positive and nega-
tive, of tampering with natural populations. These have
been usually well-meaning attempts to compensate for
the damage done by expanding human populations.
Impacts may have been either direct by past eradica-
tion of individuals or indirect through destruction or
alteration of habitats critical to the survival of one or
more amphibian species.

The introduction stresses that this survey is intended
to influence those in a position to make decisions that
could save biodiversity one case at a time. Although
recommendations are not provided, basic ecology is
included. Interventions that are discussed are not ranked
by importance or extent of their effects or evaluated
by their quality. The multitudes of references to pub-
lications of many countries are grouped with the head-
ings where they are appropriate.

The text is broken into 14 sections each with key
messages and various subheadings. Ten concentrate
on threats: residential and commercial development
(interventions specific to development); agriculture
(engage farmers and other volunteers, terrestrial and
aquatic habitat management); energy production and
mining (mist habitat); transportation and service corri-

Amphibians and Reptiles in Minnesota

dors (install culverts or tunnels, barrier fencing, modi-

fy drains and kerbs, signs and road closures, assistance

during migration); human intrusions and disturbance
(signs and access restriction); natural system modifi-
cations (prescribed fire, herbicides, mechanical re-
moval of vegetation, regulate water levels); invasive
alien and other problematic species (reduce predation,
competition and habitat alteration by other species,
reduce parasitism and disease by chytridiomycosis and
ranaviruses); pollution (agricultural and industrial);
climate change and severe weather (irrigation, ephem-
eral ponds, shelter habitat, gradients). These are fol-
lowed by habitat protection (connectivity, buffer zones);
habitat restoration and creation (terrestrial, aquatic);
species management (translocate, captive breeding,
rearing, release); education and awareness raising
(campaigns, programs, citizen science).

A 17-page index concludes the book but is a disap-
pointment. Canadian studies are not cited by country or
province forcing the reader to search through the text
for references to them. However, there are entries for
other countries and many entries by species.

This is part of an ambitious series that aims toward
accumulating a comprehensive summary of evidence
on the effects of conservation interventions on saving
the diversity of life over the entire planet. Among other
titles completed is Bee, Bird, Farmland and more are
being prepared to cover different groups and habitats.

FRANCIS R. COOK

Curator Emeritus and Research Associate, Canadian Museum
of Nature, Ottawa, ON, Canada, K1P 6P4

By John J. Moriarty, and Carol D. Hall. 2014. University of Minnesota Press, Suite 290 111 Third Avenue South. Minneapolis,

MN, USA, 55401. 372 pages, 39.95 CAD, Paper.

This new survey of the Minnesota herpetofauna
comprehensively updates Amphibians and Reptiles
Native to Minnesota by Barney Oldfield and John J.
Moriarty (1994; reviewed by FRC 1998 Canadian
Field-Naturalist 112(1): 170-171). The extent of new

information now available is reflected in a Resources
section which includes 6 Minnesota, 6 Regional (Upper
Midwest), 12 North American, and 8 General refer-
ences published since the earlier treatment. This is fur-
ther emphasized in the Literature Cited, which has 144

416

2014

post-1994 entries in its 20-page coverage (included are
some duplications from the more general Resources
listings).

Minnesota lies to the south of the boundaries of west-
ern Ontario and Manitoba. It has a primarily north-
eastern and east-central North American herpetofauna
So it is not surprising that of the 53 species recorded
for the state, 48 are shared with Canada. The represen-
tation of the two classes is unbalanced as there are more
shared reptiles but many barely enter Canada whereas,
in contrast, many of the fewer shared amphibian species
present have moved extensively into the north. In all,
22 amphibians (14 frogs and toads and 8 salamanders),
and 26 reptiles (2 lizards, 16 snakes and 8 turtles) re-
corded also occur in Canada. For seventeen (1 frog, |
toad, 3 salamanders, | lizard, 8 snakes, 3 turtles) that
are shared with Canada the Minnesota occurrences are
not linked to Manitoba or northeastern Ontario but to
populations south of the Great Lakes. Three of the spe-
cies now included have been added to the state list from
the eastern part of Minnesota since 1994 and it is sug-
gested that three additional species (1 frog, 1 toad and
| spadefoot) that occur near the western border of the
state may yet be found within it. One of these occurs in
Manitoba.

The contents open with a map of Minnesota counties
with arbitrary divisions of the large northeast counties
(this map is copied for quick referenced on the inside
back cover). A forward stresses the varied interests
which this book serves. A preface and acknowledg-
ments deal with the herpetological activity in the state
and the growth of interest in conservation. An intro-
duction covers the definition of “herp”, the history of
herpetology in Minnesota, the Minnesota Department
of Natural Resources Nongame Wildlife Program, Min-
nesota Herpetological Society, and other herpetological
efforts, amphibian and reptile habitats (with coloured
maps of ecological provinces, sections, and subsec-
tions), natural vegetation of Minnesota, average pre-
cipitation, temperature, and a table of amphibian and
reptile distributions by ecological section. Watersheds
are discussed and mapped. A table presents amphibian
and reptile distributions by habitat type. Aquatic habi-
tats are lakes and ponds, marshes, prairie wetlands, and

Amphibians of Ohio

Book REVIEWS

417

peatlands. Terrestrial habitats are flood-plain forest-
coniferous-northern hardwood forest, and prairies. Dis-
turbed areas are discussed separately as agricultural
lands and urban and suburban habitats. Next come sug-
gestions for observing and studying amphibians and
amphibians and reptiles encompassing ethical field
methods, field study and care of captives. A conserva-
tion section, divided between habitat loss and pollution
is followed by amphibian declines, diseases, harvesting
pressures, and persecution. Common amphibian and rep-
tile problems covers snakes in house, garage, and yard
invasions, salamanders in basements, turtles nesting in
the yard, snapping turtles eating ducklings, and sala-
manders, frogs, on the road or yard.

A checklist gives the original describer(s), the year
named for each genus, species and subspecies. System-
atics and taxonomy follow Frost ef a/. 2006. Bulletin of
the American Museum of Natural History, and English
names follow Crother, editor, 2012, SSAR Herpeto-
logical Circular 39.

Species accounts are grouped by class and, within
each, by family, each prefixed by summaries of fea-
tures. Each individual species account starts with cur-
rent English (common) and scientific name, and con-
tinues with description, and distribution in United
States and Canada (but statements for the latter are
vague and often omit actual extent apparently in an
effort to be concise). A small map of Minnesota coun-
ties shows museum records indicated by half-filled
(pre-1960) or solid (post-1960) circles and literature
or sighting records as open circles. Habitat, and life
history, and remarks (including legal status) complete
each account.

Species of possible occurrence include mention of
released exotics and abbreviated species accounts for
three species that further field studies in the state may
yet find. Concluding the book is a glossary giving defi-
nitions from amelanism to ventral, resources (refer-
ences and organizations), literature cited, and index to
genera, species and subspecies.

FRANCIS R. COOK

Curator Emeritus and Research Associate, Canadian Museum
of Nature, Ottawa, ON, Canada, K1P 6P4

Edited by Ralph A. Pfingsten, Jeffrey G. Davis, Timothy O. Matson, Greg Lipps, Jr., Doug Wynn, and Brian J. Armitage. 2013.
Bulletin New Series Volume 17, Number 1, Ohio Biological Survey, P.O. Box 21370, Columbus, OH, USA, 43221-0370.

916 pages, 90.00 USD, Cloth.

The US state of Ohio lies along the southwestern
end of Lake Erie, between Pennsylvania on the east
and Indiana on the west. It is adjacent on the north to
the richest area (southwestern Ontario) for species of
amphibians and reptiles in Canada. Ontario includes
the most northern of the Lake Erie’s western islands. It
is therefore unsurprising that the accounts contained in

this detailed review of Ohio amphibians which in-
cludes the islands to the south (as well as the mainland
beyond) are very relevant to the interests of Canadian
herpetologists, naturalists, and conservationists.
Publication on Ohio’s amphibians and reptiles began
early in the nineteenth century when they were in-
cluded by Jared Porter Kirkland in his 1838 list cov-

418

ering the settled region west of the Appalachians at
that time. Ohio frogs and toads were treated in detail
by Charles F. Walker in 1946 based on field surveys
he initiated with colleagues and students in the 1930s
but a companion volume on salamanders did not appear
until the multi-authored treatment edited by Ralph A.
Pfingsten and Floyd L. Downs in 1989.

The growing attention on herpetological research
after the Second World War was reflected by the ini-
tiation of the Ohio Journal of Herpetology in 1957 by
the Ohio Herpetological Society. The journal was soon
rechristened as the Journal of Herpetology and its pub-
lisher as The Society for the Study of Amphibians and
Reptiles. This became a leading international research
journal, joining the longer established Copeia (origi-
nating at the American Museum in New York in 1913
and later adopted as the publication of the American
Society of Ichthyologists and Herpetologists) and Her-
petologica begun in Kansas in 1936 which became the
publication of the Herpetologists League. Despite its
world coverage and influence, many of the society’s
members continued to focus their efforts, at least in
part, on their native herpetofauna in Ohio.

The latest contribution of the latter and their sup-
porters is a weighty volume by many measures, includ-
ing sheer size (over 900 pages). This detailed treatise
begins by presenting summaries of the influences on
Ohio herpetofauna of its geology, soils, climate, habi-
tats, and the integrated Ohio environment. Then there
are sections on amphibian systematics (including diag-
nosis of families). The species accounts which follow
are the major contribution. They begin with identifi-
cation keys for all included species followed by indi-
vidual coverage for each member of five families of
salamanders (24 species, 13 of these occur in Canada)
and four families of frogs and toads (14 species, 12 of
which occur in Canada). A bonus account covers hybrid
salamanders which are usually female and may be dip-
loid, triploid or higher polyploids. These variously have
genes from /aterale combined with one or two of jef-
fersonianum, tigrinum, texanum, and barbori. Accounts
cover etymology, synonyms, taxonomic status, com-
mon names, description, and all aspects of natural his-
tory. They also include photos of typical and variant

THE CANADIAN FIELD-NATURALIST

Vol. 128

metamorphosed juveniles and adults, salamander lar-
vae and anuran tadpoles, eggs, drawings of key mor-
phological and behavioural features, tables of meas-
urements and photos of habitats. Together these set a
new standard for regional coverage. They draw not only
on herpetology research in the state but also studies
beyond its borders wherever significant observations
and data on species present in Ohio. The distribution of
each species Ohio is presented on a vegetation base
map with dots for only those county records that have
vouchers (photographs or museum specimens). Three
time periods are distinguished (before 1952, 1952-
1989, and after 1989). A small inset map gives the
whole North American range.

Final sections cover potential occurrences and exclu-
sions, and practical applications of the data gathered
for developing priorities for conservation of Ohio’s
amphibian diversity; amphibian conservation; Ohio
amphibian distributions; environmental applications.
Indicative of the comprehensiveness of this volume and
therefore its value as a reference to the ever-growing
number of productive researchers focussing on amphib-
ians, is that the double-columned Literature Cited takes
73 pages.

Support for this publication reflects, like its multi-
ple authorship, how effective group effort can be. Its
development has been supported by the Ohio Division
of Wildlife through its Wildlife Diversity and Endan-
gered Species Fund, with additional assistance for
printing from Cincinnati Zoo, the Cleveland Museum
of Natural History, the Columbus Zoo and Aquarium,
the Crane Hollow Foundation, the Toledo Zoo, the
Akron Zoo and the Toledo Naturalists’ Association.

In recent decades there has been a succession of new
guides to other states, but this is by far more compre-
hensive and detailed than is usually attempted and rais-
es the bar to a level that its successors will find hard to
exceed or even match. All the many people contribut-
ing to the research, review, editing, and publication
may be justly proud.

FRANCIS R. COOK

Curator Emeritus and Research Associate, Canadian Museum
of Nature, Ottawa, ON, Canada. K1P 6P4

Illustrated Checklist of the Birds of the World — Non Passerines

By Josep del Hoyo, Nigel Collar, David Christie, Andrew Elliott, and Lincoln Fishpool. 2014. Lynx Edicions, Montseny, 8,
08193 Bellaterra, Barcelona, Spain. 904 pages, 145.00 EUR, Cloth.

I am not a fanatic lister, but an observer who reli-
giously keeps notes. When I retired I realised the last
time I added up my life list was 40 years earlier. I de-
cided it was time for an update. Using the Sibley and
Monroe World List of Bird Names (1999) I tried five
times over four years to complete this daunting task.
There were many times when it was not clear which

species I saw and I had to undertake much slow. re-
search to resolve each issue.

Now we have a Lynx Edicions version of the world
bird list. The authors take 54 pages to review and ex-
plain the process that lead them to declare one group
of birds as a single species. The authors use the Bio-
logical Species Concept (BSC relies on defining repro-

2014

ductively isolated populations) along with a scoring
system that evaluates differences in morphology, song,
ecology and geographical relationships. Using this pro-
cess they have produced a volume that covers 4,372
extant species of non-passerines (Volume 2 — Passerines
will be published in 2016). Also covered are 99 species
that have become extinct since 1500, with 50 of them
illustrated.

Each entry gives the currently accepted species in
English. Then lists the names in French, German and
Spanish, as well as alternative common names in Eng-
lish. There is a brief set of taxonomic notes to support
the decision to accept species designation. All species
have a distribution map and a colour illustrations, in-
cluding distinctive subspecies where appropriate. (The
artwork is high quality and mostly taken from the
Handbook of Birds of the World series). The index is
colour coded to show extant taxa, old or invalid taxa
and extinct taxa — a useful idea. There are 34 maps
and a full bibliography.

So how useful is this new book? My first thought
was to check Hen Harrier. In Europe I had seen these
birds hunting mice over upland heath. When I arrived
in North America there was a raptor called a “Marsh
Hawk.” To me this was a different bird as it was a little
grubby in appearance and I typically saw it hunting and
catching frogs and snakes over wetlands. The bird’s use
of a stall-stopping dihedral was more pronounced. To
my surprise this was the same species — Circus cyaneus.
For 50 years I have waited for an authority to split these
harriers in two. Del Hoya and his colleagues have
finally taken this plunge.

My next challenge was to review the typical tou-
cans. On every trip to South America I have had to
research which species I have seen. Handbook of Birds
of the World Volume [Josep del Hoyo, Andrew Elliott
and Jordi Sargatal (2002), Lynx Edicions, Barcelona,
Spain] listed 7 full species. Del Hoya now lists 11
species and it took me a couple of hours to evaluate
whether my own notes were correct. (Wikipedia cur-
rently lists eight species with 6 sub-species). Where
the taxonomy will settle I would hate to guess. At least
the issue of whether the Red-billed Toucan (Ram-
phastos tucanus) and Cuvier’s Toucan ( Ramphastos
cuvieri) should be considered species (they are signifi-
cantly different in their bill colour) I am sure will rage
on, because they interbreed freely.

Six years ago I saw and photographed a nightjar in
central India. Originally I listed it as a Grey Nightjar
(Caprimulgus indicus) and not an Indian Nightjar (Cap-
rimulgus asiaticus). My 1999 field guide showed the
Grey distributed India-wide. Back in Canada, when 1
reviewed my notes, | became confused. Handbook of

BooK REVIEWS

419

Birds of the World Volume 5 [Josep del Hoyo, Andrew
Elliott and Jordi Sargatal (1999), Lynx Edicions, Bar-
celona, Spain| contained a reference to Caprimulgus
indicus as the widespread Jungle Nightjar. The Indian
Nightjar (Caprimulgus asiaticus) had a similarly broad
Indian distribution. Looking at my photo | finally
decided my bird was an Indian Nightjar. This new book
shows the Grey Nightjar (Caprimulgus jotaka) as hav-
ing a very limited range in north-east India and the Indi-
an Nightjar (Caprimulgus asiaticus) occurring through-
out the Indian mainland. This confirms my sighting as
an Indian Nightjar — doesn’t it?

My other question was did I miss or misidentify
Fork-tailed Swift among the large number of “Palm
Swifts” I saw in Mongolia. This book lists Fork-tailed
Swift as an alternative name of the Palm Swift sub-
species. However one researcher has split this “super-
species” into Salim Ali’s Swift, Blyth’s Swift, Cook’s
Swift as well as Pacific Swift. This split has been ac-
cepted by the International Ornithological Commit-
tee (IOC), but not the International Union for Con-
servation of Nature, nor del Hoya et al. (but with five
sub-species). For now I am sticking with Palm Swift
and del Hoya.

The authors have changed some of the English
names too. Why list Peregrine as invalid and replace it
with Peregrine Falcon? They retain the old English
names of Merlin, Kestrel, Hobby etc. but change Pere-
grine and Gyr. (To be consistent it should be Merlin
Falcon etc.). Original texts do not have this modifica-
tion (especially as “falcon” is the term for female only).
They have also “updated” names like Fea’s Petrel (to
Cape Verde Petrel). This could be confused with the
Cape Verde Storm Petrel (Oceanodroma jabejabe)
which is now regarded as a separate species by the
British Birding Association and others, but not by del
Hoya et al. Oh, the joys of taxonomy!

So is this book really useful? Without question it
is. It is now the most current and up-to-date checklist
of world birds. It has been assembled by people who
have spent over twenty years evaluating and deciding
the taxonomic issues related to birds. It is a complete
checklist of the non-passerine species using the most
up-to-date taxonomy. Having both an illustration and a
range map alongside of the species entry is a real bless-
ing. It is an essential purchase for researchers and for
all those engaged in world-wide birdwatching. Even if
you are not a combat lister, you still need to know which
species you have seen and where. It will now be my go-
to text. There is one critical question. Will Volume | be
out-of-date before they publish Volume 2?

Roy JOHN

2193 Emard Crescent, Ottawa, ON, Canada, K1J 6K5

THE CANADIAN FIELD-NATURALIST

Vol. 128

Life on the Rocks: A Portrait of the American Mountain Goat

By Bruce L. Smith. 2014. University Press of Colorado, 5589 Arapahoe Ave., Suite 206C, Boulder, CO, USA, 80303. 192 pages,

35.00 USD, Cloth. (Also available as an e-book)

Anyone who has ever done any rock climbing or
scrambling is likely in complete awe of the feats that
mountain goats manage in the steep terrain that is their
home year around. The photos in this book attest to
their ability to survive “life on the rocks,” and the text
helps the reader understand how they do it, physically
and physiologically.

While Life on the Rocks could be described as a “‘cof-
fee table book” by virtue of its size (31 cm tall x 24 cm
wide) and plethora of colour photographs (over 100, the
majority taken by the author), to limit it to that catego-
ry would do both the book and the author, a disservice.
Smith is a scientist with the US Fish & Wildlife Serv-
ice, and the photographs complement his well-written
text on the mountain goat’s taxonomy, anatomy, be-
haviour, environment and co-inhabitors of extreme ter-
rain, rather than the text just being an expanded cap-
tion for each photograph. As well as life history and
natural history (Part I), Smith discusses current pop-
ulation estimates and reasons for declining numbers,
and the conservation challenges that impact mountain
goats (Part II). And, throughout the book Smith weaves
in tales of his personal adventures as a field biologist
studying mountain goats, particularly in the Selway-
Bitterroot Wilderness of southwestern Montana.

The majority of the photographs are excellent, and
range from intimate portraits to broad landscape images.
The quality of some of the full-page images do suffer
from apparently having been scanned from decades-
old slides or colour prints (e.g., the landscape photo on
page 109), although these are often used to illustrate
a point or theme, for which they are adequate. (As an
aside, it would be interesting to know if the photographs
reproduce more sharply in the Kindle e-edition.) Many
photographs are reproduced in two-page spreads, some
of which are truly outstanding, such as the backlit goat
(including breath vapours) on pages 134135, also used
on the book’s coyer jacket, or the close-up of a pika
with a mouthful of vegetation (pages 160-161). Smith
includes many photographs of other species that share
the mountain goat’s environment, which help to illus-
trate his text. I did notice an error on photo pages 148—
149, where the caption refers to whitebark pine dying

from disease and insects, whereas the photo clearly
depicts lodgepole pine and subalpine fir, whose red
needles are likely the result of the phenomenon known
as “red belt,” where a temperature inversion causes the
tree’s needles to transpire, while the roots are still in
frozen ground and unable to take up moisture, causing
desiccation of the needles.

It would have been useful to have had the sketches
of different ages and sexes of mountain goats near the
beginning of the book, rather than near the end, so that
the reader could refer to them when looking at some of
the photographs. Smith highlights pertinent research in
an easy manner throughout the book, but while there is
a page of suggested references (including scientific
papers), I would have liked source notes for some of the
chapters, relating his statements to specific research
papers.

Relatively early in the book Smith states that “at its
core, the goat is both product and captive of its evolu-
tionary history and specialization as a mountain climb-
er.” This could also be an introductory statement for
the chapters on local and global conservation chal-
lenges. While wildlife and land managers can reduce
harvest levels and minimise human disturbance, only
a landscape approach will ensure that mountain goats
survive in a world with a changing climate. Smith
touches on the uncertainty of how mountain goats may
respond: their generalist diet may help them cope with
vegetation changes, but they are intolerant of summer
heat so where will they go when the temperature rises,
and what might changes in parasite loads and transmis-
sion vectors mean for the goat? As such questions are
just starting to be asked of mountain goats, Smith refers
to research involving other inhabitants of these moun-
tain ranges, such as pika.

If you’ve ever looked up at a mountain goat on a
seemingly impossible ledge and wondered how come
it doesn’t fall off that cliff or what does it eat in winter?
And how does it stay warm? then you will enjoy this
book and its many photos.

CyNbI M. SMITH
P.O. Box 70, Mountain View, AB, Canada, TOK 1NO

2014

BoOoK REVIEWS

The Fish in the Forest: Salmon and the Web of Life

By Dale Stokes, Photographs by Doe White. 2014. University of California Press, (California-Princeton Fulfillment Services),
1445 Lower Ferry Road, Ewing, NJ, USA, 08618. 159 pages, 29.95 USD, Cloth.

_ “The story of the Salmon Forest is a story of the
interconnectedness of the sea and land and a fish that
has evolved and become entwined in a landscape, a
story that serves as a metaphor for all of life on earth.”

These lines set the tone for Dale Stokes’ book on the
natural history of salmon and their environment in the
north Pacific. In just under 150 pages, Stokes describes
in careful detail how the “King of Fish’ is an important
ecosystem engineer in the forests of the north Pacific,
particularly the coasts of British Columbia and Alaska.
He does so with a tangible passion that sometimes
edges into reverence — Stokes clearly loves this fish,
and uses this book to tell us why we should too. The
Fish in the Forest is not short on scientific terminology
—we’re treated to a whole host of Latin species names
and biological details. But Stokes’ obvious excitement
for the subject saves it from being a dry textbook — it
feels more like listening to a passionate teacher lec-
ture on his favourite topic, for example: “Taken from
the Latin, seme/parous means “begotten once,’ a refer-
ence to their lyrical existence: they die soon after they
reproduce.”

Stokes routinely sees the poetic in everything, which
can get a little grating at times and overly anthropomor-
phic. Female salmon sound heroic when he describes
the final stages of their life cycle: they “invest their last
metabolic energy in egg production and in guarding
their nests until they become too weak and battered to
hold their position in the current, and they drift away to
die.” All technically true, but the language almost
makes the salmon sound like martyrs — not the most
apt analogy, as fish do not have the power of free will.
Still, Stokes’ writing style makes for a very enjoyable
and informative read. The full colour photographs by
Doc White are definitely assets — my favourites are the
underwater shots of glistening salmon. White’s photos
bring to life the whole Salmon Forest, from delicate
riparian vegetation to ancient cedars towering over
gravely creeks, and powerful apex predators to gelat-
inous fish eggs.

Stokes begins this story by precisely defining key
terms and addressing basic questions: What is a fish, a
salmon, the Salmon Forest? For the most part, all fish
are aquatic, poikilothermic (a fluctuating internal body
temperature) vertebrates with gills. There are five spe-
cies of salmon that frequent the Pacific Northwest:
large Chinook (up to 50 kg); vibrantly striped Chum
or Dog; small and abundant Pink; Coho or Silver,
which spend the most amount of time in freshwater;
Sockeye, the ‘fish of fishes’, with the longest spawning
run and the ability to battle raging rapids head on, and
the fresh water variant Kokanee, which live their entire
life cycle in landlocked streams and lakes. The Salmon
Forest is defined geographically as the coastal forests

from northern California through to British Columbia
and Alaska, and across the Pacific to Russia, Japan and
Korea.

In chapter 2 we learn about the life cycle of the
salmon, and the biological traits that set these fish apart
and contribute to their special role in the forest. Two
of the defining traits of salmon are semelparity and
anadromy. Semelparous organisms die after one repro-
ductive cycle, and invest all of their metabolic energy
into the survival of their offspring. Anadromous fish
migrate upstream to spawn, either from marine to fresh-
water environments, or further upstream if they are ex-
clusively freshwater species. In salmon, this ensures
that valuable marine nutrients are brought into the
forest systems surrounding freshwater streams. These
really are fascinating fish — their ability to return to their
natal streams is particularly incredible, and it’s believed
that tiny particles of magnetite in their skulls act as a
sort of compass to guide them home, in a process called
olfactory homing — not dissimilar to the experiences
of migrating birds.

In chapter 3 we learn about how the very small — the
isotope signatures in salmon flesh — impact the envi-
ronment in a large way, leaving their mark throughout
the entire forest ecosystem. Since the development of
the mass spectrometer, scientists are able to trace a spe-
cies’ isotope signature through the environment. In this
case, salmon flesh leaves a distinct trace of marine-
derived nutrients, which has been found throughout the
Salmon Forest, from the needles of giant cedars to the
bones of coyotes.

Entitled ‘Salmon Gestalt’, chapter 4 covers several
concepts of ecological study, including the niche theo-
ry, trophic levels, food webs and keystone species. At
first it seemed unnecessary to discuss the history of
ecology, but by the end of the chapter it was interesting
to see how Stokes linked the study of salmon to the
study of all life.

After defining the salmon and how we might see its
signature throughout the Salmon Forest, Stokes uses
chapter 5 to look at the natural history of the diverse
flora and fauna that call this ecosystem home. Of par-
ticular interest are the examples of plant and insect phe-
nology that are tightly linked with the salmon life cycle,
such as the case of the blowfly and kneeling angelica
(Angelica genuflexa). The riparian angelica blooms ap-
proximately ten days after salmon spawning begins,
when adult blowfly are around to pollinate their flow-
ers. After feeding on the blooms, the blowflies lay their
eggs in the recently spawned salmon carcasses. Stokes
demonstrates how the forest is dependent on an inde-
scribably complex series of relationships between pred-
ator and prey, biogeochemical cycles and biotic/abiotic

422

interactions. There is simply no way to understand the
whole by prying apart the individual threads.

The concluding chapter brings us full circle, and
discusses the role of salmon as ecosystem engineers,
inevitably leaving their mark on every aspect of the
forest landscape in the Pacific Northwest, including the
people that call the region home. Although Stokes does
not go into great detail about recent ecosystem destruc-
tion, it is certainly implied that although resilient, these
species and this environment are nonetheless vulnerable
to our rapacious appetites for salmon, and our unfortu-

THE CANADIAN FIELD-NATURALIST

Vol. 128

nate habit for resource exploitation. It’s almost impos-
sible to quantify the importance of a keystone species
like the salmon, but according to Stokes, “if we truly
valued the intrinsic worth of the salmon as a part of the
Salmon Forest, perhaps we’d realize that we can’t
afford to exploit them — and we wouldn’t want to.”

I would certainly recommend this book to anyone
interested in natural history, and the beauty of the intri-
cate connections between a fish and its forest.

EMMA BOCKING

21 Avondale Avenue South, Waterloo, ON, Canada, N2L 2B5

Biology and Conservation of North American Tortoises

Edited by David C. Rostal, Earl D. McCoy, and Henry R. Mushinsky. 2014. The Johns Hopkins University Press, 2715 North
Charles Street, Baltimore, MD, USA, 21218-4363. 190 pages, 69.95 USD, Cloth.

The North American tortoises, or gopher tortoises,
are known for their digging ability. Most of the species
dig extensive burrows for shelter. These tortoises are
limited to the southern parts of the United States, from
California to Florida, as well as parts of Mexico, but all
of them face significant conservation challenges.

This book consists of 18 chapters on various aspects
of the biology and conservation of North American
tortoises. None of the chapters focus on individual spe-
cies. Instead, each chapter addresses what is known
about a topic for all of the species. Chapter topics in-
clude the fossil record, systematics, thermo-regulation,
reproduction, growth, health issues, habitat, movement
patterns, genetics and conservation. Over half of the
chapters have multiple authors, usually with experts
on different species collaborating on a topic.

It is difficult to name the five species covered in this
book, as the 33 authors could not agree on either the
common or scientific names. Until recently there were
four accepted species in one genus. The western Desert
Tortoise was split into two species, and it is possible
that further genetic analyses will reveal more species
within the group. Most of the authors of this volume
consider all of the North American tortoises to be in the
genus Gopherus, but a few consider that three of the
species should be placed in a different genus, Yero-
bates. Common names are also contentious. The two
Desert Tortoises are commonly referred to as the Mo-
have (or Mojave) Desert Tortoise and the Sonoran
Desert Tortoise but some herpetologists object to these
names, preferring Agassizi’s and Morafka’s Desert
Tortoises.

Overall, these chapters provide a thorough overview
of the biology of North American tortoises, including
research published up to the year 2012. For the most
part the chapters just summarize the scientific literature
on the five species, but a few chapters provide addi-

tional analysis of existing data. Some of the chapters
are vague on some topics. Climate change is mentioned
as likely resulting in more droughts which would likely
affect many of these species, but there is no attempt to
quantify such statements. Drought certainly is a signifi-
cant threat to tortoises as individuals can lose 40% of
their body weight during a severe drought. Longer or
more frequent droughts as a result of climate change
could lead to desertification and local extinctions.

Other threats are more clearly addressed. Invasive
exotic grasses are a significant issue in many western
areas, but since tortoises will eat both grasses and forbs,
it might be thought that exotic grasses pose little threat
to the tortoises. Forbs, however, are more nutritious
than grasses, so if plant communities become more
grass dominated this could be a serious threat to the
tortoises. In addition, many exotic grasses promote
severe wildfires that can kill tortoises.

The final chapter of the book provides a detailed
overview of the threats and conservation needs of these
tortoises. Despite protection of some populations dat-
ing back over 20 years, all these species are still declin-
ing. The Bolson Tortoise, limited to a small area of
northern Mexico, has even been listed as one of the
top 40 turtles or tortoises at risk of extinction. And
new threats continue to emerge. For example, frack-
ing potentially affects nearly 50% of the range of
Berlandier’s Tortoise in Texas. Not surprisingly then,
the major issues facing these species are adequate pro-
tection of populations and their habitat. This book suc-
cessfully summarizes the state of knowledge about
North American tortoises, but it is hard to be optimistic
for the future of these species.

DAVID SEBURN

Seburn Ecological Services, 2710 Clarenda Street, Ottawa.
ON, Canada, K2B 785

2014

Societies of Wolves and Free-ranging Dogs

BooK REVIEWS

By Stephen Spotte. 2012. Cambridge University Press, New York, NY, USA, 394 pages, 58.00 USD, Paper and Kindle,

133.00 USD, Cloth.

Stephen Spotte’s book, Societies of Wolves and Free-
ranging Dogs, is a detailed account of wild wolves and
free-ranging dogs (including dingoes) living through-

out the world. It is well researched with 73 pages of

endnotes, 50 pages of references, and a 16 page index.
The volume is 394 pages but ends on page 238 before
the citations noted above, and after 16 pages of Roman
numeral introductory material. The jacket cover of the
book accurately summarizes its contents: “Wolves are
charismatic emblems of wilderness. Dogs, which de-
scended from wolves, are models of urbanity. Do free-
ranging dogs revert to pack living or are their societies
only reminiscent of a wolfish heritage? Focusing on
behavioral ecology, this is the first book to assess soci-
eties of both gray wolves and domestic dogs living as
urban strays and in the feral state. It provides a com-
prehensive review of wolf genetics, particularly of New
World wolves and their mixture of wolf, coyote and
dog genomes. Spotte draws on the latest scientific find-
ings across the specialized fields of genetics, sensory
biology, reproductive physiology, space use, foraging
ecology and socialization. This interdisciplinary ap-
proach provides a solid foundation for a startling and
original comparison of the social lives of wolves and
free-ranging dogs.”

I am not sure if the manuscript is a startling com-
parison of wolves and dogs but it is certainly original
in how it combines the two canines into one volume. |
found it very odd though that Spotte, a well published
marine biologist, would write such a detailed book on
the Canidae family, considering that he does not appear
to have any prior publications on the topic and only
cites one of his previous references (on zoos) in this
book. That is not a negative comment, but rather more
of a curiosity on my part. However, I would have at
least liked for Spotte to have explained his decision
to write such a treatise, even if briefly discussed. He
certainly wrote the book with a different perspective
than a full-time canid biologist, as I’ll explain below.

I already have a fair bit of information (e.g., books,
scientific papers, personal research) on wolves and
coyotes, so in reading Societies of Wolves and Free-
ranging Dogs | learned more about dogs, both free-
ranging ferals and dingoes, than I previously had
knowledge of, and will likely use this book as a ref-
erence on them. I found it useful to have behavioural
information on city strays and feral dogs and to learn
about dingo ethology which seems to be dramatically
different than wolves and more of a mix of dog and
wolf, including their reproductive cycle and ecology
such as not living in packs and displaying questionable
parental care of offspring. However, it was pretty clear
that there isn’t nearly as much scientific information
published on dingoes and feral dogs as there is on

wolves, especially considering that most of the dingo
literature that was referenced was dated and much of
that was anecdotal (e.g., second hand reports of 1-2
sentences on some aspect of dingo ecology such as
parental care).

The book is a literal encyclopaedia of information
on what makes a wolf, what makes a dog, visual and
tactile communication, olfactory and vocal communi-
cation, space, foraging, courtship and conception, re-
production and parenting, and socialization. Spotte did
a commendable job of synthesizing knowledge on both
wolves and dogs which is an enormous task. I did find
many sections that were very “heavy” and lengthy to
read; for instance, the introduction to chapters, such as
Chapters 3.1 (p. 33-38) on metaphors and semiotics
and 4.1 (p. 61-66) on odour and pheromone reception,
were especially time consuming and philosophical
despite being information filled. Thus, the book was
more of a reference book than something to sit down
and casually read. I found that it took me multiple hours
to comprehend each of the 9 chapters and, as the author
suggested on p. xii, | had a bookmark in the endnotes
and reference sections. While I tried to simply peruse
through the book I found the scientist in me constantly
paging to the endnotes and then to the references to find
the quoted works. This took me considerable time bear-
ing in mind that each chapter had between 170-474
endnote citations. And herein lies my intrigue with the
book: Spotte cites many references (50 pages) but relies
disproportionately on older studies from the 1940s-
1970s. While many of these volumes are seminal stud-
ies, such as Adolph Murie’s (1944) The Wolves of
Mount McKinley, lan Cowan’s (1947) The Timber Wolf
in the Rocky Mountain National Parks of Canada, Lois
Crisler’s (1958) Arctic Wild, and Dave Mech’s (1970)
The Wolf, \ thought that the author cited them way too
much, especially considering the hundreds of new man-
uscripts which are more up-to-date and comprehensive
such as Dave Mech and Luigi Boitani’s (2003) Wolves:
Behavior, Ecology, and Conservation which was cited
much less frequently. However, to have 73 pages of
endnotes is a massive undertaking and there are newer
references mixed into those older works making the
book certainly usable and for the most part current.

I found Spotte’s perspective very interesting even
if I disagreed with some of his premises such as his
suggestion to name red wolves as “red coyotes” (p. 8).
calling Great Lakes wolves the same as eastern wolves
(page 7 note: most of the literature separates the two as
Great Lakes wolves are believed to be eastern wolf *
gray wolf hybrids), and his behaviorist viewpoint and
repeated mention of animals unable to recognize the
mental state of others (page 74) or that non-humans do
not possess a theory of mind (page 34). My guess is that

424

Marc Bekoff, and many other scientists that study ani-
mal cognition, would disagree with this perspective.
While Spotte casts doubt if admixed canids deserve
species status (p. 5), I disagree as new research is in-
forming us that hybridization is a natural process and
the cause of rapid evolution in many groups of animals,
such as canids in eastern North America, most notably
the eastern coyote/coywolf (Canadian Field-Naturalist,
2013, 127: 1-16).

Despite my comments noted above, I do endorse the
book as it is a valuable reference for anyone’s book
shelf and it provides a very well synthesized version of
both wolves and dogs. However, the following are my
four major criticisms of the book:

The first is simply the expense of book. When I
looked at the price on Amazon.com I was stunned how
expensive this simple soft-cover, nearly 400 page book
was. In fact, the sale price for the Kindle edition was
36.80 USD, and 52.20 USD for paperback, and 110.44-
126.35 USD for the hardcover version. I don’t under-
stand why the book is so expensive especially since it
is black and white and could be printed on demand for
significantly cheaper than that. The price alone makes
it difficult to justify purchasing except maybe for an
upper level college or graduate course.

Second and bizarrely, the book (page 238) just ends
after the dingo socialization section of Chapter 9 with
no conclusion whatsoever. I strongly believe there
should have been a Chapter 10 that synthesized the
information that I digested in the first 9 sections. With-
out this summary the reader is left wondering what the
main take home messages of the book are.

Third, Spotte is a tad critical of previous studies and
theories and is inconsistent with his criticisms. I think
this may have a bit to do with the author not having a
background studying canids even if he has a strong sci-
entific understanding. There were many sections of the
book that I just didn’t feel right with but sometimes
couldn’t place my finger on exactly why I felt that way
because I was trying to trudge through all of the infor-

BOTANY

THE CANADIAN FIELD-NATURALIST

Vol. 128

mation (this would probably be similar to me writing
a detailed book on marine mammals and then having
a marine biologist review it). For instance, on page 67
Spotte claims that wolf scent marking as a territorial
boundary is more anecdote than empirical yet he then
repeatedly makes claims and references in the end-
notes (especially some of the older citations noted
above) supporting such behavior. I found it difficult
to go back and forth from the chapter to the endnotes
when reading these types of passages, especially where
there is much scientific agreement on a given topic
like canid territoriality and scent marking. In other
words, the author is critical of the scientific literature
in many sections (using philosophical arguments of
why things like territoriality aren’t proven) yet he then
attributes many of his statements with older citations
that may even be anecdotes themselves (like the books
published in the 1930s-1950s).

And fourth, the book is repetitive in many sections,
but this isn’t a major complaint. While this may be ex-
pected of a major work I found the frequent references
to go back to a specific chapter, reference, or endnote
frustrating. But that was most likely because I was read-
ing the book from cover-to-cover instead of using it as
an encyclopedia-like reference whereby that repetition
may be useful to the reader.

Despite some of my issues with Societies of Wolves
and Free-ranging Dogs it is certainly a unique refer-
ence on wolves and dogs, which are now generally
regarded as the same species. In one book, it synthe-
sizes what is known on the most recognizable of all
canids and I don’t think that has been done before.
Thus, despite me disagreeing with certain sections and
viewpoints, I would recommend the book as a refer-
ence for canid enthusiasts if one can afford the price.
However, the book is not for beginners on the subject
matter of wolf and dog ecology and behavior as it is a
time-consuming and exhaustive read.

JONATHAN WAY
89 Ebenezer Road, Osterville, MA, USA, 02655

Plantes de milieux humides et de bord de mer du Québec et des Maritimes

By Martine Lapointe, Michel Leboeuf, and Arold Lavoie. 2014. Guides nature Quintin, 4770, rue Foster, Waterloo, QC, Canada,

JOE 2NO0. 455 pages, 34.95 CAD, Cloth.

This latest edition in the Michel Quintin nature
guides is a visually pleasing publication having numer-
ous colour photos of plants and of wetland habitats,
crisp line drawings and illustrative graphics. It covers
302 species of flowering plants, ferns, horsetails, club-
mosses as well as hepatics and mosses. The guide is
stated as being aimed at nature lovers as well as spe-
cialists undertaking wetland and seashore plant inven-
tories within Quebec and the Maritimes. However, it is

likely better considered as a good introductory guide
to the flora and vegetation of these habitats for avid nat-
uralists and students of ecology. Anyone engaged ina
comprehensive documentation of the vegetation of
wetland habitats would, by necessity, require a more
detailed floristic reference work. This would certainly
be the case to differentiate morphologically similar spe-
cies in the more difficult groups such as the sedges.
grasses, and rushes common in these habitats,

2014

The initial section of the guide provides an excellent
introduction to wetland types and their relationship to

water levels and vegetation present through the use of

illustrative graphics and photos. A very helpful feature
of the guide is the listing of locations of representative
examples of wetland habitats, along with colour photos,

of specific marshes, swamps, bogs and various types of

seashore habitats. The introduction to flowering plants
provides an oversight of the diversity found in vegeta-
tive and floral structures through drawings that illus-
trate the range in leaf types and form and of inflores-
cence structure. These illustrated glossaries aid the
non-specialist in understanding the technical descrip-
tive terminology used in the guide. At the same time, it
permits the author to provide more precise descriptions
of plant form within the individual species treatments.

The species covered are grouped into seven cate-
gories readily identified by the colour coding on a por-
tion of the unbound side and corner of the guide: trees
and shrubs; terrestrial plants; aquatic plants; maritime
plants (wet habitats, dry habitats); sedges and members
of the sedge family, as well as grasses and rushes; ferns,
horsetails, and clubmosses; hepatics and mosses. The
identification of species is based initially on the place-
ment of a plant in question into one of the seven cate-
gories. Subsequent identification to species is based
in large part on a comparison of vegetative and floral
structures exemplified in the colour photos and line
drawings. For herbs and some shrubs with distinctive
flowers or flower clusters, a colour gallery of flowers,
grouped by colour, is provided for 155 species as an
identification aid. Within difficult groups of flowering
plants, such as the sedges, grasses, and rushes, an intro-
ductory set of colour graphics help differentiate these
groups. Similarly for ferns and allies and the hepatics
and mosses, graphics of their growth form and struc-
tures are provided.

Each individual species treatment includes several
elements. The visual portion includes a colour photo
with the main distinguishing features of the plant ac-
companied by a line drawing with details of appropri-
ate vegetative and floral/fruiting structures. Colour
inserts of floral or vegetative details within the larger
plant images of some species are not always helpful
due to their small size. Additional visual aids are pro-
vided by symbols that provide further information on
the biology or usefulness of the plant: whether the
plants are obligate or facultative wetland species; light
requirements; longevity (perennial, biennial, or annu-

Book REVIEWS 425

al); use in re-vegetation of riparian banks, and whether
the plants are invasive.

The main descriptive texts include the French com-
mon name in bold typeface followed by the scientific
name, English common name and the family name.
Organization of the species within each section is al-
phabetical by the French common name. Descriptive
texts for each species cover such aspects as size and
habit, branching/bark/leaves, flowers, fruits, habitat and
distributional range. Sections on similar species or
notes on such aspects as toxicity or uses are also pro-
vided. In the texts for ferns and allies and for hepatics
and mosses, information on their distinctive reproduc-
tive structures is provided.

The characteristics of each wetland species of south-
ern Quebec, treated in the guide, are summarized in a
table organized alphabetically by scientific name fol-
lowing the species treatments. The table summarizes
and codifies species on such aspects as whether they are
obligate or facultative wetland species, and for those
obligates, whether they are species always found root-
ed in water or whether they are true aquatics, partially
or totally submerged. The species are further identi-
fied as to their growth form (e.g., tree, shrub, herb) or
type of aquatic (emergent, submerged etc.), and spatial
placement within the littoral zone. Each species is
also assessed as to its level of threat and whether it is
an invasive exotic or if useful for revegetation purpos-
es. The table is perhaps most useful to students of wet-
land ecology, with the information on level of threat
(threatened, vulnerable or possibly at risk of decline)
being of more general interest to naturalists. A glossary
of terminology, a bibliography, a list of useful inter-
net sites, credits, and an index to species complete
the guide.

The guide is a very pleasing and useful tome for
gaining a basic understanding of the plants and vege-
tation of wetlands. Its appeal is based to a large extent
on the high quality of the paper, illustrations, and pho-
tos, the informative graphics on wetland composi-
tion, the useful explanatory texts, and concise techni-
cal species treatments. It is a relatively hefty guide
(about 0.8kg) for its modest size of 13 by 20 cm. Be-
cause of this, it might be best used as a reference tool
at the roadside while exploring habitats referenced in
the guide, rather than as a pocket field guide, carried
along while slogging through a wetland.

ERICH HABER
60 Baywood Drive, Stittsville, ON, Canada, K2S 2H5

426

Trees of Eastern North America

THE CANADIAN FIELD-NATURALIST

Vol. 128

By Gil Nelson, Christopher J. Earle, and Richard Spellenberg. 2014. Princeton University Press, 41 William Street, Princeton,

NJ, USA, 08540-5237. 720 pages, 29.95 USD, Paper.

There are several field guides available for eastern
North American trees. One of the more recent is the
Sibley guide (Sibley 2009), the National Wildlife Fed-
eration field guide (Kershner ef a/. 2008), and the Peter-
son guide (Petrides 1988). The Petrides (1988) guide
covers roughly the same geographic area as this guide
whereas the other two are US and Canada. Nelson and
Spellenberg are also listed as authors of the Kershner
et al. (2008) guide.

Back to the Nelson ef a/. guide that is the subject
of this review. This guide which is a hefty 720 pages,
covers virtually all species of tree expected to be found
in the eastern part of North America which is roughly
defined as the Northwest Territories/Nunavut boundary
and a line extending southward and slightly east of the
Manitoba/Saskatchewan border continuing southward
along or near the 100" meridian until it enters Texas.
In Texas the line traces along the eastern edge of the
Edwards Plateau to Austin and then south ending at
the Gulf of Mexico at Corpus Christi. In contrast, the
Peterson guide (Petrides 1988) considers all of the trees
east of the Black Hills and Rocky Mountains. Thus,
coverage is a bit different.

The present guide includes virtually all trees that
“grow without the aid of human cultivation” (page 7)
within the above-described area. This includes a num-
ber of exotic species that have escaped cultivation,
which is a lot of species considering the guide encom-
passes all of Florida where many exotics have been
planted as ornamentals. Also, because it includes a por-
tion of south Texas, species of a more southwestern dis-
tribution are included. This species coverage is very
similar to the Sibley (2009) guide except that Sibley
includes many ornamental trees that are not at present

OTHER

known to naturally reproduce independent of human
cultivation.

Of the alternative guides mentioned, this guide has
up-to-date classification, and is reasonably compact so
that it can be used as a field guide. The Sibley (2009)
guide is not of a convenient size to carry on field trips.
The quality of illustrations is far better than the other
guides mentioned so that identification is easier. This
includes the distribution maps which are sized to be
clear. The text is error-free and no other editorial issues
were noted while reading.

This guide can be recommended for each of the
above reasons. It adds a very favorable choice to the
selection of eastern tree guides presently available and
is mid-way in price among each of the guides men-
tioned in this review. If you wish a new guide, an alter-
native guide to supplement others, or an off-the-shelf
tree reference, I recommend this book for that purpose.
I would also not hesitate to use this guide as a text for
identification in a course of dendrology or any class
involving plant identification. Foresters, horticulturists,
wildlife managers, and other professionals will also find
it usable. It adds to the many high-quality field guides
now being published by Princeton University Press.

Literature Cited

Kershner, B., D. Mathews, G. Nelson, R. Spellenberg, T. Purinton,
A. Block, G. Moore, and J. W. Thieret. 2008. Field guide to trees
of North America. New York, NY, Sterling.

Petrides, G. A. 1988. A field guide to eastern trees. New York, NY,
Houghton Mifflin.

Sibley, D. A. 2009. The Sibley guide to trees. New York, NY, Knopf.

ROGER D. APPLEGATE
57 Benzing Road, Antioch, TN, USA, 37013

The Real Weed Man: Portrait of Canadian Botanist Gerald A. Mulligan

By Julie Mulligan. 2014. Published by Julie Mulligan, P.O. Box 304, Russell, ON, Canada, K4R 1E1. 238 pages, Paperback
edition 16.84 CAN, Kindle edition 2.95 CAN (Amazon.com).

The Real Weed Man is a daughter’s very personal
account of her father’s life and career at the Central
Experimental Farm (CEF) in Ottawa. It is based on dis-
cussions with her father, family members, colleagues,
and on entries from Gerry’s daily diary begun in 1974,
which are frequently quoted. Within the book’s 25
chapters, beginning with the early years growing up in
Ottawa, the author traces Gerry’s association with the
“Farm” over a period covering more than 70 years.
Gerry followed in the footsteps of other family mem-
bers of earlier generations who worked at CEF. He
began as a summer student, then as a technician after

High School, and subsequently, following the comple-
tion of his Bachelor of Science degree in 1952 from
Macdonald College of McGill University, he was hired
as a biologist. Through considerable perseverance and
his abilities as a researcher and manager, Gerry rose
through the ranks to reach the top level of the Research
Scientist classification. Gerry eventually also served as
the Director of the Biosystematic Research Institute at
CEF from 1978 to 1987.

The author touches on many facets of Gerry’s per-
sonality and career. We learn that Gerry is an avid
sports, and especially hockey, fan, a devoted father and

2014

husband, as well as a gifted researcher and manager.
Gerry is a very disciplined individual and persistent
in achieving his goals. He has always tried to be ethi-
cal in his approach to decision-making as a manager
and was open to heading in new directions when so
required. The biographical sketch deals not only with
aspects of Gerry’s personal traits, but also reveals, over
a series of chapters, how he dealt with or views specific
issues, either as a researcher or as a manager. These
include such topics as health and safety in the work-
place, annual staff appraisals, French in the public serv-
ice, and genetically modified plants, among others.

Readers who are familiar with Ottawa, would be
interested in the details on historical land ownership
by the Mulligan family within the city and at CEF.
Changes that have occurred over the years at CEF, in
the focus of its operations and in staffing, are touched
on as well.

The most surprising aspect of the book is the frank-
ness of revelations on Gerry’s interactions with col-
leagues and managers, or of his views about the pub-
lished works of others specifically named in the book.
Because of such anecdotes, the book would be of par-
ticular interest to many botanists and biologists.

The primary subject of the book is that of a botanist,
a weed specialist, Gerald Alfred Mulligan, his strengths
and possible weaknesses, his lifestyle and views. In
this regard, I have some minor quibbles on content.
The author tends to expand topics such as Gerry’s life-
long stuttering into a long, personal and family account.
This, for the reviewer, digresses too far from its rele-
vance to Gerry’s condition and its impact on his career.
Similar, perhaps excessive tangents, dealing with fam-
ily outings and incidents, related specifically to the

Book REVIEWS

427

author and her siblings, are included rather than remain-
ing focused on Gerry’s fieldwork activities. The chap-
ter on photography might have been better treated under
a different subject heading. Much of it highlights prob-
lems in maintaining a web site on Weeds of Canada
rather than on Gerry’s photography. The author also
regularly inserts lengthy personal views rather than
reflecting those of her subject.

Considering that this biography is likely of greatest
general interest to botanists and biologists, the detailed
definitions and discussion of botanical “type” collec-
tions seem unnecessary. It is unfortunate that the error
in naming of the Gray Herbarium of Harvard Univer-
sity (GH), given as the “Great Herbarium of Harvard
University” (page 50 and in footnote page 199), was
not recognized. Similarly, the institutional citation for
the herbarium known by its acronym CAN is given as
“ Canadian National Museum herbarium” rather than as
“Canadian Museum of Nature”. The updated names of
institutions and the acronyms of their herbaria are read-
ily found on the internet by searching on Index Herbar-
iorum.

In spite of the above critique, the book is an interest-
ing read. It provides a very detailed perspective of a
Canadian botanist who is known widely for his work
on the mustard family, his numerous plant chromosome
counts and his documentation of the weeds of Canada.
Gerry’s life-time work and achievements were recog-
nized by the Canadian Botanical Association in 2006
when it presented him with its prestigious George Law-
son Medal “...for excellence in the contribution of an
individual to Canadian botany”.

ERICH HABER
60 Baywood Drive, Stittsville, ON, Canada, K2S 2H5

The Once and Future World: Nature as it was, as it is, as it could be

By J. B. MacKinnon. 2013. Random House Canada, One Toronto Street, Unit 300, Toronto, ON, Canada, MSC 2V6. 256 pages,

29.95 CAD, Cloth.

The Once and Future World by J. B. MacKinnon,
co-author of The 100-Mile Diet: A Year of Local Eat-
ing, is an eloquent and thought-provoking exploration
of nature and our place in it. It addresses the roles
humans play within the natural world, the influence we
exert on its complex systems, and the relationships we
develop with our natural surroundings. It really is a
book about Homo sapiens in nature.

The book is also an invitation. MacKinnon invites us
to stretch our thinking and our imaginations beyond
common knowledge to envision a different world and
future. He does so by blending personal narrative, SCI-
entific fact, historical discovery, recent events, case
studies, speculation about the future, and sweeping
questions that trigger reflection on where we are today
and what we would like the world, and the place of

humans in it, to look like. Along the way, McKinnon
discusses a wide range of concepts, including the an-
thropocene, change blindness, shifting baselines, cas-
cade effects, ecosystem services, relaxed selection,
progress trap, double extinction, re-wilding, habitec-
ture, and more.

“Nature is a confounding thing” he writes in the first
chapter (page 8), illustrating throughout the book how
our relationship with and within nature is equally per-
plexing. He offers particularly intriguing perspectives
on the past — a past which is itself confounding and
imperfect, filled with drama, story, extinction, human
‘progress’, forgetting and denial, resulting in what he
calls a “10 percent world” offering only one-tenth of its
former natural abundance. Accompanying the impov-
erishment of natural abundance is the loss of relation-

ships, knowledge and cultural practices relating to the
natural world —a loss of nature from human social net-
works, as he puts it.

Yet this loss and gradual impoverishment was not a
deliberate trajectory, but rather an incremental “taking”
from the environment by different arrangements of peo-
ple in different places at different points in history. All
these ‘takings’, McKinnon adds, occurred with limited
approval to degrade the environment, and subsequent
adaptation to the consequences. Now, after many mil-
lennia, the accumulated impact of the consequences
and adaptations is becoming increasingly clear.

Yet it is still a beautiful world, as the title of Chapter
5 suggests, and nature has a life and a vitality all its own
—“always expressing itself in a weed poking up from a
crack in the concrete” (page 9) — which shows us a way
forward despite challenges which appear overwhelm-
ing. McKinnon identifies one major challenge as a lack
of awareness regarding the natural world which sur-
rounds us — from weeds in sidewalk cracks, to birds of
prey hunting in urban parks, to ocean life returning to
the waters of a popular beach after the holiday season.
The nature we surround ourselves with shapes us as a
species, he points out, adding that we tend to prefer
wilder, more diverse environments. In that light, an
important and overlooked result of conservation and
ecological restoration efforts may be the recovery of a
certain wildness within ourselves. In the context, fur-
thermore, of challenges and limitations faced by na-
ture conservation and ecological restoration — in world
where, as McKinnon points out, many people spend

THE CANADIAN FIELD-NATURALIST

Vol. 128

more time in virtual landscapes than natural ones — this
‘human re-wilding’ of sorts, and deliberate reconnec-
tion and cohabitation with wild species, may be more
important that we realize.

McKinnon offers examples of human-nature recon-
nection and co-existence from different parts of the
world. He devotes most of the entire last chapter of the
book (excluding the epilogue), to an imaginary exam-
ple illustrating how humans can live in a wilder world.
McKinnon ends that chapter, and essentially the book,
in echo of an earlier passage where he encourages us
to “remember what nature can be; reconnect with it as
something meaningful in our lives; and start to remake
a wilder world” (page 157). He ends the book on a
tremendously positive note, pointing out that “it just
might be possible for seven billion people — maybe
even more — to survive on this planet, and not only to
stop the endless decline of the natural world but watch
it return to astounding, perpetual life. All it will take is
a wilder way of being human” (page 215).

It is difficult to do The Once and Future World jus-
tice in a book review. This rich, informative and opti-
muistic book defies categorization and description. Let
me just say that I recommend it highly for being well-
researched, well-written, thought-provoking, imagina-
tive, and compelling. May it compel readers toward a
wilder future where, as McKinnon puts it, humanity
can express its genius, and so can nature.

RENATE SANDER-REGIER
3, 11th Line, Bristol, QC, Canada

2014

NEw TITLES

Prepared by Roy John

+ Available for review * Assigned

NEw TITLES

429

Currency Codes — CAD Canadian Dollars, USD U.S. Dollars, EUR Euros, AUD Australian Dollars.

ZOOLOGY

* Field Guide to the Birds of Australia — A Photo-
graphic Guide. By lain Campbell, Sam Woods, and
Nick Leseberg. 2014. Princeton University Press, 41
William Street, Princeton, NJ, USA, 08540-523. 392
pages, 35.00 USD, Paper.

Birds of a Feather. By C. Rees, and D. Thomas. 2014.
Troubador Publishing Ltd, 9 Priory Business Park,
Kibworth, Leicester, UK, LE8 ORX. 200 pages, 12.99
GBP, Paper.

The Carnivore Way — Coexisting with and Consery-
ing North America’s Predators. By Cristina Eisen-
berg. 2014. Island Press, 2000 M Street NW, Suite 650,
Washington, DC, USA, 200362014. 328 pages, 30.00
USD, Cloth.

Ducks, Geese, and Swans of North America —
revised. By Guy Baldasarre. 2014. The Johns Hopkins
University Press, 2715 North Charles Street, Baltimore,
MD, USA, 21218-4363. 1,088 pages, 69.95 USD,
Cloth.

Fishes: A Guide to Their Diversity. By Philip A. Hast-
ings, H. J. Walker Jr., and Grantly R. Galland. 2015.
University of California Press, California-Princeton
Fulfillment Services, 1445 Lower Ferry Road, Ewing,
NJ, USA, 08618. 336 pages, 65.00 USD, Cloth.

* An Indomitable Beast: The Remarkable Journey
of the Jaguar. By Alan Rabinowitz. 2014. Island Press,
2000 M Street NW, Suite 650, Washington, DC, USA,
20036. 304 pages, 30.00 USD, Cloth.

Marmot Biology — Sociality, Individual Fitness, and
Population Dynamics. By K. Armitage. 2014. Cam-
bridge University Press, 32 Avenue of the Americas,
New York, NY, USA, 10013-2473. 405 pages, 120.00
USD, Cloth.

The Passenger Pigeon. By Errol Fuller. 2014. Prince-
ton University Press, 41 William Street, Princeton, NJ,
USA, 08540-5237. 184 pages, 29.95 USD, Cloth.

Pilgrims of the Air. By John W. Foster. 2014. Notting
Hill Editions, Widworthy Barton, Widworthy, Devon,
UK, EX14 9JS. 240 pages, 25.00 USD, Cloth.

* Penguins — The Ultimate Guide. By Tui De Roy,
Mark Jones, and Julie Cornthwaite. 2014. Princeton

University Press, 41 William Street, Princeton, NJ,
USA, 08540-5237. 240 pages, 35.00 USD, Cloth.

RSPB Seabirds. By Marianne Taylor. 2014. Blooms-
bury Publishing PLC., 50 Bedford Square, London,
UK, WC1B 3DP. 240 pages, 25.00 GBP, Cloth.

Woodpeckers of the World. By G. Gorman. 2014.
Christopher Helm-Bloomsbury Publishing PLC., 50
Bedford Square, London, UK, WC1B 3DP. 528 pages,
35.00 GBP, Cloth.

BOTANY

Common Lichens of Northeastern North America
— A Field Guide. By Troy McMullin, and Frances
Anderson. 2014. Koeltz Scientific Books, P.O. Box
1360, D-61453 Koenigstein, Germany. 194 pages,
49.00 EUR or 63.70 USD, Paper.

* Sedges of Maine: A Field Guide to Cyperaceae
Paperback. By Matt Arsenault, Glen H. Mittelhauser,
Don Cameron, Sally C. Rooney, and Jill E. Weber.
2013. The University of Maine Press, 126A College
Avenue, Orono, ME, USA, 04473. 712 pages, 29.50
USD, Paper.

OTHER

Experimental Evolution and the Nature of Biodi-
versity. By Rees Kassen. 2014. Roberts and Company
Publishers, 4950 S. Yosemite Street, F2 #197, Green-
wood Village, CO, USA, 80111. 288 pages, 36.00 USD,
Paper.

The Analysis of Biological Data, Second Edition.
By Michael Whitlock, and Dolph Schluter. 2014.
Roberts and Company Publishers, 4950 S. Yosemite
Street, F2 #197, Greenwood Village, CO, USA, 80111.
864 pages, 99.00 USD, Cloth.

* Climber’s Paradise — Making Canada’s Mountain
Parks, 1906-1974. By Pearl A. Reichwein. 2014. Uni-
versity of Alberta Press, Ring House 2, Edmonton, AB,
Canada, T6G 2E1. 432 pages, 45.00 CAD, Paper.

Conservation Science: Balancing the Needs of Peo-
ple and Nature, Second Edition. By Peter Kareiva,
and Michelle Marvier. 2014. Roberts and Company
Publishers, 4950 S. Yosemite Street, F2 #197, Green-
wood Village, CO, USA, 80111. 672 pages, 72.00 USD,
Paper.

430

A Field Guide to Economics for Conservationists.
By Brendan Fisher, Robin Naidoo, and Taylor Rick-
etts. 2014. Roberts and Company Publishers, 4950 S.
Yosemite Street, F2 #197, Greenwood Village, CO,
USA, 80111. 208 pages, 36.00 USD, Paper.

* Conrad Kain — Letters from a Wandering Moun-
tain Guide, 1906-1933. By Conrad Kain, and Zac
Robinson et al. 2014. University of Alberta Press, Ring
House 2, Edmonton, AB, Canada, T6G 2E1. 472 pages,
34.95 CAD, Paper.

THE CANADIAN FIELD-NATURALIST

Vol. 128

* Ian McTaggart-Cowan -The Legacy of a Pioneer-
ing Biologist, Educator and Conservationist. Edited
by Wayne Campbell, Dennis A. Demarchi, and Ronald
D. Jakimchuck. Harbour Publishing, P.O. Box 219,
Madeira Park, BC, Canada, VON 2HO0. 320 pages,
49.95 CAD, Cloth.

* The Real Weed Man: Portrait of Canadian Bota-
nist Gerald A. Mulligan. By Julie M. Mulligan (author
and publisher; available from Amazon.com). 2014.
228 pages, 19.99 USD, Paper, e-book 2.99 USD.

News and Comment

The Canadian Herpetologist (TCH) 4(2), Fall 2014

The Canadian Herpetologist (TCH) is a publication pro-
duced twice each year by the Canadian Association of Her-
petologists and the Canadian Amphibian and Reptile Conser-
vation Network.

CONTENTS: CHS Executive Members — Instructions for
Authors — Editorial Notes —Meetings— Feature Articles:
Unveiling of an Educational Panel Commemorating Over
40 Years of Turtle Research in Algonquin Provincial Park
by P. Moldowan, J. Litzgus and R. Brooks; Nest Site Over-
wintering by Northern Map Turtle (Graptemys geographica)
Hatchlings and Nest Site Protection at a Major Nesting Site in
Quebec by D. Fournier — Field Notes: An Overview NCC’s
Conservation Efforts for Endangered Herpetofauna in the
Ottawa Valley (Quebec) by M. Saumur; Calgary Zoo Centre
for Conservation & Research 2014 Fieldwork Update by L.
Randall; Fall Update on the Ojibway Massasauga Recovery
Program by J. Choquette; First Northern Map Turtle (Grapte-

Upcoming Meetings and Workshops

Northeast Natural History Conference 2015

The 15" Northeast Natural History Conference (NENHC)
hosted by The Association of Northeastern Biologists to be
held 18-20 April 2015 at the Sheraton Springfield Hotel in
Springfield, MA. This is a 3 day conference which promises
again to be the largest forum for researchers, natural resource
managers, students, and naturalists to present current informa-
tion on the varied aspects of applied field biology (freshwater,
marine, and terrestrial) and natural history for the Northeast-
ern United States and adjacent Canada. Registration is cur-
rently open. More information is available at http://www.eagle
hill.us/NENHC_2015/NENHC2015.shtml.

William Beverly Scott 1917-2014

An outstanding Canadian ichthyologist died 18 August
2014. William Beverly Scott (Bev) was born 7 July 1917 and
raised in Toronto. A fter serving in the Canadian army in Europe
in WWII he obtained his PhD at the University of Toronto in
1950 and was appointed Curator of Fishes at the Royal Ontario
Museum with a cross-appointment to the Department of Zool-
ogy, University of Toronto. In 1950-1 976 he built the research
fish collection at the ROM to be the largest in Canada and it has
continued to grow under E. J. Crossman, A. Emery, R. Win-
terbottom and H. Lopez-Fernandez. It now contains 97,600
cataloged lots containing a little over 1,1 27,636 specimens rep-
resenting 6,677 valid species in 415 families (Erling Holm: 27
November 2014). For contrast, the next largest Canadian col-
lection is held by the Canadian Museum of Nature has 60, 828
catalogued lots for a total of 679,522 specimens representing
4,025 species (Sylvie Laframboise: 28 November 2014).

In 1976, Bev moved to St. Andrews, New Brunswick, where
he became Director of the Huntsman Marine Science Centre,
a laboratory run by a consortium of Canadian Universities.
There he established the Atlantic Reference Center, one of
North America’s largest collections of larval fishes. After Bev’s
retirement from the Huntsman in 1982, he served as a Hunts-
man Senior Scientist, even after moving to Kingston, Ontario

in 1998.

While with the ROM, Bev coauthored Freshwater Fish of

Canada (1973) with E. J. Crossman (Ed) who had come to the
ROM in 1957 after obtaining a MSc from the University of
Toronto and a PhD from the University of British Columbia.
Ed succeeded Bev as Curator of Fishes. He died in December

mys geographica) Capture in Algonquin Provincial Park by
P. Moldowan, Record Size Female Blanding’s Turtle (Emy-
doidea blandingi) Found in Algonquin Provincial Park by P.
Moldowan; Toad Tales from Long Point, Ontario (PART 1)
by K. Yagi; Toad Tales from Long Point, Ontario (PART 2)
by F. Papini — Thesis Abstracts in Canadian Herpetology:
Moldowan, P.D. M.Sc. 2014. Laurentian University, Sudbury,
Ontario. (Supervisor: J.D. Litzgus). Sexual dimorphism and
alternative reproductive tactics in the Midland Painted Turtle
(Chrysemys picta marginata) — Crosby, J. MES. 2014. Uni-
versity of Waterloo, Waterloo, Ontario. (Supervisors: S. Mur-
phy and S. Ashpole). Amphibian occurrence on south Okana-
gan roadways (2010-2012): Investigating movement patterns,
crossing hotspots and roadkill mitigation structure use at the
landscape scales. — Recent Publications in Canadian Her-
petology — News and Announcements — CHS/SHC Mem-
bership Form.

Annual Conference of the Animal Behavior Society
2015

The 52™ Annual Conference of the Animal Behavior Soci-
ety to be held 10-14 June 2015 in Anchorage, Alaska. More
information is available at http://abs2015.org/.

International Conference on Biodiversity 2015

The 4" International Conference on Biodiversity to be held
15-17 June 2015 in Las Vegas, USA. The theme: “Share and
Enhance Ecological & Geological Conservation research”.
Registration is currently open. More information is available
at http://biodiversity.conferenceseries.com/index.php.

2004 and an often discussed revision never materialized. Still
at the ROM, Bev completed a monograph Fishes of the Atlantic
Coast of Canada (1966) started earlier by A. H. Leim, but left
unfinished at his death. Subsequently at Huntsman, as more
information accumulated for the region, a completely new vol-
ume was undertaken by Bev with his wife, Milly, as coauthor:
Atlantic Fishes of Canada (1988). Bev and Milly moved to
Kingston when Milly’s health deteriorated and she died there
in 2006. Bev subsequently moved to Lindsay, Ontario, where
both his children live, but remained in contact with colleagues
at the ROM and elsewhere.

Bev frequently served The Canadian Field-Naturalist, as he
did for other scientific journals, as a reviewer of ichthyology
papers. His last contribution was a review of a book by long-
time ROM colleague Glenn B. Wiggins: Biological Notes on
an Old Farm: Exploring Common Things in the Kingdoms of
Life. Canadian Field-Naturalist 123(4): 387.

Appreciation for contributions and corrections to an earlier
draft are due (by institution) Erling Holm, Rick Winterbottom,
Ross MacCulloch (Royal Ontario Museum) Sylvie Lafram-
boise, Brian Coad (Canadian Museum of Nature), C. G. Gru-
chy (retired, formerly Canadian Museum of Nature), and Lou
Van Guelpen (Curator of Fishes and Collections manager, At-
lantic Reference Centre The Huntsman Marine Science Cen-
tre). Also consulted was the tribute based on an interview by
David G. Smith,; William Beverly Scott Copeia 2006: 307-315,
and a DFO Science and Research feature on-line article:
Not the Retiring Kind: Scientist emeritus Bev Scott.

FRANCIS R. COOK

43]

The Ottawa Field-Naturalists’ Club Awards for 2013, presented April 2014

ELEANOR ZURBRIGG, IRWIN BRODO, JULIA CIPRIANI, CHRISTINE HANRAHAN, AND ANN MACKENZIE

On April 26'", 2014 members and friends of the Ot-
tawa Field-Naturalists’ Club gathered for the Club’s
Awards Night at St. Basil’s Church in Ottawa to cele-
brate the presentation of awards for achievements in
the previous year. Awards are given to members or
non-members who have distinguished themselves by
accomplishments in the field of natural history and

Member of the Year: Mark Brenchley

The OFNC’s Member of the Year award recognizes
the member judged to have contributed the most to
the Club in the previous year.

We are recognizing Mark Brenchley for his positive,
creative and enthusiastic commitment. Mark Brenchley
joined the OFNC in 2012 upon his return to Ottawa.
In late 2012 and throughout 2013 he jumped in to sup-
port the Education and Publicity Committee. His ener-
gy, skills and ideas were pivotal to the completion of
several projects designed to enhance the profile of the
Club.

When Ontario Nature invited naturalist clubs to spon-
sor a youth to attend the Ontario Nature Youth Summit
last year, the request landed in the hands of the Edu-
cation and Publicity Committee. Mark stepped up to
design an application process to select a candidate to
represent the Club. He communicated with school
boards, scouting groups and the members of the Club
to generate interest in the opportunity. Three applica-
tions came back to him. Mark, Suzanne Deschénes and
Lynn Ovenden interviewed the candidates and selected
Sarah Wray, a grade 10 student at Nepean High School.
After the Summit, Mark invited Sarah to describe her
experience at a monthly meeting in the Fall and asked
her to write a report which was published in the 2014
Winter issue of Trail and Landscape.

Had Mark not taken the Ontario Nature initiative,
the club would have missed an opportunity to support
the Summit and Sarah. Thanks to his initiative, there is
a framework in place to support the Club’s future par-
ticipation.

Over the year, Mark worked on a series of promo-
tional items and signs, all bearing the OFNC’s owl
logo, the club name and often the website on a forest

conservation or by extraordinary activity within the
Club. Three awards were presented for 2013, recog-
nizing achievement in enhancing the profile of the
Club, long time service as editor for one of the Club’s
publications, and success in raising awareness of envi-
ronmental stewardship and respect for nature in the pri-
mary school program.

green background. He made an 8' long banner that tops
our tent and display table at public events. When Edu-
cation and Publicity wanted to offer hand lenses on a
club lanyard at the monthly meeting sales table, Mark
created the logo file and placed the order with the
lanyard imprinting company. Then he coordinated the
logo design and purchase of the lens wipe pouches
which clip onto a lanyard or binocular strap. He led
the design of a promotional bookmark which includes
photos and the Club website.

The OFNC Council realized a few years ago that
the sign at the entrance to Fletcher Wildlife Garden is
a highly visible spot, a great place to increase public
awareness of OFNC. Although there has been a sign on
Prince of Wales Drive to direct people to the Garden,
there was nothing to indicate the OFNC’s involvement
with the Garden. Since Mark is in the sign business, he
was asked what could be done. As a result he designed
and installed a dark green aluminum panel with the
OFNC owl and our website atop the existing Fletcher
sign clearly identifying the Fletcher Wildlife Garden
with the OFNC, all at minimal cost to the Club.

Through these products Mark has developed several
graphic files with the OFNC’s logo and wordmark. He
continues to work at “branding” the OFNC with the
goal to create a recognizable visual identity for all Club-
related materials, promotional items and signage.

Mark is a cheerful, engaging salesman behind the
display table at most monthly meetings and at some
of OFNC’s public events. It is for his many contribu-
tions to promoting the Club in 2013 that we want to
recognize Mark as Member of the Year.

(Written by Julia Cipriani based on information
and feedback from Lynn Ovenden)

George McGee Service Award: Karen McLachlan Hamilton

The OFNC George McGee Service Award is given in
recognition of a member who has contributed signifi-
cantly to the smooth running of the Club over several
years.

This year, the award goes to Karen McLachlan
Hamilton, long-serving editor of Trail & Landscape.

All club members are familiar with our quarterly pub-
lication, more commonly known as T&L, in which news
of the club, regional biological and conservation reports
and articles by members, lists of upcoming events,
poems, photos and much else is published. Every three
months, this lively and informative publication arrives

432

2014

in our mailboxes, to be enjoyed and savoured. | would
guess, however, that many of us give little thought to the
work and effort that ensures 7&L lands in our laps four
times a year!

From its inception in 1967, when it was produced
five times a year, to today’s much more substantial
quarterly, Trail & Landscape has had only 6 editors,
as each tends to stay for a long time. Karen
McLachlan Hamilton has served in that capacity
since 2001 and has overseen the production of more
issues (52 to the end of 2013) than anyone since
founding editor Anne Hanes (65 issues).

Karen is responsible for seeking articles, editing
them for technical accuracy and presentation, provid-
ing feedback and suggestions to authors, and making
sure the publication is ready for the printer. In this, she
is helped by an associate editor, a production coordina-
tor and a mailing team. But the ultimate responsibility
for ensuring the quality and production standards of
T&L (presentation and timing etc.), is Karen’s. No soon-
er has she put one issue “to bed”, than she is working

CLUB AWARDS

433

on the next, for there is little downtime in this business.
It is astonishing that Karen, with a full-time job, is able
to maintain this hectic schedule with professionalism
and grace. While an honorarium goes along with the
position of Editor, it does not begin to compensate for
the actual work and time that goes into this process.
And besides, Karen has always refused to accept the
stipend. It is indeed, a labour of love.

Karen not only edits T&L, she also writes many arti-
cles for it, such as the annual summary of the OFNC’s
Soiree and Awards Ceremony. She has also written nu-
merous book reviews for T&L, which are characterized
by a forthright discussion of the books in question, inci-
sive comments on content, and descriptive information
on the books themselves. Karen is also a long-serving
member of the OFNC Board of Directors and the club’s
Publications Committee.

For all these reasons, we believe that Karen is very
well deserving of this award to recognize her long serv-
ice on behalf of the OFNC.

(Written by Christine Hanrahan)

Mary Stuart Education Award: Angelika Skevington

The Mary Stuart Education Award is given in recog-
nition of outstanding achievements in the field of nat-
ural history education in the Ottawa Region.

Angela Skevington is the recipient of this award for
2013. Angela is a primary school teacher who is pas-
sionate about natural history education. For over 15
years she has taught a variety of grades in different
schools, sharing her dedication to good environmental
stewardship and her knowledge and wonder of nature
with her students every day, every year. Currently An-
gela teaches a combined Grade 4/5 class at Huntley
Centennial Public School in the village of Carp near
Ottawa.

Wherever she works, Angela has championed such
environmental programs as EarthCare Canada and more
recently Ontario EcoSchools. Last year her school was
recognized with a silver level EcoSchool certification.
This achievement required years of diligent effort on
Angela’s part to engage students and the school com-
munity in activities across the 6 areas of environmen-
tal practice rated by the EcoSchool Program, including
waste minimization and energy conservation. Each
school year, she instigates a student-led club (previ-
ously called an EarthCare Club, currently dubbed The
Dragonflies) that undertakes environmental projects,
such as a periodic audit of their school’s energy and
waste practices. If the audit indicates that there 1s a
problem, such as improper waste disposal, then The
Dragonflies club delivers classroom presentations to
fellow students demonstrating the proper practice, such
as how to separate garbage for recycling. The students
in The Dragonflies club also put on plays at school as-

semblies to demonstrate proper energy and waste prac-
tices. Last year, Angela piloted “litterless lunches” with
her class. The Dragonflies promoted the idea and this
year the whole school is litter-free at lunch. There are
many more examples. Through this hands-on practice,
students learn how to be active environmental stewards.

For the past 3 years, Angela has participated in Otta-
wa’s Clean up the Capital Day. She enlisted classes that
were interested in being involved, applied through the
City of Ottawa to get materials (bags, gloves) and then
they cleaned up the school grounds.

On Earth Day, Angela has conducted student plays at
the school assemblies featuring pollution, anti-littering,
proper recycling. She also encourages the students to
create and perform skits, songs and presentations. She
does this every year to promote celebration of the nat-
ural environment and good environmental stewardship.

Field trips and nature appreciation are important parts
of Angela’s teaching. Habitat is part of the Grade 4 cur-
riculum. She has successfully applied through Ducks
Unlimited Canada’s Project Webfoot for funding for
field trips to a local wetland (such as the Bill Mason
Outdoor Education Centre). They have also had field
trips to the Bonnechere Caves.

In addition to all of this, Angela leads informal
lunchtime field trips for interested students to a nearby
natural area, to learn about and develop an appreciation
for nature. One day they may focus on birds, the next
day on insects, and the third on plants. She applied for
and received funding to organize a field trip for The
Dragonflies club for a day to learn about bird banding
at Innis Point Bird Observatory. The children were excit-

434

ed to each hold a chickadee or nuthatch. On breaks they
played nature games in the field, and went on a nature
hike.

Angela teams up with her husband Jeff Skevington
to help him lead “kid friendly” field trips for The Ottawa
Field-Naturalists’ Club as well as for other nature clubs
in the Ottawa Valley. Angela assisted Jeff and colleagues
in hosting a well-attended Bug Day held at the Fletcher
Wildlife Garden last year.

Angela’s efforts are recognized and appreciated, as
evidenced by fellow teacher Donna Christie who said:

THE CANADIAN FIELD-NATURALIST

Vol. 128

“I feel honoured to work with Angela and know that
she has made a positive impact on her students and our
school .... She is a scientifically-oriented, dedicated,
hard-working educator.”

The OFNC is delighted to present the Mary Stuart
Education Award for 2013 to Angela in recognition of
her success in raising awareness of environmental stew-
ardship and respect for nature at her school and more
broadly.

(Prepared by Eleanor Zurbrigg
with input from Donna Christie)

Index to Volume 128
Compiled by William Halliday

Abundance, |, 327 Least, 350
Abundismus, 63 Bleeker, W., 385
Ackerman, M.R., 191 Bonasa umbellus, 327
Alaska, Fairbanks, 80 Bond, A.L., 72
Alberta, Breeding, 50
Hannah, 145 Breeding Bird Survey, 119
Southern, 50 British Columbia,
Western-Central, 223 Coastal, 57
Albino, 250 Southwestern, 393
Alien, Invasive, 358 Brodo, F., 314, 432
Alvar, 243 Brooding, 145
Amelanistic, 250 Broods, Double, 44
Ammodytes hexapterus, 57 Bubo virginianus, 145, 393
Amos, A., 377 Bunting, Snow, 269
Amphibian, 50, 151 Burbot, 377
Anadromy, 260 Burgess, N.L., 1
Analysis, Pellet, 393 Burn, Prescribed, 400
Anomaly, Colour, 250 Buteo,
Arachnids, 363 lagopus, 145
Archibald, H.L. The Enigma of the 10-year Wildlife regalis 145

Population Cycle Solved? Evidence that the Peri- Butterflies, Local, 358
odicity and Regularity of the Cycle Are Driven —_ Callaghan, C., T. Rytwinski. Editor’s Report for Volume

by a Lunar Zeitgeber, 327-340 127 (2013), 220-222
Arctic, 269 Calls, Night-Flight, 135
Ardetta neoxena, 350 Cambarus bartonii, 111
Ascalapha odorata, 77 Canada, 119
Bags, Leaf Litter, 200 Atlantic, 72
Banding, 135 Eastern, 25, 363
Barcoding, DNA, 77 Canary, Atlantic, 269
Barrier(s), 80 Canis lupus, 189
Ecological, 135 Capture, Hand, 111
Geographic, 135 Capture Success, 111, 191
Bartramia longicauda, 341 Care, Parental, 145
Bat(s), 25 Carex acutiformis, 358
Activity, 158 Carnivora, 25
Big Brown, 158 Catling, P.M. Impact of the 2012 Drought on Woody
Diet, 158 Vegetation Invading Alvar Grasslands in the
Eastern Red, 158 Burnt Lands Alvar, Eastern Ontario, 243-249
Hoary, 158 Catling, P.M., and B. Kostiuk. Use of a Marsh Domi-
Bay, nated by the Introduced European Lake Sedge,
Hudson, 77, 385 Carex acutiformis, by Highly Localized Native
of Fundy, 72 Butterflies, 358-362
Witless, 72 Cavallaro, M.C., A.R. Main, C.A. Morrissey. Muskrat
Bear, Grizzly, 223 (Ondatra zibethicus) Interference with Aquatic
Behaviour, Destructive, 200 Invertebrate Traps, 200-203
Benthic, | Cedar, Eastern White, 243
Birds, Chalmers, L.D., 50
Breeding, 385 Chelydra serpentina, 179
Water, 235 Chiroptera, 25
Bittern, Chronology, Denning, 223
Cory’s, 350 Chrysemys picta marginata, 179

(e.g., article, note), and as such are not always to the exact word location.

435

Se
Note: Page references are to the first page of the item

436

Cipriani, J., 432
Collar (GPRS; 223
Collins, P. A Naturalist for All Seasons: Richard Merrill
Saunders, 1904-1998, 289-294
Colouration, 250
Communication, 195
Connectivity, Wetland, 151
Conservation, 50
Cook, E.R. A Tribute to Kenneth William Stewart,
1936-2011, 84-90
Cook, E.R. Retirement of Associate Editor C.D. Bird,
218
Cook, F.R. Obituary — Farley Mowat 1921-2014, 219
Cook, E.R. A Tribute to Warren Baxter Ballard, 1947—
2012, 276-288
Cook, F.R. William Beverly Scot 1917-2014, 431
Corvus brachyrhnchos, 195
County,
Essex, 135
Fayette, 191
Hardeman, 191
Shelby, 191
Count GeSe45
Cove, Bird, 77
Crayfish,
Allegheny, 111
Appalachian Brook, I11
Calico, 111
Northern Clearwater, 111
Rusty, 111
Virile, 111
Crow, American, 195
Cylce,
10-year Wildlife, 327
Lunar Nodal, 327
Decline,
Amphibian, 151
Population, 165
Dendrochonology, 243
Dentary, 25
Depredation, Nest, 179
Detection, Predator, 179
Detector, Ultrasonic, 158
Development, Embryo, 50
Devlin, J., 111-118
Dicrostonyx nunatakensis, 265
Dictwoy7593
Dimorphism, Sexual, 377
Discrimination, Song, 408
Dispersal, 80
Distribution, |, 77, 204, 265, 363
Divergence, Genetic, 77
Downes, C., 119
Drought, 243
Duncan, J.R., 38
Dussureault, J., 235
Ecology, 179
Bag, oUn /2
Eider, Common, 235

THE CANADIAN FIELD-NATURALIST

Vol. 128

Ekrem, T., P.G. Kevan, T.S. Woodcock, and P.D.N.
Hebert. The Most Northerly Black Witch (As-
calapha odorata): A Tropical Moth in the Cana-
dian Arctic, 77-79

Electrofishing, 111

Elliott, J.E., 393

Elmer, K.R., 377

Entanglement, 173

Eptesicus fuscus, 158

Euphyes dion, 358

Erythristic, 250

Experiment, Playback, 408

Fabianek, F., 25

Farming, Organic, 158

Fauteux, D., G. Lupien, F. Fabianek, J. Gagnon, M.
Séguy, and L. Imbeau. An Illustrated Key to the
Mandibles of Small Mammals of Eastern Cana-
da, 25-37

Feeding, 145, 189

Feierabend, D., and K. Kielland. Multiple Crossings of
a Large Glacial River by Canada Lynx (Lynx
canadensis), 80-83

Fish, Freshwater, 377

Fisher, 272

Fishing, Line Hooking, 377

Florida, 350

Food Provisioning, 145

Forest, Boreal, 80

Francis, C.M., 119

Frog,

Boreal Chorus, 151
Northern Leopard, 50

Gagnon, J., 25

Galbraith, M. 57

Gartersnake, Maritime, 63

Geothlypis philadelphia, 408

Gerard, M.A., 145

Gilhen, J., and F.W. Scott. Melanistic Diversity in the
Maritime Gartersnake, Thamnophis sirtalis pal-
lidulus, in Nova Scotia, Canada, 63-71

Gilliland, S.G., 235

Graham, H., and G.B. Stenhouse. Home Range, Move-
ments, and Denning Chronology of the Grizzly
Bear (Ursus arctos) in West-Central Alberta.
223-234

Grouse, Ruffed, 327

Grousberry, 400

Gull,

Great Black-backed, 165
Herring, 165

Gunson, K., 151

Habitat, 38, 204
Amphibian, 151
Edge, 191
Meso-, 341
Micro-, 341
Winter, 272

Hamer, D., 400

Hanrahan, C., 432

2014

Hare(s), 25
Arctic, 189
Snowshoe, 327

Harris, L.N., J-S. Moore, C.G. McDermid, and H.K.
Swanson. Long-distance Anadromous Migra-
tion in a Fresh Water Specialist: the Lake Trout
(Salvelinus namaycush), 260-164

Hawk,
Ferruginous, 145
Rough-legged, 145

Hearne, Samuel, 269

Hebert, P.D.N., 77

Hendricks, P., S. Lenard. Pygmy Shrew (Sorex hoyii)
in Montana East of the Rocky Mountains with
Comments on its Distribution Across the North-
ern Great Plains, 204-206

Hindmarch, S., and J.E. Elliott. Comparing the Diet of

Great Horned Owls (Bubo virginianus) in Rural
and Urban Areas of Southwestern British Col-
umbia, 393-399

Hipfner, J.M., and M. Galbraith. Diet of the Pacific
Sand Lance (Ammodytes hexapterus) in the Sal-
ish Sea, British Columbia, in the 1960s, 57-62

Hudson, M.-A.R., 119

Identification, 25, 250

Imbeau, L., 25

Invasive Species, 111, 358

[ridistic, 250

Island(s),
Belcher, 385
Boot, 165
Coastal, 63
Ellesmere, 189
Gull, 72
Machias Seal, 72
Nastapoka, 385
Pelee, 135
Tukarak, 385
Victoria, 260

Ixobrychus, “neoxenus” exilis, 350

exilis, 350

Jaw, Lower, 25

Jung, T.S., B.G. Slough, D.W. Nagorsen, and P.M.
Kukka. New Records of the Ogilvie Mountains
Collared Lemming (Dicrostonyx nunatakensis)
in central Yukon, 265-268

Juniper, Common, 243

Juniperus communis, 243

Kansas, Wichita, 1955

Kennedy, A.C., 165, 191

Kevan, P.G., 77

Key, Dichotomous, 25

Kielland, K., 80

Kostiuk, B., 358

Kukka, P.M., 265

Kurta, A., 158

Labrador, 72, 363

Lac, Guillaume-Delisle, 385

Lagomorpha, 25

INDEX TO VOLUME 128

437

Lake(s),
Acidic, |
Erie, 135
Kawartha, 111

Lance, Pacific Sand, 57

Lane, Flight, 195

Langley, W. Effect of Food Patch Discovery on the
Number of American Crows (Corvus brachy-
rhynchos) Using a Flight Lane, 195-199

Lariurus,
borealis, 158
cinereus, 158

Larus,
marinus, 165
argentatus, 165

Lead-backed, 250

Learning, 189

Lemming(s), 25
Ogilvie Mountain Collared, 265

Lenard, S., 204

Lepus,
americanus, 327
arcticus, 189

Lethe Eurydice, 358

Leucism, 350

Leucistic, 250

Limit, Northern, 341

Lithobates pipiens, 50

Litzeus,.JD:, 179

Woche, 377

Long, B.L., A. Kurta. Activity and Diet of Bats in
Conventional versus Organic Apple Orchards
in Southern Michigan, 158-164

Lota lota, 377

Lupien, G., 25

Lynx, Canada, 80, 327

Lynx canadensis, 80, 327

MacKenzie, A., 432

MacKinnon, C.M., A.C. Kennedy. Decline in Breeding
of the Great Black-backed Gull, Larus marinus,
and the Herring Gull, L. argentatus, on Boot
Island, Nova Scotia, 1986 to 2010, 165-172

Macroinvertebrates, |

Main, A.R., 200

Maine, Oxbow, 408

Management, 50
Invasive, 358

Mandible, 25

Manitoba,
Churchill, 77
Duck Mountain Provincial Park, 408
Northern, 77
Southern, 38, 44

Mariposa de la Muerte, 77

Marsh, Delta, 44

Martes pennant, 272

McDermid, C.G., 260

Mech, L.D. A Gray Wolf (Canis lupus) Delivers Live
Prey to a Pup, 189-190

438

Melanism, 63, 350
Mosaic, 63
Pure, 63

Melanistic, 250
Predominantly, 63

Mennill, D.J., 135

Mice, 25

Michigan, Southern, 158

Microtus townsendii, 393

Migration, 77, 260
Bird, 135

Migratory, 77

Miller, V., E. Nol, L.P. Nguyen, and D.M. Turner. Habi-
tat Selection and Nest Success of the Upland
Sandpiper (Bartramia longicauda) in Ivvavik
National Park, Yukon, Canada, 341-349

Minutes of the 135" Annual Business Meeting of the
Ottawa Field-Naturalists’ Club January 14,
2014, 314-326

Model,

Hierarchical Bayesian, 119

Maximum Likelihood, 119

Moehrenschlager, A., 50

Moles, 25

Monitoring, 111
Acoustic, 135
Population, 119
Small Mammal, 191

Montana,

Eastern, 204

Yellowstone River, 173

Moore, J.-D., and M. Ouellet. A Review of Colour
Phenotypes of the Eastern Red-backed Sala-
mander, Plethodon cinereus, in North Ameri-
ca, 250-259

Moore, J.-S., 260

Morph, Colour, 250

Morphometrics, 25

Morrissey, C.A., 200

Mortality, 173

Movement, 223

Murre, Common, 72

Muskeg, 272

Muskrat, 200

Nagorsen, D.W., 265

National Historic Site, Kejimkujik, |

National Park,

Banff, 400

Ivvavik, 341

Jasper, 223

Kejimkujik, |

National Wildlife Area, 165

Nelson, W., 145

Nesting, 38, 165

Net, Mist, 44, 158

New Brunswick, 72

New Hampshire, Twin Mountain, 408

Newfoundland, 72, 408
Southeastern, 235
Stephenville Crossing, 408

THE CANADIAN FIELD-NATURALIST

Vol. 128

Nguyen, L.P., 341

Nigrismus, 63

Noakes, D.L.G., and J.D. Noakes. A Note on Bird
Song: Samuel Hearne’s Observations on the
Snow Bunting (Plectrophenax nivalis), 269-
271

Noakes, J.D., 269

Nol, E., 341

North America, 250

Nova Scotia, |
Boot Island, 165
Coastal, 63
Eastern, 408
Western, 408
Wreck Cove, 408

Nunavut,
Ellesmere Island, 189
Rankin Inlet, 145

Nussbaumer, C., N.L. Burgess, and R.C. Weeber. Dis-
tribution and Abundance of Benthic Macroin-
vertebrates and Zooplankton in Lakes in Kej-
imkujik National Park and National Historic
Site of Canada, Nova Scotia, 1-24

Obituary, 219

Observations, 145

Ondatra zibethicus, 200

Ontario, 179, 350
Eastern, 243
Marathon, 408
Muskoka/Haliburton, 111
Ottawa, 151, 358
Southwestern, 135

Orchard, Apple, 158

Orconectes,
immunis, 111
obscurus, 111
propinquus, 111
rusticus, 111
virilis, 111

Osprey, 173

Ostovar, K., 173

Ottawa Field-Naturalists’ Club Minutes of the 134"
Annual Business Meeting, 314-326

Ouellet, M., 250

Owl,
Barred, 38
Great Horned, 145, 393

Pandion haliaetus, 173

Pardy, S., 363

Patch, Food, 195

Pekania pennant, 272

Pengelly, I, and D. Hamer. Grouseberry (Vaccinium
scoparium Leiberg ex Coville) Fruit Production
in Forest Openings in Banff National Park.
Alberta, 400-407

Peninsula, Avalon, 235

Period, Spawning, 50

Perry, R.C., J.R. Pickavance, and S. Pardy. Spiders of
the Southern Taiga Biome of Labrador, C anada,
363-376

2014 INDEX TO VOLUME 128 439

Pesticides, 158

Phenology, 50

Phenotype(s), 63, 250

Picea glauca, 243

Pickavance, J.R., 363

Pigment, Black, 63

Pine, Eastern White, 243

Pinus strobus, 243

Pirie-Hay, D.W., and A.L. Bond. Thickness of Com-
mon Murre (Uria aalge) Eggshells in Atlantic
Canada, 72-76

Pitocchelli, J. Song Discrimination by Male Mourning
Warblers (Geothlypis philadelphia) and Impli-
cations for Population Divergences across the
Breeding Range, 408-415

Plains,
Boreal, 272
Northern Great, 204

Plant, Larval Food, 358

Playback, Audio, 38

Plectrophenax nivalis, 269

Plethodon cinereus, 250

Poanes,
massasoit, 358
viator, 358

Pollution, Oil, 235

Population Trend, 119

Potholes, Prairie, 204

Power, K., 235

Predators, 179

Production, Fruit, 400

Proulx, G. Late-winter Habitat Use by the Fisher, Peka-
nia pennant (Erxleben, 1777), in the Boreal
Plains Ecozone of Northwestern Saskatchewan,
Canada, 272-275

Provincial Park,
Algonquin, 179
Cypress Hills, 50
Duck Mountain, 408

Pseudacris maculata, 151

Radiation,
Photosynthetically Active, 400
Solar, 400

Randall, L.A., L.D. Chalmers, A. Moehrenschlager,
and A.P. Russell. Asynchronous Breeding and
Variable Embryonic Development Period in the
Threatened Northern Leopard Frog (Lithobates
pipiens) in the Cypress Hills, Alberta, Canada:
Conservation and Management Implications,
50-56

Range, Home, 80, 223

Raptor, 38

Rats, 393

Rattus, 393 .

Recknagel, H., A. Amos, and K.R. Elmer. Morphologi-
cal and Ecological Variation among Populations
and Subspecies of Burbot (Lota lota [L, 1758])
from the Mackenzie River Delta, Canada,

377-384

Recovery, Species, 179

Red-backed, 250

Region, Great Lakes, 243

Reid, S.M., J. Devlin. Effectiveness of Stream Sam-
pling Methods in Capturing Non-native Rusty
Crayfish (Orconectes rusticus) in Ontario, |11-
118

Relationship, Weight-Length, 377

Reserve, Witless Bay Ecological, 72

Restani, M., 173

Riley, J.L., J.D. Litzgus. Cues Used by Predators to
Detect Freshwater Turtle, 179-188

River,
Halokvik, 260
Mackenzie, 377
Tanana, 80
Yellowstone, 173

Robertson, G.J., S.G. Gilliland, P.C. Ryan, J. Dussur-
eault, K. Power, and B.C. Turner. Mortality of
the Common Eider, Somateria mollissima (Lin-
naeus, 1758), and other Water Birds during two
Inshore Oiling Events in Southeastern New-
foundland, 2005 and 2006, 235-242

Rodentia, 25

Rodenticide, 393

Roost, Winter, 195

Russell, A.P., 50

Ryan, PIC,,.235

Rytwinski, T., 220

Salamander,
Eastern Red-backed, 250

Salvelinus namaycush, 260

Sanders, C.E., D.J. Mennill. Acoustic Monitoring of
Migratory Birds over Western Lake Erie: Avian
Responses to Barriers and the Importance of
Islands, 135-144

Sandpiper, Upland, 341

Saskatchewan,
Alvena, 200
Hudson Bay, 408
Northwestern, 272
Saskatoon, 145

Schueler, F.W., 151

Schumtz, J.K., M.A. Gérard, G.S. Court, R.W. Nelson.
Parental Care by Lone Male Ferruginous Hawks
(Buteo regalis), Rough-legged Hawks (Buteo
lagopus), and Great Horned Owls (Bubo virgin-
ianus) was Limited to Providing Food, 145-150

Scott, F.W., 63

Sea, Salish, 57

Seacor, R., K. Ostovar, M. Restani. Distribution and
Abundance of Baling Twine in the Landscape
Near Osprey (Pandion haliaetus) Nests: Impli-
cations for Nestling Entanglement, 173-178

Sealy, S.G. Yellow Warbleys (Setophaga petechia) Rear
Second Broods in Some Years at Delta Marsh,
Manitoba, 44-49

Seburn, D.C., K. Gunson, F.W. Schueler. Apparent
Widespread Decline of the Boreal Chorus Frog

440)

(Pseudacris maculata) in Eastern Ontario, 151-
157

Sedge, European Lake, 358

Séguy, M., 25

Seining, 111

Selection, Nest-Site, 341

Serinus canaries, 269

Setophaga petechia, 44

Shorebirds, 341

Shrew(s), 25, 204
Pygmy, 204

Size, Egg, 72

Skull, 25

Slough, B.G., 265

Smith, A.C., M.-A.R. Hudson, C. Downes, C.M. Fran-
cis. Estimating Breeding Bird Survey Trends
and Annual Indices for Canada: How Do the
New Hierarchical Bayesian Estimates Differ
from Previous Estimates?, 119-134

Smith, P. Cory’s Bittern, Lvobrychus “neoxenus”’ exilis:
Morph, Mutant, or Mixed Bag? 350-357

Somateria mollissima, 235

Song, 408
Bird, 269

Sorex hoyii, 204

Soricomorpha, 25

Spiders, 363

Spruce, White, 243

Starvation, 145

Stenhouse, G.B., 223

Strait of Georgia, 57

Streams, 111

Strix varia, 38

Subspecies, Parapatric, 377

Succession, 243

Survey,
Aerial, 235
Audio, 39, 50, 393
Auditory, 151
Transect, 38, 50, 165, 400
Visual, 173, 179, 235, 341, 363, 385, 393

Swanson, H.K., 260

Swarth, C.W., and W. Bleeker. Observations of the
Summer Birds of Tukarak Island (Belcher Is-
lands, Nunavut), Nastapoka Islands (Nunavut),
and Lac Guillaume-Delisle (Northern Quebec),
385-392

Switch, Food Plant, 358Telemetry, GPS, 80, 223

Tennessee, Southern, 191

Territorial Park, Tombstone, 265

Thamnophis sirtalis pallidulus, 63

Thickness, Eggshell, 72

Thuja occidentalis, 243

Tracks, Winter, 272

Trait, Recessive, 63

Transmitter,
Acoustic, 260
GESH228

Trap (s),

THE CANADIAN FIELD-NATURALIST

Vol. 128

Aquatic Insect Emergence, 200

Light, 158
Orientation, 191
Pitfall, 363
Sherman Live, 191
Snap, 265

Success, 19]
Susceptibility, 191

Tribute, 84, 276, 289, 431

Trout, Lake, 260

Tundra, Alpine, 265

timer BC 235

Turner, D.M., 341

Turtles, 179
Midland Painted, 179
Snapping, 179

Twine, Baling, 173

Uria aalge, 72

Ursus arctos, 223

Vaccinium scoparium, 400

Variation,
Decadal, 57
Ecological, 377
Interannual, 57

Vegetation, Woody, 243

Vocalization, 269

Voles, 25, 393
Townsend’s, 393

Warbler,
Mourning, 408
Yellow, 44

Water, Chemistry, |

Weasels, 25

Weeber, R.C., 1

Weir, 260

Wetlands, Prairie, 200

Whiklo, T.M., and J.R. Duncan. Characteristics of
Barred Owl (Strix varia) Nest Sites in Manitoba,
Canada, 38-43

Wilderness Area, Wilmore, 223

Wildfire, 400

Witch, Black, 77

Wolcott, D.M., M.R. Ackerman, M.L. Kennedy. As-
sessing Capture Success of Small Mammals Due
to Trap Orientation in Field-Forest Edge Habitat,
191-194

Wolf, Gray, 189

Woodcock, T.S., 77

Yukon, 341
Aklavik, 377
Fort McPherson, 377
Inuvik, 377
Ogilvie Mountains, 265
Tstigehtchic, 377

Zooplankton, |

Zurbrigg, E., I. Brodo, J. Cipriani, C. Hanrahan, and
A. MacKenzie. The Ottawa Field-Naturalists’
Club Awards for 2013, presented April 2014,
432-434

2014

Index to Book Reviews

Botany

Applegate, R.D. “Trees of Eastern North America”
by Gil Nelson et al., 2014, 426

Bickerton, H.J. “Field Manual of Michigan Flora” by
E.G. Voss and A.A. Reznicek, 2013, 99

Catling, P. “Plants of Southern Ontario, Trees,
Shrubs, Wildflowers, Grasses, Ferns and
Aquatic Plants” by Richard Dickinson and
France Royer, 2014, 305-306

Haber, E. “Plantes de milieux humides et de bord de
mer du Québec et des Maritimes” by Martine
Lapointe et al., 2014, 424-425

Smith, C.M. “North Pacific Temperate Rainforests:
Ecology and Conservation” by Gordon H.
Orians and John W. Schoen, 2013, 100-101

Environment

Applegate, R.D. “Ecology 3™ Edition” by Michael
Cain et al., 2014, 214

John, R. “The Crossley ID Guide to Britain and Ire-
land” by Richard Crossley and Dominic
Couzens, 2013, 95

McBride, B. “The Once and Future Great Lakes
Country: An Ecological History” by John
Riley, 2013, 215

Other

Beaudcin, A.B. “Bootstrap Geologist: My Life in
Science” by Gene Shinn, 2013, 103-104

Beaudoin, A.B. “Alfred Russel Wallace (2013) On
the Organic Law of Change: A Facsimile Edi-
tion and Annotated Transcription of Alfred
Russel Wallace’s Species Notebook of 1855-
1859”, 2013, 105-107

Fox. G. “The Dismal State of the Great Lakes” by
James P. Ludwig, 2013, 307-310

Haber, E. “The Real Weed Man: Portrait of Canadian
Botanist Gerald A. Mulligan” by Julie Mulli-
gan, 2014, 426-427

Houston, C.S. “A Love Affair With the Birds: The
Life of Thomas Sadler Roberts” by Sue Leaf,
2013, 101-103

John, R., New Titles, 108-109, 216, 311-313, 429-
430

John, R. “Bird Lady — A Lifelong Love Affair with
Birds” by Elizabeth Le Geyt, 2014, 306-307

Sander-Regier, R. “The Once and Future World:
Nature as it was, as it is, and as it could be” by
J.B. MacKinnon, 2013, 427-428

Seburn, D. “Tracks and Shadows: Field Biology as
an Art” by Harry W. Green, 2013, 104-105

INDEX TO VOLUME 128

44]

Zoology

Bocking, E. “The Fish in the Forest: Salmon and the
Web of Life” by Dale Stokes and Doc White,
2014, 421-422

Brooks, R. “Snapper” by Brian Kimberling, 2013,
93-95

Brooks, R. “A Feathered River Across the Sky. The
Passenger Pigeon’s Flight to Extinction” by
Joel Greenberg, 2014, 298-303

Cook, F.R. “A Pocket Guide to Salamanders of Penn-
sylvania” by Walter E. Meshaka, Jr., and
Joseph T. Collins, 2012, 93

Cook, F.R. “Amphibian Conservation — Global Evi-
dence for the Effects of Interventions” by
Rebecca K. Smith and William J. Sutherland,
2014, 416

Cook, F.R. “Amphibians and Reptiles in Minnesota”
by John J. Moriarty and Carol D. Hall, 2014,
416-417

Cook, F.R. “Amphibians of Ohio” by Ralph A. Pfin-
gsten et al., 2013, 417-418

Foster, R. “Animals of the Serengeti and Ngorongoro
Conservation Area” by Adam S. Kennedy and
Vicki Kennedy, 2014, 207-208

Foster, R. “Birds of the Serengeti and Ngorongoro
Conservation Area” by Adam S. Kennedy,
2014, 207-208

Foster, R. “Beetles of Eastern North America” by
Arthur V. Evans, 2014, 295-296

Foster, R. “Birds of the Kenya’s Rift Valley” by
Adam S. Kennedy, 2014, 296-297

Gawn, M. “The Warbler Guide” by Tom Stephenson
and Scott Whittle, 2013, 96

John, R. “Rare Birds of North America” by Steve
Howell, 2013, 209

John, R. “Dolphin” by Alan Rauch, 2014, 211-212

John, R. “The Amazing World of Flyingfish” by
Steve N.G. Howell, 2014, 297

John, R. “A Field Guide to the Larger Mammals of
Tanzania” by Charles Foley et al., 2014, 297-
298

John, R. “A Sparrowhawk’s Lament — How British
Birds of Prey are Faring” by David Cobham,
2014, 303-304

John, R. “Illustrated Checklist of the Birds of the

World — Non Passerines” by Josep del Hoyo et

al., 2014, 418-419

B.K. “Into the Night: Tales of Nocturnal

Wildlife Expeditions” by Rick A. Adams,

2013, 91-92

O'Neill, J. “Yellowstone Wildlife: Ecology and Nat-
ural History of the Greater Yellowstone

Lim,

442
Ecosystem” by Paul A. Johnsgard, 2013, 96-
OF

Sander-Regier, R. “Owls” by Marianne Taylor, 2012,
92

Seburn, D. “Biology and Conservation of North
American Tortoises” by David R. Rostal et al.,
2014, 422

Smith, C.M. “Life on the Rocks: A Portrait of the
American Mountain Goat” by Bruce L. Smith,
2014, 420

THE CANADIAN FIELD-NATURALIST

Vol. 128

Way, J. “Yellowstone Wildlife: Ecology and Natural
History of the Greater Yellowstone Ecosys-
tem” by Paul A. Johnsgard, 2013, 97-98

Way, J. “Deer” by John Fletcher, 2014, 210-211

Way, J. “Wild Again: The Struggle to Save the
Black-footed Ferret” by David Jachowski,
2014, 212-214

Way, J. “Societies of Wolves and Free-ranging Dogs”
by Stephen Spotte, 2012, 423-424

TABLE OF CONTENTS (concluded) Volume 128, Number 4

Book Reviews

ZOOLOGY: Amphibian Conservation — Global Evidence for the Effects of Interventions — Amphibians and
Reptiles in Minnesota — Amphibians of Ohio — Illustrated Checklist of the Birds of the World — Non
Passerines — Life on the Rocks: A Portrait of the American Mountain Goat — The Fish in the Forest:
Salmon and the Web of Life — Biology and Conservation of North American Tortoises — Societies of
Wolves and Free-ranging Dogs

BoTANyY: Plantes de milieux humides et de bord de mer du Québec et des Maritimes — Trees of Eastern
North America

OTHER: The Real Weed Man: Portrait of Canadian Botanist Gerald A. Mulligan — The Once and Future World:
Nature as it was, as it is, as it could be

NEw TITLES

News and Comment

The Canadian Herpetologist (TCH) 4(2), Fall 2014
Northeast Natural History Conference 2015

Annual Conference of the Animal Behavior Society 2015
International Conference on Biodiversity 2015

William Beverly Scott 1917-2014
Club Awards

Index to The Canadian Field-Naturalist Volume 128 By WILLIAM HALLIDAY

Mailing date of the previous issue 128(3): 20 October 2014

2014

416

THE CANADIAN FIELD-NATURALIST Volume 128, Number 4 2014

Articles
The enigma of the 10-year wildlife population cycle solved? Evidence that the periodicity and ,
regularity of the cycle are driven by a lunar zeitgeber HERBERT L. ARCHIBALD 28)

Habitat selection and nest success of the Upland Sandpiper (Bartramia longicauda) in Ivvavik

National Park, Yukon, Canada
VALERIE MILLER, ERICA NOL, LINH P. NGUYEN, and DEVIN M. TURNER 34]

Cory’s Bittern, xobrychus “neoxenus” exilis: morph, mutant, or mixed bag? PAUL SMITH 350

Use of a marsh dominated by the introduced European Lake Sedge, Carex acutiformis, by highly
localized native butterflies PAUL M. CATLING and BRENDA KOSTIUK 358

Spiders of the southern Taiga biome of Labrador, Canada
R. C. PERRY, J. R. PICKAVANCE, and S. PARDY 363

Morphological and ecological variation among populations and subspecies of Burbot (Lota lota [L,
1758]) from the Mackenzie River Delta, Canada
HANS RECKNAGEL, AMy AMOS, and KATHRYN R. ELMER 377

Observations of the summer birds of Tukarak Island (Belcher Islands, Nunavut), Nastapoka Islands
(Nunavut), and Lac Guillaume-Delisle (northern Quebec)

CHRISTOPHER W. SWARTH and WOUTER BLEEKER 385

Comparing the diet of Great Horned Owls (Bubo virginianus) in rural and urban areas of south-
western British Columbia SOFI HINDMARCH and JOHN E. ELLIOTT 393

Grouseberry (Vaccinium scoparium Leiberg ex Coville) Fruit Production in Forest Openings in
Banff National Park, Alberta TAN PENGELLY and DAVID HAMER 400

Song discrimination by male Mourning Warblers (Geothlypis philadelphia) and implications for
4» population divergence across the breeding range JAY PITOCCHELLI 408

Te

(continued on inside back cover)

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