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Recent occurrences of wild-origin wolves (Canis spp.) in Canada south of the St. Lawrence River revealed by stable isotope and genetic analysis

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Abstract

A free-ranging canid killed near Caraquet, New Brunswick, Canada, in 2012 exhibited a mitochondrial DnA sequence of Gray Wolf (Canis lupus) origin and a Y-chromosome haplotype of eastern Wolf (C. lycaon) origin. the animal, which is the first wolf recorded in New Brunswick since 1862, was identified as a Gray-eastern Wolf hybrid (C. lupus × C. lycaon) based on analysis of its autosomal microsatellite genotype. Stable carbon isotope values (δ13C) suggest that the Caraquet wolf was of wild origin. Likewise, δ13C analysis suggests that a wolf-coyote hybrid killed in Quebec south of the St. Lawrence River in 2002 was also of wild origin. however, δ13C values for a wolf from the same region in 2006 suggest that this animal spent most of its life feeding predominantly on non-wild-source food items. Recent occurrences of wild-origin animals south of the St. Lawrence River demonstrate that wolves are capable of dispersal to formerly occupied areas in southeastern Canada and the United States. however, limited natural dispersal alone will likely not be sufficient to re-establish wolves in northeastern North America.
386
Introduction
Information concerning the history of the wolf (Can-
is spp.) in Maritime Canada (New Brunswick, Nova
Scotia, and Prince Edward Island) is sparse. The scant
literature suggests that wolves were rare in the region
at the time of European settlement in the early 1600s
(Ganong 1908; Scott and Hebda 2004; Sobey 2007).
Reports of wolves increased in New Brunswick start-
ing in 1774 and peaked during 1840–1860 (Ganong
1908; Parker 1995). Reports of occurrence in Nova Sco-
tia are likewise limited, with the last known wolf in
Nova Scotia killed for bounty in 1845–1847 (Ganong
1908; Scott and Hebda 2004). A bounty on wolves was
introduced in New Brunswick in 1792, apparently in
response to the loss of domestic sheep; a further New
Brunswick bounty on wolves was in effect from 1858
to 1870, with the last bounties paid for three wolves
killed in 1862 (Ganong 1908). Historic information on
the status of wolves on Prince Edward Island is anec-
dotal and limited, but suggests that animals may have
moved onto the Island over the ice from adjacent New
Brunswick and Nova Scotia (Sobey 2007). In 1900,
Thaddeus Thurber collected mammals across parts of
northern New Brunswick and southern Quebec, report-
ing at that time that wolves were rare throughout the
region, with only a few still present in Quebec south of
the St. Lawrence River outside the Gaspé (Elliot 1901).
Although wolves may have continued to occur in New
Brunswick until 1921 (Lohr and Ballard 1996), Ganong
(1908) considered the wolf nearly extirpated in the
province by 1867.
Wolves were, therefore, never common in Maritime
Canada, and Lohr and Ballard (1996) concluded that
there was insufficient historical information to ascertain
whether New Brunswick ever supported a minimum
viable wolf population. By the early 1970s, following
an eastward range expansion, Coyote (Canis latrans)
were abundant in New Brunswick. Recent evidence
suggests that Coyotes now inhabiting eastern Canada
are hybrid, having interbred with wolves as they moved
east (Kays et al. 2010). Regardless, it is now more than
a century since any canid deemed to be a wolf has been
confirmed to be free-ranging in New Brunswick. Thus,
it is significant that in the late winter of 2012 a large
wolf-like canid was shot and killed in northern New
Brunswick.
Recent Occurrences of Wild-origin Wolves (Canis spp.) in Canada
South of the St. Lawrence River Revealed by Stable Isotope and
Genetic Analysis
DONALD F. MCALPINE1, 9, DAVID X. SOTO2, LINDA Y. RUTLEDGE3, 4, TYLER J. WHEELDON5, BRADLEY N.
WHITE6, JAMES P. GOLTZ7, and JOSEPH KENNEDY8
1New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada
2Canadian Rivers Institute and Department of Biology, University of New Brunswick, Fredericton, New Brunswick E3B 5A3
Canada
3Biology Department, Trent University, Peterborough, Ontario K9J 7B8 Canada
4Current address: Department of Ecology and Evolutionary Biology, Princeton University, Princeton, New Jersey 08544 USA
5Environmental and Life Sciences Graduate Program, Trent University, Peterborough, Ontario K9J 7B8 Canada
6Wildlife Forensics DNA Laboratory, Trent University, Peterborough, Ontario K9J 7B8 Canada
7Provincial Veterinary Laboratory, Department of Agriculture and Fisheries, New Brunswick Agriculture Research Station,
P.O. Box 6000, Fredericton, New Brunswick E3B 5H1 Canada
8Fish and Wildlife Branch, Department of Natural Resources, P.O. Box 6000, Fredericton, New Brunswick E3B 5H1 Canada
9Corresponding author: donald.mcalpine@nbm-mnb.ca
McAlpine, Donald F., David X. Soto, Linda Y. Rutledge, Tyler J. Wheeldon, Bradley N. White, James P. Goltz, and Joseph
Kennedy. 2015. Recent occurrences of wild-origin wolves (Canis spp.) in Canada south of the St. Lawrence River
revealed by stable isotope and genetic analysis. Canadian Field-Naturalist 129(4): 386–394.
A free-ranging canid killed near Caraquet, New Brunswick, Canada, in 2012 exhibited a mitochondrial DNA sequence of Gray
Wolf (Canis lupus) origin and a Y-chromosome haplotype of Eastern Wolf (C. lycaon) origin. The animal, which is the first wolf
recorded in New Brunswick since 1862, was identified as a Gray–Eastern Wolf hybrid (C. lupus × C. lycaon) based on analysis
of its autosomal microsatellite genotype. Stable carbon isotope values (δ13C) suggest that the Caraquet wolf was of wild origin.
Likewise, δ13C analysis suggests that a wolf–coyote hybrid killed in Quebec south of the St. Lawrence River in 2002 was also
of wild origin. However, δ13C values for a wolf from the same region in 2006 suggest that this animal spent most of its life feeding
predominantly on non-wild-source food items. Recent occurrences of wild-origin animals south of the St. Lawrence River
demonstrate that wolves are capable of dispersal to formerly occupied areas in southeastern Canada and the United States.
However, limited natural dispersal alone will likely not be sufficient to re-establish wolves in northeastern North America.
Key Words: Gray Wolf; Canis lupus; Eastern Wolf; Canis lycaon; Coyote; Canis latrans; conservation; stable isotopes; genetic
analysis; New Brunswick; Quebec; St. Lawrence River
2015 MCALPINE ET AL.: WOLVES IN CANADA SOUTH OF THE ST. LAWRENCE RIVER 387
Over the past several decades, the few free-ranging
wolves reported south of the St. Lawrence River in
Canada and through the northeastern United States have
generally been considered of wild origin (Elder 2000;
Villemure and Jolicoeur 2004). However, Kays and
Feranec (2011) recently determined that among eight
animals collected in New England, most were of cap-
tive origin, although three were likely of wild origin. In
addition, the assignment of free-ranging animals to spe-
cific types within the genus Canis, especially in regions
of hybridization among Coyotes, domestic dogs, and
wolves, is rarely obvious phenotypically (Chambers et
al. 2012). The origin and identity of wolf-like canids in
the northeastern United States and Canada south of the
St. Lawrence River can, therefore, not be assumed, al -
though such occurrences could be significant to poten-
tial wolf re-introduction or re-establishment in these
regions.
Here we identify the genetic status and origin (wild
versus captive) of a purported wolf shot in New
Brunswick in 2012. We also report on the origin of two
wolves from Quebec south of the St. Lawrence River
killed in 2002 and 2006 and discuss the significance
of these results. Acknowledging that wolf taxonomy
in North America remains controversial, we follow a
three-species model (Chambers et al. 2012; Rutledge
et al. 2012) that includes the Gray Wolf (Canis lupus),
Eastern Wolf (C. lycaon), and Coyote (C. latrans), and
their hybrids, rather than the more traditional two-
species approach, C. lupus and C. latrans (Koblmüller
et al. 2009; vonHoldt et al. 2011).
Methods
Collection and Necropsy
On 6 April 2012, a large canid was shot at a bait
station in a regenerating clear-cut forest about 3 km
southwest of Caraquet (47.7280°N, 64.9474°W; Fig-
ure 1), in northwest New Brunswick, Canada. New
Brunswick Department of Natural Resources (NB
DNR) personnel collected tissue for genetic analysis
and the carcass of the animal was subsequently turned
over to NB DNR. A full necropsy of the animal by a
qualified veterinary pathologist (JPG) was performed;
standard external measurements and body mass were
recorded and alimentary tract contents were collected
for later examination. The skeleton and taxidermied hide
were deposited in the New Brunswick Museum (NBM
M11985). We used the tooth wear criteria of Gipson et
al. (2000) developed for Gray Wolves, in conjunction
with a series of wolf skulls from northern Quebec and
Labrador deposited in the NBM mammal collection
that had been aged by tooth-sectioning (Parker and Lut-
tich 1986), to estimate the age of the animal. We refer to
this specimen hereafter as the “Caraquet wolf.”
Genetic Analysis
DNA was extracted from tongue tissue of the Cara-
quet wolf with a Qiagen DNeasy Blood and Tissue Kit
(Qiagen Inc., Toronto, Ontario, Canada). Maternal,
paternal, and bi-parental genetic markers were inves-
tigated for species identification. The mitochondrial
DNA (mtDNA) control region was amplified as in
Wheeldon et al. (2010), and the polymerase chain reac-
tion product was sequenced in both forward and reverse
directions on an ABI3730 DNA analyzer (Life Tech-
nologies, Burlington, Ontario, Canada). Sequences
were edited in MEGA (version 5; Tamura et al. 2011),
and the consensus sequence was assigned a specific
haplotype based on a search of the National Centre for
Biotechnology Information sequence database using the
Basic Local Alignment Search Tool (BLAST) and com-
parison with previously described sequences (Wilson
et al. 2000).
Four Y-chromosome microsatellite loci and 12 auto-
somal microsatellite loci were genotyped as in Wheel-
don et al. (2010). The Y-chromosome microsatellite
genotype was combined into a haplotype. The autoso-
mal microsatellite genotype of the Caraquet wolf was
analyzed in the Bayesian-clustering program STRUC-
TURE (version 2.3; Pritchard et al. 2000; Hubisz et al.
2009) using default settings (i.e., F-model, infer alpha),
including genotypes from five reference populations
(data from Wheeldon 2009; Wheeldon et al. 2013):
Coyotes from southeastern Ontario (n= 100); Eastern
Wolves from Algonquin Provincial Park, Ontario (n=
62); Gray–Eastern Wolf hybrids from northeastern On -
tario (n= 62; which are known to cluster together with
wolves from parts of Quebec; Wheeldon 2009); Gray
Wolves from Northwest Territories (n= 55); and
Domestic Dogs (C. lupus familiaris; n= 75). The ad -
mixture model of STRUCTURE was run five times
as suming K= 5 for 106iterations following an initial
burn-in of 105iterations and Qvalues for the Caraquet
canid were averaged. We used an exclusion test with
10 000 simulated genotypes and the frequencies-based
method (Paetkau et al. 1995, 2004) in GENECLASS
(version 2; Piry et al. 2004) to determine the probability
of the Caraquet wolf originating from each of the five
reference populations.
Stable Isotope Analyses
Stable isotopes are increasingly used in wildlife ecol-
ogy and forensics to determine the geographic origin or
diet of animals (Bowen et al. 2005; Moore and Sem-
mens 2008; Hobson et al. 2012). Here, we used a multi-
tissue stable isotope approach to determine the wild ver-
sus captive origin of wolf samples based on short- and
long-term trophic history, following the method of Kays
and Feranec (2011). The method assumes that captive-
origin animals have been fed a diet derived from C4
plant material (i.e., corn), whereas wild-origin wolves
have fed on tissue derived from C3plants, which are
dominant in northeastern North America. Free-ranging
urbanor suburban canids may rely on food from domes-
tic sources (i.e., C4) to some degree. This should be
reflected in a carbon stable isotope signature that is
intermediate between those of captive and wild animals.
As isotope turnover rates vary among tissues, the stable
388 THE CANADIAN FIELD-NATURALIST Vol. 129
isotope composition of different tissues can incorporate
trophic information from specific time periods (Tieszen
et al. 1983). For mammals, hair samples provide iso-
topic data since the last molt, while bone should pro-
vide information over the entire life of a canid (Kays
and Feranec 2011). Kays and Feranec (2011) acknowl-
edge that interpreting such signatures with the limited
data currently available can sometimes present chal-
lenges, and they review the limitations and caveats of
the approach. They also investigated the use of nitrogen
stable isotopes to discriminate between wild- and cap-
tive-origin wolves, but did not find this useful.
Collagen was extracted from bone (caudal vertebrae,
scapula, and metatarsals) and hair of the Caraquet wolf
in the manner of Kays and Feranec (2011). To place the
Caraquet wolf in a broader context, we also obtained
hair and bone for stable carbon isotope analysis (δ13C)
from two wolves killed in 2002 and 2006 in Quebec
south of the St. Lawrence (Figure 1).
The 29.1-kg male “Lingwick wolf” was snared in
January 2002 near the village of Sainte-Marguerite-
de-Lingwick (45.6042°N, 71.2875°W). Villemure and
Jolicoeur (2004) reported that the mtDNA profile for
this animal was consistent with an Eastern Wolf–Coyote
hybrid, the microsatellite genotype suggesting 95.0%
shared ancestry with Eastern Wolf from Algonquin
Provincial Park, Ontario. On this basis, Villemure and
Jolicoeur (2004) identified the animal as an Eastern
FIGURE 1. Collection location of the Caraquet wolf (*), a Gray–Eastern Wolf hybrid (Canis lupus × C. lycaon), and extralimital
records of wolves in adjacent Maine and Quebec determined by δ13C to be wild (■), domestic (), or of unknown
origin (?). Wolves 3 and 4 are the most northerly records analyzed by Kays and Feranec (2011). The stippled area
marks the southeastern margin of the current distribution of Gray and Eastern Wolves and their hybrids (C. lupus, C.
lycaon, C. lupus × C. lycaon) in Quebec. 1 = Lingwick wolf, 2 = Sainte-Marguerite wolf, 3 = Museum of Compara-
tive Zoology, Harvard (MCZ) 62506, 4 = MCZ 62507.
2015 MCALPINE ET AL.: WOLVES IN CANADA SOUTH OF THE ST. LAWRENCE RIVER 389
Wolf. In a more recent study using 12 nuclear markers,
this animal clustered with Quebec Coyotes (Stronen
et al. 2012). However, the sample size of Eastern
Wolves from Algonquin Park in Stronen et al. (2012)
was insufficient to identify a distinct cluster of Eastern
Wolves in the program STRUCTURE. The Lingwick
sample may, therefore, have been inaccurately as signed
and A. V. Stronen (Aalborg University, Denmark, per-
sonal communication) has acknowledged that this ani-
mal should be considered a wolf–coyote hybrid. Al -
though the skull of the Lingwick canid is decidedly
smaller and less robust than that of the Caraquet wolf
(ortheSainte-Marguerite wolf, see below; Figure 2), the
body weight of this animal is above the mean for
male Coyote, Coyote–Eastern Wolf hybrids, or Eastern
Wolves provided by Benson et al. (2012). The skull and
mounted skin of this animal are now in the Musée de la
nature et des sciences in Sherbrooke, Quebec (acces-
sion no. 2003.2).
The 48.6-kg “Sainte-Marguerite wolf” was trapped
in November 2006 near Sainte-Marguerite-de-Beauce
(45.514°N, 70.9415°W) and has been identified genet-
ically as an Eastern–Gray Wolf hybrid (J.-F. Dumont,
Québec Ministère des Forêts, de la Faune et des Parcs,
personal communication); the mounted skin is in a pri-
vate collection and the skull is held by the Ministère
des Ressources naturelles et de la Faune, Quebec.
Bone samples were decalcified using 0.5 N HCl at
room temperature for 24–48 h. Samples were rinsed
with distilled water and then decanted once the mineral
portion of the bone was fully dissolved. Lipids were
extracted in a 2:1 (v/v) chloroform:methanol solution
and the resulting bone collagen was oven dried. Hair
samples were washed in the same solution to remove
surface oils and air dried. Samples (about 1 mg) were
analyzed for δ13C using a continuous flow isotope-ratio
mass spectrometer (Thermo-Finnigan, Bremen, Ger-
many) at the Stable Isotopes in Nature Laboratory,
Fred ericton, New Brunswick. Carbon isotope measure-
ments were expressed as isotope delta (δ) in parts per
thousand (‰) relative to the international standard,
Vienna Pee Dee Belemnite. Isotope values were nor-
malized using in-house standards calibrated against
International Atomic Energy Agency reference materi-
als. Analytical precision, estimated by repeated analy-
ses of laboratory standards, was better than ± 0.2‰.
Canid δ13C values were adjusted using the same diet-
tissue discrimination values that Kays and Feranec
(2011) applied to wolves and Coyotes from the north-
eastern United States (+5.0‰ for bone collagen and
+2.6‰ for hair; Roth and Hobson 2000). Here, we
assume that the isotopic composition of the wild diet of
canids in eastern Canada is similar to that of canids
in the northeastern United States. Although potential
FIGURE 2. Comparison of skulls of the Sainte-Marguerite (A), Caraquet (B), and Lingwick (C) wolves. Note the relatively
smaller size and less robust character of C, an Eastern Wolf–Coyote hybrid (Canis lycaon ×C. latrans) compared
with A and B, both Gray–Eastern Wolf hybrids (C. lupus ×C. lycaon). Photo: New Brunswick Museum.
390 THE CANADIAN FIELD-NATURALIST Vol. 129
regional isotopic variation could introduce uncertainty,
we believe it should be negligible given that eastern
Canada and New England are geographically proximate
and that Kays and Feranec (2011) found relatively good
separation in isotopic values between wild and captive
canid diets.
We categorized New Brunswick and Quebec wolves
as “wild” or “captive” using an assignment test based
on likelihood analysis (Rogers et al. 2012). From each
measured wolf δ13C value, we depicted the likely origin
of individuals and summarized the likelihood of origin
for each wolf based on the average probability across
our replicate samples. The likelihood that a wolf sam-
ple originated from a captive versus wild animal was
determined by ƒ( y*µff), assessing a normal prob-
ability density function as follows:
for each wolf subsample (y*), given the expected mean
f) and standard deviation (σf) of δ13C for individuals
growing their tissues on domestic food (mean = −18.2,
standard deviation [SD] = 3.4 for commercial meat and
dog foods typically given to captive canids) or wild
food sources (mean = −27.8, SD = 1.8 for medium and
large mammals and wild fruits that generally domi-
nate their natural diet) analyzed by Kays and Feranec
(2011). Thus, values from replicate samples for each
wolf showed a probability of origin for both domes-
tic and wild food sources, with the highest values of
ƒ(y*µff) indicating the greatest likelihood of the ani-
mal being associated with that food source.
Results
Collection and Necropsy
The Caraquet wolf (Figure 3) was a male; testes were
scrotal and measured 40.6 mm by 29.1 mm. Shortly
after death, the animal weighed 39.9 kg. Tooth wear
suggests that it was 3–4 years old at time of death.
Measurements for the animal were as follows: total
length 1573 mm, tail vertebrae 397 mm, hind foot
278 mm, ear 107.4 mm. Physically, the animal was in
good body condition; it was negative for mange and
carried 12.2–22.6 mm of extra-visceral body fat across
the lateral pelvic region, about 2 mm over the ribs, and
819.2 g of visceral fat. Stomach contents (266.4 g wet
FIGURE 3. Taxidermied skin of the Caraquet wolf, a Gray–Eastern Wolf hybrid (Canis lupus × C. lycaon) (NBM M11985).
Photo: New Brunswick Museum.
2015 MCALPINE ET AL.: WOLVES IN CANADA SOUTH OF THE ST. LAWRENCE RIVER 391
weight) consisted mostly of pig meat from the bait.
However, some residual hairs in the stomach and a
20.4-g (wet weight) fecal pellet of hair proved to be
Moose (Alces americanus). Mature taenid cestodes
were present in the intestine.
Genetic Analysis
Genetic analysis identified the Caraquet wolf as a
Gray–Eastern Wolf hybrid. The mtDNA sequence
(C22; Wilson et al. 2000) was of Gray Wolf origin and
the Y-chromosome haplotype (AA; Wilson et al. 2012)
was of Eastern Wolf origin. STRUCTURE assigned
the Caraquet wolf 96% to the northeastern Ontario
Gray–Eastern Wolf hybrid reference population and
GENECLASS analysis excluded (i.e., P< 0.01) all
populations except northeastern Ontario Gray–Eastern
Wolf hybrids (P= 0.305) as a probable population of
origin.
Stable Isotope Analyses
The δ13C value for bone collagen of −26.85 ± 0.38
for the Caraquet wolf clearly places it within the “wild”
category (range −30.2 to −24.6) of Kays and Feranec
(2011), although hair samples hint at some reliance on
domestic food sources since the last molt (Table 1).
The Lingwick wolf, with a δ13C value for bone collagen
of −28.34 ± 0.16 is also classified as of wild origin. Hair
samples from the Lingwick wolf likewise suggest an
animal that fed on wild food sources since its last molt.
Analysis of bone collagen from the Saint-Marguerite
wolf suggests that this animal spent most of its life sub-
sisting on prey that included substantial domestic food
sources, and hair samples strongly suggest that this ani-
mal had been feeding on or was fed largely domestic
food sources since its last molt. Although the overall
δ13Cvaluefor the Saint-Marguerite wolf, −23.34 ± 1.02,
is ambiguous in terms of categorizing this animal as a
once-captive or a free-ranging urban animal, it strongly
suggests that this wolf did not spend most of its life as
a free-ranging wild animal. Likelihood-based assign-
ment methods support these characterizations (Table
1).
Discussion
Regardless of the historical genetic composition of
Maritime wolves, the Caraquet wolf is the first free-
ranging, wild-origin wolf recorded in New Brunswick
since 1862, presumably dispersing south to the province
from northeastern Ontario or western Quebec north of
the St. Lawrence River. Wolves were extirpated from
Maritime Canada by about 1900 (Harrison and Chapin
1998) and their possible re-establishment, whether by
natural or assisted means, is controversial (Lohr et al.
1996; Nie 2001; Williams et al. 2002; Musiani and
Paquet 2004). However, determining whether wolves
outside their current distributional range represent the
vanguard of re-establishment can be difficult. Such
animals may represent nothing more than isolated
oc currences of once-captive wolves or domesticated
wolf–dog hybrids (Prendergast 1989; Kays and Feranec
2011), particularly as an estimated 300 000 wolf–dog
hybrids are kept as companion animals in the United
States (Fischer 2003). Complicating the sociopolitical
issues surrounding wolf re-establishment has been an
unresolved taxonomy (Chambers et al. 2012), lack of
information on historical distribution of forms (Wilson
et al. 2003; Rutledge 2010a), and possible contempo-
rary hybridization among wolves, coyotes, and dogs
(vonHoldt et al. 2011; Stronen et al. 2012; Monzón
et al. 2013; Way 2013).
The literature suggests that wolves (of unknown
genotype) were not common in Maritime Canada in the
past, so it is not surprising that historical wolf speci-
mens from New Brunswick do not appear to exist (Lohr
and Ballard 1996). Wilson et al. (2000) include Mar-
itime Canada within the historical range of the Gray
Wolf. Wheeldon and White (2009) suggest that admix-
ing of the Gray Wolf and the Eastern Wolf may pre-date
European settlement in the western Great Lakes region,
whereas Wilson et al. (2003) showed that historical
samples from New York and Maine carried only East-
TABLE 1. Stable carbon isotope analysis of bone collagen and hair samples from New Brunswick and Quebec wolves south
of the St. Lawrence River. Trophic discrimination values of δ13C +5.0‰ for bone collagen; +2.6‰ for hair.
Sample Number of
source Mean δ13C ± SD Likelihood of wild origin, % replicates
Caraquet wolf
Collagen −26.85 ± 0.38 98 6
Hair −24.52 ± 0.18 67 4
All samples −25.92 ± 1.19 10
Lingwick wolf
Collagen −28.34 ± 0.16 99 6
Hair −26.79 ± 0.16 98 3
All samples −27.82 ± 0.79 9
Sainte-Marguerite wolf
Collagen −24.26 ± 0.13 57 6
Hair −22.42 ± 0.47 66
All samples −23.34 ± 1.02 12
392 THE CANADIAN FIELD-NATURALIST Vol. 129
ern Wolf mtDNA. Based on 16th century archaeolog-
ical remains, Rutledge et al. (2010a) suggest that the
Eastern Wolf, or perhaps an Eastern–Gray Wolf hybrid
occupied the eastern temperate forest before European
arrival. Kyle et al. (2006) state that after European set-
tlement, it was the Gray Wolf that was extirpated from
southeastern Ontario and Quebec; this was followed by
land clearing, changes in forest cover, and the concomi-
tant movement of the Eastern Wolf northward with
White-tailed Deer (Odocoileus virginianus).
Unfortunately, none of this information resolves
questions about the genetic makeup of the wolves that
may have occupied New Brunswick historically. Har-
rison and Chapin (1998) and Wydeven et al. (1998) felt
it was unclear whether wolves could recolonize avail-
able habitat in northeastern North America without hu -
man assistance and recommended a re-introduction
program. Although Harrison and Chapin (1998) note
that low human population and the extensive forests of
northern New Brunswick provide suitable habitat for
wolves, they also suggest that the St. Lawrence River
and associated human development may present bar-
riers to wolf dispersal south from Ontario and Quebec.
Likewise, Larivière et al. (2000) were uncertain whether
individual wolves from populations in Quebec north of
the St. Lawrence River might disperse to available wolf
habitat in the northeastern United States.
Until 2002, wolves had not been reported in the wild
in Canada south of the St. Lawrence River for more
than a century (Wydeven et al. 1998; Villemure and
Jolicoeur 2004). Nonetheless, δ13C values suggest that
the Caraquet and Lingwick wolves were of wild origin,
indicating that wolves are capable of dispersing from
north of the St. Lawrence River into southern Quebec
and New Brunswick and probably the northeastern
United States, in spite of natural and human barriers.
Kays and Feranec (2011) also report that some free-
ranging wolves in the northeastern United States appear
to be of wild origin based on δ13C values. Furthermore,
in the course of our work, we were made aware of other
purported (and assumed wild) wolves from Quebec
south of the St. Lawrence River and New England (M.
Hénault, Québec Ministère des Forêts, de la Faune et
des Parcs, personal communication; M. McCollough,
Endangered Species Specialist, United States Fish and
Wildlife Service, personal communication; Monzón
2012), in addition to those examined by Kays and
Feranec (2011).
The outcome of any future wolf dispersal into New
Brunswick or New England is unclear. Although Coy-
otes in the east will readily interbreed with the Eastern
Wolf (Kyle et al. 2006; Rutledge et al. 2010b), such
hybridization occurs more rarely with Gray–Eastern
Wolf hybrids (Wheeldon and Patterson 2012). Kyle
et al. (2006) note that, although hybridization may be
reducing the distinctiveness of the Eastern Wolf, it may
also enhance the adaptive potential of wolves. Con-
versely, Coyote hybridization with wolves appears to
have enhanced the adaptive ability of Coyotes in the
northeast (Kays et al. 2010). Kyle et al. (2006) and Way
(2013) both suggest that a Gray–Eastern Wolf hybrid is
a more efficient predator of Moose than a Coyote–
Eastern Wolf hybrid, which is adapted to prey on deer.
Moose is the predominant ungulate in northern New
Brunswick (Forbes et al. 2010).
Wydeven et al. (1998) observed that wolves in south-
ern Ontario and Quebec appeared to be heavily exploit-
ed. Together with the fate of the Lingwick, Saint-
Marguerite, and Caraquet wolves, this suggests that
mortality among wolves dispersing into regions south
of the St. Lawrence River may be high. In light of pos-
sible high mortality and the dense network of roads
adjacent to the St. Lawrence River (well above the
< 0.70 km/km2 selected by Harrison and Chapin [1998]
as a proxy for maximum levels of human presence
compatible with wolf habitat), natural dispersal alone
may be insufficient to re-establish the wolf in the north-
east. Nonetheless, the importance of individual dis-
persers to the evolutionary potential of whatever wolf-
like phenotype populates the region south of the St.
Lawrence River in the future should not be underesti-
mated (Vilà et al. 2003).
The δ13C values for the Saint-Marguerite wolf are
ambiguous, and we are unable to categorize this animal
as either of captive or free-ranging urban origin. The
δ13C values of Kays and Feranec (2011) mark this ani-
mal as an “urban canid” and not wild, but attest to the
statement of Darimont and Reimchen (2002) that the
interpretation of isotopic signals can be challenging
without relevant ecological information. Apparently, the
animal had been resident for some time in an agricul-
tural area and was feeding on domestic animals (J.-F.
Dumont, Québec Ministère des Forêts, de la Faune et
des Parcs, personal communication), but the origin of
this wolf remains uncertain.
Although the approach presented by Kays and Fer-
anec (2011) appears to have considerable utility in sep-
arating wild from formerly captive wolves, its value
could be enhanced with a wider range of sample iso-
tope values from animals of known diet and of wild,
domestic, and urban origin. Wydeven et al. (1998) con-
clude that there is a need for better collection of data
relating to dispersing wolves in the northeast, and Lar-
ivière et al. (2000) argue for the monitoring of wolf
populations in Quebec outside wildlife reserves.
We concur, noting that we had difficulty locating the
remains of the Lingwick and Saint-Marguerite wolves.
Although, the collection of tissue for DNA analysis
from purported wolves occurring outside their normal
range now seems routine, ensuring that skeletal and hair
samples from such animals are deposited in publicly
maintained museum collections should likewise be a
priority. The outcome of analyses of such samples may
well influence future management decisions for the
wolf in northeastern North America
2015 MCALPINE ET AL.: WOLVES IN CANADA SOUTH OF THE ST. LAWRENCE RIVER 393
Acknowledgements
We are grateful to Jacques Mallet, who shot the Cara-
quet wolf, for generously meeting our requests for
information. François Chiasson, Bernard Godin, and
Gérald Richardson Fish and Wildlife Branch, New
Brunswick Department of Natural Resources, retrieved
tissue from the Caraquet wolf for our analysis. We
thank Emily Kerr, Wildlife Forensics DNA Laboratory,
Trent University, for conducting genetics laboratory
work and generating the raw genetic data for the Cara-
quet wolf. Graham Forbes and Karen Vanderwolf as -
sisted with the necropsy and the late Dale Robinson,
Tratton Run Wilderness Company, made himself avail-
abletoskintheanimal following examination. We thank
Rick Cunjak, Heather Burke, Anne McGeachy, Chris-
tine Paton and Mireille Savoie, Stable Isotopes in Na -
ture Laboratory, University of New Brunswick, for
support with stable isotope analysis. The following peo-
ple generously provided information during our efforts
to locate details on purported wolves in the northeast:
Jean-François Dumont, Michel Hénault, Mark McCol-
lough, Astrid Vik Stronen, and Mario Villemure. Lau-
rent Cloutier provided access to the Lingwick wolf
skull and generously allowed us to deposit it in the
Musée de la nature et des sciences, Sherbrooke, Que-
bec, at the conclusion of our analyses. Serge Gauthier,
Musée de la nature et des sciences, Sherbrooke, was
kind enough to allow hair samples to be taken from the
mounted Lingwick wolf for analysis. We are very grate-
ful to Jean-François Dumont, Québec Ministère des
Forêts, de la Faune et des Parcs, for permitting access
to the Sainte-Marguerite skull and collecting hair sam-
ples on our behalf from the privately held mounted
animal.
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Received 11 January 2015
Accepted 28 September 2015
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