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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



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.
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
Recent Occurrences of Wild-origin Wolves (Canis spp.) in Canada
South of the St. Lawrence River Revealed by Stable Isotope and
Genetic Analysis
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
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:
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
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
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).
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
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.
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.
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.
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.
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
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
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
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
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
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Received 11 January 2015
Accepted 28 September 2015
... We hypothesize that natural range expansion and higher trophic roles for coyotes (in the absence of wolves) have newly enabled the life cycle of E. canadensis in Nova Scotia, raising the possibility that the parasite is present in New Brunswick, Prince Edward Island, and even the island of Newfoundland (McNeill et al., 1984;Lichtenwalner et al., 2014). Because of the lack of physical specimens from the Maritime provinces, it is not possible to determine if the gray wolf, C. lupus, or the eastern wolf, C. lycaon was historically native to this region (McAlpine et al., 2015;Whitaker and Beazley, 2017). It is possible that E. canadensis may have been introduced to Cape Breton Island as early as the 1940s, but in the absence of wild canids at the time, the parasite was unable to maintain transmission. ...
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Echinococcus spp. tapeworms can cause serious diseases in mammals, including humans. Within the E. granulosus species complex, metacestodes produce unilocular cysts that are responsible for cystic echinococcosis in animal intermediate hosts. Canids are definitive hosts, harbouring adult cestodes in their intestines. Adult E. canadensis were recovered from the small intestine of 1 of 262 coyotes (Canis latrans) from Nova Scotia, Canada. Subsequently, we found unilocular cysts in lungs and livers of 4 of 8 sympatric moose (Alces alces) from Cape Breton Island. DNA was extracted from three cysts using the Qiagen DNeasy Blood and Tissue kit and assayed by polymerase chain reaction (PCR) with primers (cest4 and cest5) for a 117-bp region of the small subunit of ribosomal RNA of E. granulosus sensu lato, and further validated as E. canadensis G8 using primers targeting nicotinamide adenosine dinucleotide dehydrogenase subunit 1 (ND1) and cytochrome c oxidase subunit 1 (CO1) mitochondrial genes. These are the first records of E. canadensis in any of the three Maritime provinces, which include Nova Scotia, New Brunswick, and Prince Edward Island. The parasite was thought to be absent in this region due to extirpation of wolves (Canis spp.) in the 1800s. These findings suggest that further wildlife surveillance and risk assessment is warranted.
... While a preference for larger prey (i.e., moose and caribou) could theoretically differentiate resource-use between eastern wolves and coyotes (but see Benson and Patterson, 2013a), large prey are similarly utilized by gray wolves and their hybrids (Benson et al., 2012), thereby reinforcing evidence in support of competitive challenges for eastern wolves. Further investigations that examine fine-scale diet preference in overlapping populations, such as via space use patterns (i.e., radio-telemetry) or dietary comparison (i.e., prey selection, scat or stable isotope analysis; Leighton et al., 2020;McAlpine et al., 2016), could elucidate patterns of overlap and differentiation across the clade beyond what was possible in our large-scale investigation. Nonetheless, for the variety of environmental variables examined eastern wolves appear to be at a distinct disadvantage compared to other canid groups, and this could translate to further genetic homogenization and ultimately, functional extinction. ...
The eastern wolf (Canis lycaon), a species of conservation concern in Canada, is currently restricted to small fragmented populations in south-central Ontario and hybridizes with both encroaching gray wolves and coyote-like canids. We examined niche dynamics in canids undergoing hybridization to determine whether competition among individuals with coyote (Canis latrans) or gray wolf (Canis lupus) ancestry, or their related hybrids, could threaten persistence of the eastern wolf in south-central Canada. Our integrative approach combined extensive genotyping and comparative niche analyses across the hybrid zone to assess available and utilized niche space across all three parental species and their hybrids within the zone of admixture. We focused on detecting niche imbalances within the canid clade that indicate competitive threats, and used these data to identify specific geographic regions that disproportionately favor eastern wolves and might confer natural exclusion of competing canids. We detected low genetic and ecological differentiation among groups across the region. Niche dynamics in the admixture zone were dominated by gray wolf and coyote-like canids, with coyote-like canids in particular exhibiting niche space that overlapped entirely with eastern wolves. Conservation action for eastern wolves must either exploit the narrow niche space that differentiates them from other canids, representing ~5% of their currently occupied space, or accept whichever group dominates the landscapes regardless of genetic makeup. This study suggests that competitive disadvantage can limit species' recovery efforts, and thereby potentially warrant management that targets factors promoting ecological differentiation between groups.
... While there are no existing plans to reintroduce the wolf to Nova Scotia, there is a small possibility that individuals could disperse from populations in Ontario and Québec, or from future reintroductions nearby (Carroll 2003, 2005, McAlpine et al. 2015. Nova Scotia could be proactive in devising a wolf management plan for the province based on this possibility. ...
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This study investigated the ecological and social potential for wolf (Canis spp.) recovery in Nova Scotia, Canada. Reintroduction potential was considered through a GIS-based analysis of land cover, human population density, land ownership, prey density, and road density. Two disconnected areas of adequate habitat for wolves were identified. Qualitative interviews were conducted with seven identified groups on public attitudes towards the wolf and its potential recovery in the province. Opinions ranged from ‘love’ to a strong dislike of wolves, and many interviewees associated wolves with fear and expressed concern that they would come into contact with wolves on or near their properties. It would likely not be advisable to introduce an active wolf reintroduction program in NS at this time, due to the absence of effective habitat connectivity between the two identified areas of suitable habitat, and the public unease about wolf proximity. However, a proactive public education initiative is recommended in case of future reintroductions or natural immigrations of wolves and other top carnivores from nearby populations.
Technical Report
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Les connaissances sur les espèces de grands canidés sauvages occupant le nord-est de l’Amérique du Nord ont connu un essor considérable au cours des deux dernières décennies, notamment grâce à l’utilisation d’outils moléculaires qui ont permis de mieux distinguer les espèces entre elles. Au Québec, on reconnaissait jusqu’à tout récemment le loup gris (Canis lupus), le coyote (C. latrans) ainsi que, possiblement, le loup de l’Est (C. l. lycaon ou C. lycaon). La présente étude, s’appuyant sur un échantillon de 438 grands canidés provenant de différentes régions du Québec (dont 435 canidés sauvages et 3 chiens domestiques), a permis de déceler 4 regroupements génétiques distincts parmi les canidés sauvages : le loup gris (0,7 %), le loup de l’Est (3,0 %), le loup boréal (C. lupus x C. lycaon; 27,8 %) et le coyote de l’Est (C. lycaon x C. latrans; 40,9 %), en plus des hybrides entre ces regroupements, lesquels représentaient plus du quart (27,6 %) de l’échantillon total. Les divers regroupements génétiques de canidés semblent être structurés spatialement selon la végétation dominante. Ainsi, le loup de l’Est, dont la reproduction a été détectée officiellement pour une première fois au Québec, fréquentait spécialement la forêt décidue sur la rive nord du fleuve Saint-Laurent, alors que le loup boréal occupait principalement la forêt boréale, également sur la rive nord du fleuve. Puisque seuls trois individus ont été associés au loup gris, il n’a pas été possible de statuer sur le type de végétation que ces derniers sont les plus susceptibles de fréquenter, bien qu’il semble que ce canidé soit davantage associé aux milieux nordiques (c’est-à-dire à la taïga et à la toundra). Le coyote de l’Est, pour sa part, a surtout été trouvé en forêt décidue au sud du fleuve Saint-Laurent, mais quelques sujets ont été collectés aussi en forêt boréale et dans la taïga. Dans l’échantillon étudié, aucun loup ayant une probabilité d’assignation (Q) supérieure à 0,80 n’a été détecté au sud du fleuve Saint-Laurent, ce qui n’exclut cependant pas la possibilité que des loups puissent occuper ce territoire, puisqu’on y a trouvé un hybride de loup gris et de loup boréal et quelques hybrides de loups et de coyotes. Bien que la taille de l’échantillon utilisé dans le cadre de cette étude fût jugée satisfaisante, la répartition des spécimens analysés n’était cependant pas homogène sur tout le territoire. Certaines régions, telles que l’Outaouais, la Gaspésie–Îles-de-la-Madeleine, la Mauricie, le Centre-du-Québec, Lanaudière, la Côte-Nord et le Nord-du-Québec, étaient sous- représentées. Les regroupements génétiques répertoriés et l’hybridation décelée entre ces derniers dans le cadre de la présente étude suggèrent que l’identification d’un individu appartenant au genre Canis en milieu naturel sur la base de son apparence pourrait être un exercice plus complexe qu’on ne le pensait auparavant. L’étude comparative des grands canidés sur le plan morphologique (phénotype) à partir de l’identité génétique obtenue à l’aide d’outils moléculaires (génotype) semble être la prochaine étape requise pour mieux départager ces derniers entre eux, idéalement grâce à une clé d’identification.
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Thirty-one chapters by a variety of species experts document and describe the biological diversity of the Atlantic Maritime Ecozone, a region encompassing Maritime Canada, the Gaspe and the eastern townships of Quebec south of the St. Lawrence River. Introductory chapters also provide an over-view Atlantic Canadian biodiversity generally, the environmental history of the region, forest landscapes, inland waters, and protected areas. Taxa rarely reviewed for the region, include aquatic hypomycetes, fleshy fungi, zooplankton, watermites, and oribatid mites, among others.
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The latest taxonomic, distributional, habitat and conservation status information is presented for a total of 91 species of marine and terrestrial mammals presently or historically known from Nova Scotia. Four of them are extirpated and one is extinct. Of the 86 remaining species 51 are terrestrial (46 native, 5 introduced). The terrestrial mammals include 9 species of Insectivora, 6 (+ one tentative) of Chiroptera, one Primate, 12 Carnivora, 1 Perrissodactyla, 3 Artiodactyla, 19 Rodentia and 1 Lagomorpha. Native species include Boreal, Transition Zone and Austral elements, the result of Nova Scotia’s mid-latitude position on the continental coast.The 32 recorded marine species include 6 Carnivora (5 seals, walrus) and 26 Cetacea: Delphinidae (10 species), Phocoenidae (1), Monodontidae (1), Kogiidae (2), Physeteridae (1), Ziphiidae (4), Eschrichtiidae (1, extirpated), Balaenopteridae (5) and Balaenidae (1).Since 1971 four small mammal species (Sorex gaspensis, S. dispar, Glaucomys volans and Microtus chrotorrhinus) have been added to the provincial fauna, as a result of the first systematic and intensive sampling ever done in Nova Scotia. All are disjunct and three of them are restricted to forested talus habitats in the Cobequid Mts. or the Cape Breton Highlands. With the two disjunct species already known (Sorex arcticus maritimensis and Peromyscus leucopus caudatus), there are 6 disjunct mammals in the province, comprising 14.3 % of native non-volant terrestrial mammals, 33 % of insectivores and 18.8 % of native rodents. Two of them (Sorex gaspensis and Microtus chrotorrhinus) occur only on Cape Breton Island. No other area of similar size north of Mexico has a comparable proportion of disjunct mammals.Cape Breton Island historically has had a depauperate mammal fauna, lacking 8 species that were present on the adjacent mainland. The building of the connecting Canso Causeway in 1953-55 had a major zoogeographic impact, as it caused the western third of the strait to freeze over in winter and allowed the invasion and establishment of four large mammal species (Canis latrans, Procyon lotor, Mephitis mephitis and Lynx rufus).Le document présente l’information la plus récente sur la taxonomie, la répartition, l’habitat et la situation de 91 espèces de mammifères marins et terrestres qui vivent actuellement ou ont déjà vécu en Nouvelle-Écosse. Quatre d’entre elles ont disparu et une est éteinte. Sur les 86 autres espèces, 51 sont terrestres (46 espèces indigènes et 5 espèces introduites). Les mammifères terrestres comprennent 9 Insectivora, 6 (+ une espèce provisoire) Chiroptera, un Primate, 12 Carnivora, un Perrissodactyla, 3 Artiodactyla, 19 Rodentia et un Lagomorpha. La Nouvelle-Écosse étant située à une latitude moyenne sur la côte continentale, les espèces indigènes qui y vivent sont des éléments de la zone boréale, de la zone de transition et de la zone australe.Parmi les 32 espèces marines signalées, on compte 6 Carnivora important a (5 phoques et le morse) et 26 Cetacea: Delphinidae (10 espèces), Phocoenidae (1), Monodontidae (1), Kogiidae (2), Physeteridae (1), Ziphiidae (4), Eschrichtiidae (1, disparue), Balaenopteridae (5) et Balaenidae (1).Depuis 1971, à la suite du premier échantillonnage systématique et intensif mené en Nouvelle-Écosse, quatre espèces de petits mammifères (Sorex gaspensis, S. dispar, Glaucomys volans et Microtus chrotorrhinus) se sont ajoutées aux espèces fauniques de la province. Ce sont toutes des espèces disjointes, et trois d’entre elles sont confinées aux talus d’éboulis boisés des monts Cobequid ou des hautes-terres du Cap-Breton. Si on inclut les deux espèces disjointes déjà connues (Sorex arcticus maritimensis et Peromyscus leucopus caudatus), il existe 6 espèces disjointes de mammifères dans la province, qui représentent 14,3 % des mammifères terrestres indigènes qui ne volent pas, 33 % des Insectivores et 18,8 % des Rongeurs indigènes. Deux d’entre elles (Sorex gaspensis et Microtus chrotorrhinus) ne sont présentes que dans l’île du Cap-Breton. Au nord du Mexique, aucune autre région de taille semblable n’a une telle proportion d’espèces disjointes de mammifères.Dans le passé, l’île du Cap-Breton renfermait relativement peu de mammifères : on y trouvait 8 espèces de moins que dans la partie continentale de la Nouvelle-Écosse. La construction de la levée de Canso en 1953-1955 a eu un impact important zoogéographique du fait que, depuis, le tiers ouest du détroit gèle durant l’hiver, ce qui a permis à quatre espèces de gros mammifères (Canis latrans, Procyon lotor, Mephitis mephitis et Lynx rufus) de gagner l’île et de s’y établir.
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Wolf management can be controversial, reflecting a wide range of public attitudes. We analyzed wolf management case histories representing a spectrum of approaches in Canada and the United States. During the early 20th century, wolves were considered undesirable. They were subject to persecution and were extirpated from large areas of their original range. With increased environmental awareness in the 1970s, attitudes toward wolves began to change. Wolf conservation became a focus of public interest, providing conditions that favored regional wolf recovery. However, in regions where livestock production or big-game hunting is valued, wolves have continued to be controlled by management authorities or through the actions of individual citizens. With US wolf populations recovering in the conterminous states, a rule was approved to delist the species from endangered to threatened status under the Endangered Species Act. Notwithstanding the intent of legal instruments, history has demonstrated that societal values ultimately determine the survival of species such as the wolf.
The origin and taxonomy of the red wolf (Canis rufus) have been the subject of considerable debate and it has been suggested that this taxon was recently formed as a result of hybridization between the coyote and gray wolf. Like the red wolf, the eastern Canadian wolf has been characterized as a small "deer-eating" wolf that hybridizes with coyotes (Canis latrans). While studying the population of eastern Canadian wolves in Algonquin Provincial Park we recognized similarities to the red wolf, based on DNA profiles at 8 microsatellite loci. We examined whether this relationship was due to similar levels of introgressed coyote genetic material by comparing the microsatellite alleles with those of other North American populations of wolves and coyotes. These analyses indicated that it was not coyote genetic material which led to the close genetic affinity between red wolves and eastern Canadian wolves. We then examined the control region of the mitochondrial DNA (mtDNA) and confirmed the presence of coyote sequences in both. However, we also found sequences in both that diverged by 150 000 - 300 000 years from sequences found in coyotes. None of the red wolves or eastern Canadian wolf samples from the 1960s contained gray wolf (Canis lupus) mtDNA sequences. The data are not consistent with the hypothesis that the eastern Canadian wolf is a subspecies of gray wolf as it is presently designated. We suggest that both the red wolf and the eastern Canadian wolf evolved in North America sharing a common lineage with the coyote until 150 000 - 300 000 years ago. We propose that it retain its original species designation, Canis lycaon.
We describe a model-based clustering method for using multilocus genotype data to infer population structure and assign individuals to populations. We assume a model in which there are K populations (where K may be unknown), each of which is characterized by a set of allele frequencies at each locus. Individuals in the sample are assigned (probabilistically) to populations, or jointly to two or more populations if their genotypes indicate that they are admixed. Our model does not assume a particular mutation process, and it can be applied to most of the commonly used genetic markers, provided that they are not closely linked. Applications of our method include demonstrating the presence of population structure, assigning individuals to populations, studying hybrid zones, and identifying migrants and admixed individuals. We show that the method can produce highly accurate assignments using modest numbers of loci—e.g., seven microsatellite loci in an example using genotype data from an endangered bird species. The software used for this article is available from
Restoration of gray wolves (Canis lupus) to their original range depends not only on a sound ecological basis but also on public acceptance. We samplest 4 special interest groups in New Brunswick about a hypothetical reintroduction to this area. Two white-tailed deer (Odocoileus virginianus) hunter groups and 2 naturalist groups were sampled by questionnaire to test the hypothesis that deer hunters would have more negative attitudes and be less, willing to reintroduce wolves to New Brunswick than would members of naturalist groups. Deer hunters in northern New Brunswick, where deer bunting was closed due to low numbers of deer, were more negative about a reintroduction than southern deer hunters (deer seasons open) and members of naturalist groups. None of the groups were willing to reintroduce wolves to New Brunswick. Positive attitude anti greater willingness to support reintroduction were correlated with higher education, not having previously hunted big game, and less fear of hiking in the woods knowing wolves were present. Knowledge-of-wolf scores for all groups were low. The most common reason given for opposing wolf reintroduction was that it would result in a deer population decline. If wolf reintroduction were ever to be contemplated for New Brunswick, education programs would be necessary to placate public fear of deer population declines.
This article examines the political cultural context and sociopolitical dimensions of wolf management and restoration in the United States. Drawing on the experiences of various wolf programs throughout the country, including New England, the Northern Rockies, Upper-Midwest, Southwest, and Yellowstone National Park, it documents how wolves are often used as a political symbol and surrogate for a number of socially significant policy issues. It also examines the politics of problem definition in the policymaking process. A "politics" model of public policy is used as an analytical framework to examine the following dimensions that are inextricably tied to the debate over wolf management and recovery: land use and the politics of ecosystem management; wilderness preservation; The Wildlands Project and the role of conservation biology in political decisionmaking; the merits and future of the Endangered Species Act; rural culture, concerns and interests; and the contested role of science and public participation in wildlife policymaking and management. The article ends with a discussion of how these sociopolitical and contextual variables affect political decisionmakers and those responsible for wolf management.
We conducted a literature review and contacted several museums to determine the historical occurrence of Wolves (Canis lupus) in the Maritime Provinces. Although there were many anecdotal sighting prior to 1870, no museum specimens originating from the Maritimes were located. Our review suggested that although Wolves were historically present, they were probably not numerous, and were probably extirpated from the Maritime Provinces between 1870 to 1921.
The eastern Coyote or Coywolf (Canis latrans × C. lycaon) inhabiting northeastern North america resulted from hybridization between the expanding population of the western Coyote (Canis latrans) and the remnant population of Eastern Wolf (C. lycaon) and possibly domestic dogs (C. lupus familiaris) in the early 20th century. This study compares the body mass of eastern (i.e., northeastern) Coyotes, western Coyotes, and Eastern Wolves and synthesizes the recent literature to gain better insight into the taxonomic relations and differences of closely-related Canis species. Northeastern Coyotes (males = 16.5 kg; females = 14.7 kg) were statistically (P < 0.0001) intermediate in mass between western Coyotes (males = 12.2 kg; females = 10.7 kg) and Eastern Wolves (males = 28.2 kg, females = 23.7 kg), consistent with their hybrid origin, but were numerically closer to western Coyotes. Large Cohen's d (3.00-8.56),2 (0.915-0.929), and Cohen's f (3.28-3.62) values indicated large effect sizes from the body mass comparisons. Eastern Wolves were 61-71% heavier than the same sex in the northeastern Coyotes, which in turn were ca. 35-37% heavier than the same sex in the western Coyotes. alternatively, western Coyotes were 73-74% of the size of the same sex in the northeastern Coyotes, which in turn were 59-62% of the size of the same sex in the Eastern Wolves. I also attempted to relate mitochondrial DNa (mtDNa) haplotypes to body mass. Six of 17 (35.3%) adult female northeastern Coyotes captured in Massachusetts weighed ≥18 kg, heavier than any other described Coyote from outside northeastern North america. Mitochondrial DNa haplotypes associated with these heavy female northeastern canids were C9 = 4, C19 = 1, and C48 = 1. Body mass (kg) and mtDNa haplotype data of 53 northeastern Coyotes (males = 28, females = 25) showed no difference between haplotype and body mass for males (P < 0.852) or females (P < 0.128), suggesting that there is not a particular haplotype (e.g., C1) that is associated with the heavier animals. I propose that the most appropriate name for this hybrid animal is Coywolf (Canis latrans × C. lycaon), rather than a type of Coyote. Coywolves are distinct, being larger than any other population of Coyotes but smaller than Eastern Wolves. I propose that the 5 distinct types of Canis be recognized as: western Coyote, Coywolf (northeastern Coyote), Eastern Wolf (including Red Wolf C. rufus), Gray × Eastern Wolf hybrids ('Great Lakes' Wolves; C. lupus × C. lycaon or C. lycaon × C. lupus), and Gray Wolf (C. lupus). The implications for wolf recovery in the northeastern United States is discussed.
A search was carried out for historical records, both published and unpublished, that make reference to the native mammalian fauna of Prince Edward Island. Based on documents dating from 1721 to 1890, a comprehensive list of the records for the native mammals of the island has been compiled. Among the new information found is evidence for the presence of the Grey Wolf(as well as the Woodland Caribou) at the time of the first French settlement in 1720, and for the absence of the Beaver and Moose. Historical information has been assembled on the abundance and food-chain relationships of each of the mammalian species, as well as on their interactions with the European population, including the attitudes of the new settlers towards each species. The records indicate that seven of the mammals were extirpated: the Grey Wolf, American Black Bear, American Marten, River Otter, Canada Lynx, Atlantic Walrus and Woodland Caribou. All of these extirpations were due to the activities of the European population, with the attitudes of the settlers contributing to four of them: an indifference to the survival of the otter and Marten, and a direct hostility to the bear and lynx (due to their predation on livestock), leading to the payment of bounties.