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Eastern Wolf - Canis (lupus?) lycaon
Eastern Wolf - Canis (lupus?) lycaon

[Image: wolf-small.jpg]

Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: Canis lupus or Canis lycaon
Subspecies: Canis lupus lycaon or none

The Eastern Wolf, also known as Eastern Canadian Wolf or Eastern Canadian Red Wolf, may be a subspecies of Grey Wolf (Canis lupus lycaon), a distinct species of canid (Canis lycaon) or a hybrid species (Canis lupus X Canis latrans) native to the eastern part of North America since the Pleistocene era. It seems to be closely related to the Red Wolf. Some populations may contain instances of hybridization with Coyotes, known as coywolves.
Many names were proposed, including the Eastern Wolf, Eastern Gray Wolf, Eastern Timber Wolf and Algonquin Wolf, although Eastern Wolf has appeared to gain the most recognition.

Eastern Wolf was recently recognized as a potentially distinct species, but closely related to Red Wolf. Some authors disagree and the status as a distinct species is not official. Now, many international and government organizations carry out scientific research for their taxonomy and genetics to answer this question, as well as researching their ecology and influence on the ecosystem.
The Eastern Wolf is smaller than the Grey Wolf and has a grey-reddish coat with black hairs covering the back and sides of the thorax. The mtDNA analysis confirms that eastern wolf belonged to an ancient form of primitive wolf (with Red Wolf) originating some 750,000 years ago in the eastern part of North America (Nowak 1979, 1992). This distribution of haplotypes shows elements similar to the Red Wolf and probably is a part of this species. Red Wolf populations were extirpated from the wild in the southeastern United States, were reintroduced to the wild in recent decades and are now critically endangered.
On March 31, 2010, a presentation by Ontario Ministry of Natural Resources research scientist Brent Patterson outlined key findings about the Eastern Wolf (and coyotes): Most coyotes in Eastern Ontario are wolf-coyote hybrids; wolves in Algonquin Park are, in general, not inter-breeding with coyotes; and the buffer zone around Algonquin Park is a great success with mortality rates down and populations remaining stable.
Proponents of distinct species designation believe that the Eastern Canadian Wolf is just the remnant northern range of a once continuous range of a native canid - the Eastern wolf (E.C. w & Red Wolf). The pre-Columbian range was thought to include U.S. states east of the Mississippi and south of the Canadian Shield-St. Lawrence corridor.
Unlike the Gray Wolf, the Eastern Wolf in Algonquin Park has never been recorded with an all-black or all-white coat (wolf research in Al. P. cited 2008). Eastern wolf mainly exist in Algonquin Park in Canada-USA border. Type Algonquin is a largely pure genetic population of Eastern wolf while type-Ontario is hybrid with Grey Wolf (possible with C. l. nubilus or C. l. griseoalbus ad etc. ) (Wilson et al. 2000). Mech and Frenzel (1971) suggest that the northeastern Minnesota timber wolves are assigned to C. l. lycaon but are found in an area within 150 km of the range of C. l nubilusas described by Goldman (1944).

[Image: algonquin-park-wolf.jpg]

Physical attributes
The Eastern Wolf is smaller than the Gray Wolf. It has a pale greyish-brown pelt. The back and the sides are covered with long, black hairs. Behind the ears, there is a slight reddish color. These differences in attributes are thought to be a result of their Red Wolf ancestry. The Eastern Wolf is also skinnier than the Gray Wolf and has a more coyote-like appearance. This is because wolves and Coyotes often mate and breed wolf/coyote hybrid pups in the park. The Canadian Parks and Wilderness Society states: "Hybridization with Coyotes has historically been a precursor to the decline of Eastern wolf populations. The Committee on the Status of Wildlife in Canada (COSEWIC) has identified hybridization with Coyotes as one of the major threats facing the Eastern wolf, and hybridization continues to pose a serious challenge to Red Wolf recovery efforts in North Carolina." Because the two animals looks so much alike, a ban on the hunting of Algonquin wolves and Coyotes has been in place to make sure no accidental deaths occur.
Grey Wolves will attack, kill or drive out Coyotes if they find them, but recent studies by John and Mary Theberge suggest that Eastern wolf males possibly mate with and accept Coyote females. John Theberge states that, because Coyotes are smaller than wolves, that female wolves would be less likely to accept a smaller mate.

The Eastern Wolf mainly occupies the area in and around Algonquin Provincial Park in Ontario, and also ventures into adjacent parts of Quebec, Canada. It also may be present in Minnesota and Manitoba. In the past, this species might have ranged south into the United States, but after the arrival of Europeans, these wolves were heavily persecuted and became extirpated from the United States. In Canada, exact numbers of Eastern Canadian Wolves are unknown.

[Image: 230px-Presentdistribution_of_wolf_subspe...ttimbr.jpg]

In Algonquin wolves often travel outside the park boundaries, and enter farm country where some are killed. "Of all the wolf deaths recorded from 1988 to 1999, a minimum of 66% was caused by humans. Shooting and snaring outside park boundaries were the leading causes of death for wolves radio-collared in Algonquin Park". One wolf that was radio-collared in July 1992 was located in October in Gatineau Park (north of Ottawa), which is 170 km from Algonquin Park. By mid-December it had made its way back to Algonquin and then, in March 1993, this wolf's severed head was found nailed to a telephone pole in Round Lake.

The Eastern Wolf preys on white-tailed deer, moose, lagomorphs, and rodents including beaver, muskrat, and mice. Preying on American black bear was also reported. Studies in Algonquin Provincial Park showed that three species accounted for 99% of the wolves' diet: moose (some of which is scavenged), white-tailed deer, and beaver (ca. 33% each). The wolves tend to prey more frequently on beaver in the summer, and on white-tailed deer in the winter.

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[Image: wildcat10-CougarHuntingDeer.jpg]
Eastern Wolves Deemed Separate Species

OurAmazingPlanet Staff - Nov 26, 2012 05:38 PM ET

[Image: gray-wolf.jpg] 
A gray wolf (Canis lupus). The eastern wolf, Canis lupus lycaon, qualifies as a separate species, according to a new review by the U.S. Fish and Wildlife Service. 

Eastern wolves, which used to live in the northeastern United States, but now remain only in southeastern Canada, qualify as a distinct species from their western cousins, according to a review by U.S. Fish and Wildlife Service scientists.

The finding may be important for the future of North American wolves and could help scientists understand how the animals evolved, as noted by USA Today.

In the study, published in October in the journal North American Fauna, the scientists reviewed decades of research on North American wolves, much of it complicated and contradictory. Some studies found 8 subspecies of gray wolves; others suggested as many as 27.

Previously, scientists considered eastern wolves a subspecies of gray wolf, Canis lupus lycaon (pronounced LY-can). However, the new review of reams of genetic data suggests that the animal should be classified as a separate species of wolf entirely.

A tale of three wolves

Eastern wolves would join two universally recognized species of wolves in North America: gray wolves (Canis lupus) and red wolves (Canis rufus). Gray wolves once ranged throughout most of modern-day America, but were hunted and poisoned to the brink of extinction, maintaining only a single population in northern Minnesota, the study noted. The animals have since recovered slightly and been reintroduced to Wyoming's Yellowstone National Park (although hunting has since resumed in Minnesota, Wyoming and elsewhere).

Red wolves were also wiped out from their native range, but have been reintroduced into North Carolina and are thought to be breeding in the wild, according to news reports.

The study found that eastern wolves are most closely related to red wolves, and that both species evolved from a common ancestor shared with coyotes. This helps explain why eastern wolves can still mate with and form hybrid offspring with coyotes, so-called "coywolves." Gray wolves, on the other hand, are known to kill any coyotes they come across.

Smaller than their western cousins, eastern wolves weigh from 62 to 77 pounds (28 to 35 kilograms), according to the study. They preferentially prey on white-tailed deer, unlike gray wolves, which have a more wide-ranging diet, USA Today reported.

Darwin's observations

According to USA Today, the recent study lends support to an account made by Charles Darwin in his 1859 book "On the Origin of Species," in which he wrote: "There are two varieties of the wolf inhabiting the Catskill Mountains in the United States, one with a light greyhound-like form, which pursues deer, and the other more bulky, with shorter legs, which more frequently attacks the shepherd's flocks."

This had looked like another one of Darwin's mistakes, but the recent study suggests his words may have been prescient.

The study could impact the reintroduction of wolves in North America, as it may not be appropriate to move eastern wolves into areas where they weren't previously found, for example. However, study's potential uses remain far from clear.

The authors are careful to state that their findings don't have any bearing on the actions of the U.S. Fish and Wildlife Service, which delisted gray wolves from the Endangered Species List in the Great Lakes in 2011, according to USA Today. 
[Image: wildcat10-CougarHuntingDeer.jpg]
Sicilianu Wrote:

Ecol Evol. 2012 September; 2(9): 2325–2332.
Published online 2012 August 13. doi:  10.1002/ece3.301
PMCID: PMC3488682
Y-chromosome evidence supports widespread signatures of three-species Canis hybridization in eastern North America
Paul J Wilson,1 Linda Y Rutledge,1 Tyler J Wheeldon,2 Brent R Patterson,3 and Bradley N White1
Author information ► Article notes ► Copyright and License information ►
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There has been considerable discussion on the origin of the red wolf and eastern wolf and their evolution independent of the gray wolf. We analyzed mitochondrial DNA (mtDNA) and a Y-chromosome intron sequence in combination with Y-chromosome microsatellites from wolves and coyotes within the range of extensive wolf–coyote hybridization, that is, eastern North America. The detection of divergent Y-chromosome haplotypes in the historic range of the eastern wolf is concordant with earlier mtDNA findings, and the absence of these haplotypes in western coyotes supports the existence of the North American evolved eastern wolf (Canis lycaon). Having haplotypes observed exclusively in eastern North America as a result of insufficient sampling in the historic range of the coyote or that these lineages subsequently went extinct in western geographies is unlikely given that eastern-specific mtDNA and Y-chromosome haplotypes represent lineages divergent from those observed in extant western coyotes. By combining Y-chromosome and mtDNA distributional patterns, we identified hybrid genomes of eastern wolf, coyote, gray wolf, and potentially dog origin in Canis populations of central and eastern North America. The natural contemporary eastern Canis populations represent an important example of widespread introgression resulting in hybrid genomes across the original C. lycaon range that appears to be facilitated by the eastern wolf acting as a conduit for hybridization. Applying conventional taxonomic nomenclature and species-based conservation initiatives, particularly in human-modified landscapes, may be counterproductive to the effective management of these hybrids and fails to consider their evolutionary potential.

Keywords: Canis, eastern wolf, hybridization, microsatellites, Y-chromosome, Y-intron
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Elucidating the taxonomic relationships and evolutionary origin of North American “Canis” has been controversial, with considerable discussion over the number of contemporary wolf species. Originally centered on the red wolf (Canis rufus) (Nowak 1979; Wayne and Jenks 1991; Roy et al. 1994; Nowak et al. 1998; Wayne et al. 1998), the controversy has been extended to the eastern wolf (Canis lycaon) (e.g., Koblmüller et al. 2009; Fain et al. 2010). Both species have been identified as smaller wolves that readily hybridize with coyotes. Initial genetic studies proposed an origin of red and eastern wolves through gray wolf (C. lupus) and western coyote (Canis latrans) hybridization based on a lack of distinct genetic material (Wayne and Jenks 1991; Roy et al. 1994). More recent genetic analyses, however, identified distinct mitochondrial DNA (mtDNA) that supports a North American evolution of the eastern wolf (Rutledge et al. 2010a, 2010b). The debate over the number of North American wolf species has been confounded by various proposed hybridization scenarios (Leonard and Wayne 2008; Koblmüller et al. 2009; Wheeldon and White 2009; Wilson et al. 2009; Wheeldon et al. 2010; vonHoldt et al. 2011). Interestingly, extensive Canis hybridization appears limited to the historic distribution of eastern wolves and red wolves (i.e., primarily east of the Mississippi River within the eastern temperate forests, which probably included Wisconsin and Michigan) with notable limitations to hybridization in more western geographies, particularly between coyotes and gray wolves (Pilgrim et al. 1998; Leonard et al. 2005; Hailer and Leonard 2008).

The difficulty with interpreting the evolutionary history of Canis using mtDNA is that hybridization between eastern wolves (see Figure S1) and coyotes would have caused introgression of closely related sequences from a proposed common New World lineage, both recently (Wilson et al. 2000, 2009) and potentially historically (Wilson et al. 2003; Rutledge et al. 2010b). To test the hypothesis that the eastern wolf, that includes the red wolf for the purpose of this study, evolved in eastern North America independent of the gray wolf, and that it is more closely related to the coyote (Wilson et al. 2000, 2003), we assessed the geographic distribution of paternally inherited Y-chromosomes in male wolves and coyotes in combination with previously described mtDNA sequences proposed to originate from the eastern wolf (Wilson et al. 2000; Wheeldon and White 2009; Rutledge et al. 2010a). This approach has been applied to a regional study in Texas that described localized hybridization among three historically sympatric species: the gray wolf, the coyote, and the red wolf (Hailer and Leonard 2008). That study identified species-specific Y-chromosome microsatellite alleles for gray wolves and coyotes, but it did not consider the relationship of the eastern wolf in the larger Canis evolutionary model and it did not consider the Y-intron sequences in association with the Y-microsatellite haplotypes. Here, we provide novel analysis of Y-intron sequences in conjunction with Y-chromosome microsatellite alleles across a wide geographic range to test the hypothesis of a distinct eastern wolf paternal lineage. We predicted that a North American evolved wolf would have evolved Y-chromosome haplotypes, concordant with previously published mtDNA results (Wilson et al. 2000, 2003; Rutledge et al. 2010a), that were divergent from gray wolves and coyotes and that were geographically localized to the historic distribution of C. lycaon (i.e., in general, east of the Mississippi River within the eastern temperate forest region). As a result of extensive levels of hybridization, these species-specific DNA markers would persist in current eastern Canis hybrids, but would be absent from nonhybridizing western coyotes.

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Materials and Methods
We extracted DNA from Canis samples (see Table 2 for sample sizes) using a DNeasy Blood & Tissue Kit (Qiagen Inc., Mississauga, Ontario, Canada). Samples were collected under capture and handling procedures approved by the Ontario Ministry of Natural Resources' animal care committee or were submitted by registered hunters and trappers. Red wolf samples were provided by the red wolf captive breeding program.
Table 2
Distribution of species-specific Y-chromosome microsatellite and mtDNA haplotypes in North American Canis specimens
Sex was determined by amplification of the last intron of the Zfx/Zfy genes (Shaw et al. 2003). Confirmed males were then profiled at four Y-chromosome microsatellite loci (MS34A, MS34B, MS41A, and MS41B) (Sundqvist et al. 2001) and at a 658 bp fragment of the Zfy intron with primers LGL-331 (5′-CAA ATC ATG CAA GGA TAG AC-3′) and Yint2-335 (5′-GTC CAT TGG ATA ATT CTT TCC-3′; Shaw et al. 2003). The polymerase chain reaction (PCR), chemical and cycling conditions for the Y-chromosome microsatellite loci were as follows: For MS34, 5–10 ng of DNA was amplified in a 15 μL reaction with 1× PCR buffer, 0.2 mm dNTPs (Invitrogen, Burlington, Ontario, Canada), 1.5 mm MgCl2, 0.1 μm MS34A-F primer, 0.15 μm MS34B-F primer, 0.2 μm MS34-R primer, and 1 U Taq DNA polymerase (Invitrogen). PCR cycling included an initial denaturation at 94°C for 5 min followed by 30 cycles of 94°C for 30 sec, annealing at 60°C for 1 min, and extension at 72°C for 1 min, with a final cycle of 60°C for 45 min and storage at 4°C. Conditions for MS41 were similar to MS34, except that primer concentrations were 0.15 μm MS41A-F primer, 0.2 μm MS41B-F primer, 0.2 μm MS41-R primer, and the annealing temperature was 58°C. The Y-intron was amplified under the following PCR conditions in a 20 μL reaction: approximately 5–10 ng of DNA, 1× PCR buffer, 0.2 mm dNTPs, 1.5 mm MgCl2, 0.2 mm each primer, 0.1 μg bovine serum albumin, and 1 U Taq DNA polymerase. PCR steps included initial denaturation at 94°C for 5 min followed by 35 cycles of 94°C for 30 sec, 52°C for 30 sec, and 72°C for 30 sec, followed by a final extension at 72°C for 10 min. All sequencing and microsatellite fragment separation and visualization were performed on a MegaBACE 1000 (GE Healthcare, Baie d'Urfé, Quebec, Canada).

Composite haplotypes were determined based on the alleles present at the four loci. Y-microsatellite haplotypes were standardized to previously published data (Hailer and Leonard 2008) (Table S1). We generated a 400 bp sequence of the last Zfy intron for each microsatellite Y-haplotype. Sequences of the mtDNA control region were generated with primers and conditions previously described in Wilson et al. (2000, 2003). In total, we analyzed 364 wolves and coyotes (Table 1) plus an additional 71 coyotes from previously published literature (Table 2) at the Y-chromosome (Table 1), and 718 wolves and coyotes at the mtDNA control region plus an additional 124 coyotes from previously published literature (Table 2). We used the Y-intron data in combination with the Y-microsatellite data to generate a median-joining network in NETWORK v.4.516 ( using methods described in Bandelt et al. (1999) and using nucleotide states to describe the Zfy intron sequence variation with the microsatellite allele haplotype combinations. A 2:1 weighting was assigned to transversions over transition site differences for the Zfy intron, and the intron sequence variation was weighted twice as high as microsatellite loci. Nucleotide diversity (Pi) of the four Zfy intron sequences was estimated using the software DnaSP v5 (Librado and Rozas 2009). We used the prop.test function in R 2.13.1 (R Development Core Team 2011) to test the null hypothesis that the frequency of the putative eastern wolf Y-chromosome haplotypes associated with Zfy-4 was the same in western coyotes (0/121) as observed in eastern Canis populations (90/288) (see Table 2).
Table 1
Summary of sampled regions including the number of individuals (N) and frequency of occurrence of Y-chromosome haplotypes (in brackets) per geographic region
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Four different sequences were identified within 400 bp of the Zfy intron (Zfy-1, -2, -3, and -4; Genbank Accession numbers: FJ687618, FJ687619, JQ394817, and FJ687620) with three segregating sites. Zfy-2, -3, and -4 each differed by one nucleotide from Zfy-1. Zfy-2 was the only sequence found in Northwest Territories gray wolves (Table 1, Fig. 1a) and was associated with the specific Y-chromosome microsatellite allele size, previously identified as gray wolf (i.e., 208 at MS41a, [Hailer and Leonard 2008]). Intron-3 was observed in western coyotes and captive red wolves (Table 1) and was associated with the allele range identified as a coyote lineage (i.e., 212–218 at MS41a [Hailer and Leonard 2008], Table S1). Zfy-1 haplotypes were found in western coyotes from Saskatchewan and Texas (Table 1, Table S1), and in eastern geographies (Table 1). Zfy-4 was associated with Y-chromosome microsatellite alleles in the size range identified for coyotes, but this sequence was only found in the proposed historic range of the eastern wolf (including the red wolf, i.e., Louisiana), and was not observed in western coyotes (Table 1, Fig. 1a). These eastern wolf Y-chromosome haplotypes were found in 22% of eastern coyotes through southeastern Ontario and into the eastern United States (excluding Louisiana; Table 1, Fig. 1a). Also, gray wolf-like Y-chromosome haplotypes (associated with Zfy-2) were found in eastern coyotes throughout their range and in the captive red wolf population (Table 1, Fig. 1a). Overall nucleotide diversity per site (Pi) based on the four 400 bp sequences of the Y-intron was 0.00375 (±SD, 0.00091) and overall nucleotide divergence with Jukes–Cantor correction (K[JC]) was 0.00188. The average number of nucleotide substitutions per site (Dxy) for each intron sequence compared with Zfy-1 was 0.0025. The proportion of Zfy-4 haplotypes in western coyotes was significantly lower than expected, based on the proportion of Zfy-4 introns found in eastern Canis populations (P = 8.371 × 10–12; 95% CI = 0.25–0.37).
Figure 1
(a) Map of the distribution of North American Canis Y-chromosome haplotypes based on four microsatellite loci and an intron of the Zfy gene. Haplotypes are classified by species: gray for gray wolf (C. lupus); yellow for coyote (C. latrans); and red for ...
The Y-chromosome network (Fig. 1b) shows clear distinctions between the haplogroups associated with coyotes (Zfy-1 and Zfy-3), gray wolves (Zfy-2) and eastern wolves (Zfy-4) when incorporating Zfy intron sequences with Y-chromosome microsatellite haplotypes. The pattern of divergent eastern-specific Y-chromosomes is comparable with previously published phylogenetic analyses and geographic distribution of Canis mtDNA (Fig. 2a,b [Rutledge et al. 2010a]). Similar to the Y-chromosome patterns, there is a stark contrast in the mtDNA composition of western coyote populations compared with that of eastern Canis populations that contain C. lycaon mtDNA, specifically the reciprocally monophyletic clade that includes C1 and C3 (Fig. 2b). As noted elsewhere, mtDNA haplotypes C2 and C13 that group within the coyote clade are of possible C. lycaon origin because they are not found in western regions (Wheeldon and White 2009; Fain et al. 2010; Wheeldon et al. 2010). Three eastern wolf mtDNA haplotypes (C1, C3, and C13) occur in high frequency in the western Great Lakes states (Fain et al. 2010; Wheeldon et al. 2010) and/or Ontario, but they are absent in coyotes sampled from western populations (Table 2). C2 occurs in the captive red wolf population and has typically been identified as the red wolf haplotype (Hailer and Leonard 2008).
Figure 2
(a) Map of the distribution of North American Canis mitochondrial DNA control region haplotypes, classified by species: gray for gray wolf (C. lupus) (includes putative dog haplotypes); yellow for coyote (C. latrans); and red for eastern wolf (C. lycaon ...
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Here, we present new Y-chromosome intron sequence data and provide a novel network analysis of the intron haplotypes in connection with new and previously published Y-microsatellite haplotypes. We also connect the Y-chromosome data to a geographic distribution, and provide comparison of the Y-chromosome data with maternal Canis mtDNA haplotype lineages across a wide geographic distribution. The presence of eastern-specific Y-chromosome and mtDNA haplotypes, absent in nonhybridizing gray wolves and coyotes, supports the origin of a North American evolved eastern wolf. Both the branching patterns of the Y-chromosome and mtDNA suggest an independent divergent lineage of haplotypes closely associated with coyotes and distinct from gray wolves. However, both the monophyletic eastern wolf mtDNA clade (C1 and C3) and the eastern wolf Y-chromosome Zfy-4 haplotypes are clearly divergent from coyotes, and the Zfy-4 lineage appears to be as divergent from coyotes as coyotes are from gray wolves.

One potential alternative interpretation of these patterns is that these variant haplotypes represent coyote-specific maternal and paternal haplotypes within the overall variation in the species. The absence of these haplotypes in western geographies would then result from either localized extinctions due to genetic drift of those mtDNA and Y-chromosomes or a failure to sample them in the population. However, this explanation is highly improbable because the eastern-specific haplotypes are divergent at both the mtDNA and Y-chromosome. The likelihood of haplotype extinctions occurring in western coyotes independently twice for mtDNA (C1 and C3) and three times for the Y-chromosome (4AA, 4BB, 4BR) makes the alternative explanation of genetic drift statistically unlikely. Similarly, the geographic distribution of haplotypes could not be the result of recent coyote expansion within the last century because the timeframe is inconsistent with mutation rates of both markers.

Given the high frequency of mtDNA haplotypes C1, C3, and C13 in wolves from the western Great Lakes states (Fain et al. 2010; Wheeldon et al. 2010) and/or Ontario, and their absence from coyotes sampled from western populations (Table 2), it seems unlikely that these putative eastern wolf haplotypes would not have been detected in nonhybridizing coyotes. This criterion could also apply to the coyote-clade C2 haplotype found in the captive red wolf population (Hailer and Leonard 2008) and in the Louisiana population. Although loss of coyote-clustering sequences in western coyotes through random genetic drift following introgression cannot be ruled out, this scenario is much less likely for the eastern-specific monophyletic grouping of the C1 and C3 haplotypes (Wilson et al. 2000; Rutledge et al. 2010a). Although our study and previous studies have not provided a comprehensive survey of coyotes at mtDNA and Y-chromosomes farther to the west, evidence suggests the central US regions summarized in our study represents the core historical source of where coyotes expanded and colonized North America (Nowak 1979; Parker 1995). Additionally, the western coyote samples analyzed here (Texas, Nebraska, and Saskatchewan) are along the eastern front of recent coyote expansion and are even more likely to have similar haplotypes to those animals found within eastern regions.

Previously analyzed historical specimens further support the mtDNA haplotypes as having an eastern North America origin independent of gray wolves and western coyotes. MtDNA haplotypes observed in specimens collected from the mid-to-late 1800s in New York and Maine, prior to coyote colonization, had a C1 haplotype and a haplotype closely related to C13, thus excluding them as originating from gray wolves (C. lupus) (Wilson et al. 2003). This finding is consistent with the divergent eastern-specific haplotypes further characterized in this study. Although we cannot exclude the possibility of occasional pre-European introgressive hybridization between eastern wolves and coyotes, eastern-specific divergent mtDNA and Y-chromosome haplotypes originating from contemporary coyote expansion and colonization is highly unlikely.

Given the unlikelihood of alternative scenarios, we conclude that the data presented here further support the inclusion of two wolf species, in addition to coyotes, into interpretations of populations, such as the Great Lakes wolf (Wheeldon and White 2009; Fain et al. 2010; Wheeldon et al. 2010) and the eastern coyote (Kays et al. 2010). However, the extent of hybridization among Canis species is so prevalent in eastern North America that essentially all eastern populations of wolves and coyotes surveyed show evidence of mtDNA or Y-chromosome introgression. This includes the historic distribution of the red wolf, specifically the area in Texas where the red wolf animals used to breed the original founders were collected (Wayne and Jenks 1991) (as inferred from captive red wolves; see also Hailer and Leonard 2008). These animals may or may not contain eastern wolf mtDNA, depending on the origin of C2, and they lack eastern wolf Y-chromosomes. The evidence for limited direct C. lupus × C. latrans hybridization in western geographies (Pilgrim et al. 1998; Leonard et al. 2005) is further supported by the absence of gray wolf introgression into western coyotes that would have overlapped with declining gray wolf populations (Hailer and Leonard 2008). Ultimately, the lack of extensive hybridization in the west may reflect the eastern wolf's potential role as an intermediate conduit for mixing of coyote and gray wolf genomes with its own at a significantly broader regional and taxonomic scale than previously reported (Hailer and Leonard 2008; Koblmüller et al. 2009; Wheeldon and White 2009; Kays et al. 2010).

The contemporary hybrid species-complex represents various forms. Specifically, a spectrum of coyote to eastern wolf to gray wolf phenotypes exists in a range of natural to human-modified landscapes, including regional differences in wolves (Mech and Paul 2008) and eastern coyotes (Kays et al. 2010). These differences demonstrate the range of hybrid forms likely resulting from a combination of differential population histories, disproportionate contributions from parental Canis species (Rutledge et al. 2010c), and potentially adaptive divergence on ecological factors, such as prey type. As a result, standard taxonomic nomenclature is difficult to apply to the classification, conservation, and management of wolves and coyotes in eastern North America. We encourage managers and policy makers to consider the evolutionary potential of these hybrid genomes because they may support the adaptability necessary to refill the ecological role once occupied by the purer wolf species that existed prior to European colonization. However, we also recognize that in situations where sufficient habitat exists for recolonization of historic species, efforts to minimize anthropogenic factors that exacerbate hybridization are an important aspect of conservation.

Assuming a three-species model of C. lycaon, C. latrans, and C. lupus, comparing the distributional patterns of Y-chromosomes and mtDNA revealed evidence of extensive multispecies hybridization across the eastern distribution, and the patterns were contrasted in different geographic regions at the population-level. In areas with previously described hybridizing wolves, such as northern Ontario, Manitoba, and Quebec (Grewal et al. 2004; Wheeldon and White 2009), Y-chromosomes from both wolf species were found and coyote Y-chromosomes were notably absent. Wolves in Quebec and northeastern Ontario had some coyote mtDNA, consistent with gene flow from Algonquin Park wolves (Grewal et al. 2004; Wilson et al. 2009). Despite an absence of gray wolf mtDNA in eastern coyotes, there was a surprisingly high frequency of gray wolf-like Y-chromosomes in eastern coyotes that were different from the haplotypes found in northern gray wolves in our study. This may reflect an origin of introgression related to the declining Plains wolves (C. lupus nubilus) or alternatively, these Y-chromosomes may have originated from dogs, as the majority of the Zfy-2 haplotypes in eastern coyotes are common in dog breeds (Sundqvist et al. 2006). The presence in eastern coyotes of Y-chromosome haplotypes observed in gray wolves but not dogs (i.e., 2CE, 2CF) certainly supports some level of gray wolf introgression, possibly via an eastern wolf conduit as coyotes expanded east through Ontario (Kays et al. 2010).

Although we cannot exclude the possibility that the eastern wolf originated from a more complex Pleistocene or early Holocene interaction of gray wolves and coyotes in eastern North America, overall, the sequence divergence and eastern-specificity of Zfy-4 haplotypes suggests a longer standing history of an eastern North American evolved wolf, and the majority of genetic markers evaluated to date suggest a closer relationship of C. lycaon to a North American coyote lineage than the gray wolf lineage. As the Plains wolf has been extirpated and there is apparent Y-chromosome haplotype sharing between European gray wolves and dogs (i.e., FF and HT: Sundqvist et al. 2006; Sundqvist et al. 2001), these alternatives cannot be tested with our data set. Increasing representative data sets from nonhybridizing Canis populations, historic samples, and increased genomic surveys will facilitate the ability to reconstruct these population and species histories.

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We thank the Ontario Ministry of Natural Resources, NSERC, and the Canada Research Chair Program for funding to P. J. W. and B. N. W. Thanks to J. Leonard for providing allele scores for previously published Y-microsatellite haplotype data (Sundqvist et al. 2006).

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Conflict of interest
None declared.

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Supporting Information
Additional Supporting Information may be found in the online version of this article:

Figure S1. Eastern Wolf from Algonquin Provincial Park. Photo by Michael Runtz.

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Table S1. Y-chromosome haplotypes observed in this study: haplotype codes correspond to the Zfy intron sequence followed by the allele letter designations for loci MS34 (first letter) and MS41 (second letter). Allele sizes of haplotypes were compared with those from previous studies (17 [Hxx], 24 [#], 25 [X]) to identify matching haplotypes and their corresponding locations: Nebraska (NE), Texas (TX), Alaska (AK), Northwest Territories (NWT), Saskatchewan (SK), Manitoba (MB), Northwestern Ontario (NWON), Northeastern Ontario (NEON), Algonquin Provincial Park (APP), Southeast Ontario (SEON), New York (NY), North Carolina (NC), Louisiana (LA), Maine/New Brunswick (ME/NB), Quebec (QC), and captive red wolves (RU).

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Please note: Wiley-Blackwell are not responsible for the content or functionality of any supporting materials supplied by the authors. Any queries (other than missing material) should be directed to the corresponding author for the article.

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North America Has Only 1 True Species of Wolf, DNA Shows

By Megan Gannon, Live Science Contributor | July 29, 2016 07:04am ET

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Gray wolves, which are not always gray, are protected under the Endangered Species Act.
Credit: Dan Stahler, courtesy of UCLA

DNA tests of wolves across North America suggest that there is just one species of the canid: the gray wolf.

What's more, populations of red wolves and eastern wolves, thought to be distinct species, are actually just hybrids of gray wolves and coyotes that likely emerged in the last couple hundred years, the study found.

The findings, published in the journal Science Advances on Wednesday (July 27), could have implications for the conservation of wolves considered endangered in the United States, the researchers say.

Shared genes

For the study, scientists sequenced the whole genomes of 28 canids, including gray wolves, eastern wolves, red wolves and coyotes in North America.

The study revealed that gray wolves and coyotes are not very different from each other, genetically speaking. According to the DNA results, the two species likely diverged from a common ancestor in Eurasia about 50,000 years ago —much more recently than previous estimates of 1 million years ago.

Meanwhile, red wolves, thought to be native to the southeastern United States, and eastern wolves from the Great Lakes region, were found to be genetic hybrids.

"These gray-wolf-coyote hybrids look distinct and were mistaken as a distinct species," study author Robert Wayne, a professor of ecology and evolutionary biology at the University of California, Los Angeles, said in a statement.

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Gray-wolf-coyote hybrids (like the one shown here) were once thought to be a distinct species.
Credit: Dave Mech, courtesy of UCLA

Compared with eastern wolves, red wolves were more coyote-like in their genetic makeup, the study found, which makes sense historically. Before the hybridization, humans dramatically altered the habitat of wolves in the southeastern U.S. Once gray wolves started to get hunted out of the region, the hybrid red wolves could mate only with other hybrids and coyotes, the researchers said.

"If you did this same experiment with humans —human genomes from Eurasia —you would find that 1 to 4 percent of the human genome has what looks like strange genomic elements from another species: Neanderthals," Wayne explained.

The researchers thought they would see a bigger chunk of such "strange genomic elements" in red wolves and eastern wolves, perhaps at least 10 to 20 percent of the genome that could not be explained by the animals' relation to gray wolves and coyotes."However, we found just 3 to 4 percent, on average —similar to that found in individuals from the same species when compared to our small reference set," Wayne said.

Conservation implications

Wolves were nearly exterminated from the contiguous United States by the mid-20th century. After gray wolves and red wolves were given protections under the Endangered Species Act in the 1970s, conservation efforts led to a modest comeback in the animals' populations. Red wolves have been reintroduced in North Carolina, and gray wolves have been restored in several areas of the western United States, notably in Yellowstone National Park. But the predators' endangered species listing is still sometimes a controversial, even politically charged issue, opposed by ranchers and farmers who see wolves as a threat to their livestock.

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Compared with eastern wolves, red wolves (like the one shown here) are more coyote-like in their genetic makeup.
Credit: Dave Mech, Courtesy of UCLA

A few years ago, the U.S. Fish and Wildlife Service (FWS) put forth a controversial proposal to remove gray wolves from the endangered species list. Technical distinctions about wolf species were at the heart of the plan. The FWS argued that gray wolves had been restored in enough of their original habitat. The agency relied on a 2012 study to designate a new species, the eastern wolf, as a separate species from the gray wolf; if that were true, it would mean gray wolves had never lived in the eastern United States, and thus the FWS claimed it wasn't responsible for restoring gray wolves in that area.

"The recently defined eastern wolf is just a gray wolf and coyote mix, with about 75 percent of its genome assigned to the gray wolf," Wayne said in the statement. "We found no evidence for an eastern wolf that has a separate evolutionary legacy. The gray wolf should keep its endangered species status and be preserved, because the reason for removing it is incorrect. The gray wolf did live in the Great Lakes area and in the 29 eastern states."

The new results may also call into question whether the red wolf can be listed as an endangered species if further research proves this population is not even a true species.

But Wayne and his co-authors argued that it is "antiquated" to require full species status for an organism to get an endangered listing. The researchers recommend that policy makers take a more flexible approach to applications of the Endangered Species Act so that they can also protect hybrids that fill important ecological niches (i.e., keeping deer populations in check).

Journal Reference:
Bridgett M. Vonholdt, James A. Cahill, Zhenxin Fan, Ilan Gronau, Jacqueline Robinson, John P. Pollinger, Beth Shapiro, Jeff Wall and Robert K. Wayne. Whole-genome sequence analysis shows that two endemic species of North American wolf are admixtures of the coyote and gray wolf. Science Advances, 2016 DOI: 10.1126/sciadv.1501714

Protection of populations comprising admixed genomes is a challenge under the Endangered Species Act (ESA), which is regarded as the most powerful species protection legislation ever passed in the United States but lacks specific provisions for hybrids. The eastern wolf is a newly recognized wolf-like species that is highly admixed and inhabits the Great Lakes and eastern United States, a region previously thought to be included in the geographic range of only the gray wolf. The U.S. Fish and Wildlife Service has argued that the presence of the eastern wolf, rather than the gray wolf, in this area is grounds for removing ESA protection (delisting) from the gray wolf across its geographic range. In contrast, the red wolf from the southeastern United States was one of the first species protected under the ESA and was protected despite admixture with coyotes. We use whole-genome sequence data to demonstrate a lack of unique ancestry in eastern and red wolves that would not be expected if they represented long divergent North American lineages. These results suggest that arguments for delisting the gray wolf are not valid. Our findings demonstrate how a strict designation of a species under the ESA that does not consider admixture can threaten the protection of endangered entities. We argue for a more balanced approach that focuses on the ecological context of admixture and allows for evolutionary processes to potentially restore historical patterns of genetic variation.

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Fig. 1 Admixture proportions, hypothesized branching patterns, and the geographic distribution of Canis in North America.
Top: Previously proposed phylogenetic relationships among Canis lineages, with gray lines indicating putative admixture events (5). Bottom: Geographic distributions of Canis in North America. Sample locations are indicated by dots and abbreviations are described in Table 1. Ancestry proportions from vonHoldt et al. (5) are indicated (proportion gray wolf/proportion coyote; see also new values in Table 3). IRNP, Isle Royale National Park; Ma, million years ago. to this post:[Image: attach.png] Whole_genome_sequence_analysis_shows_that_two_endemic_species_of_North_American_wolf_are_admixtures_of_the_coyote_and_gray_wolf.pdf (910.18 KB)
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Big-game jitters: Coyotes no match for wolves' hunting prowess
Eastern coyote lacks the chops to replace wolves in the ecosystem

Date: March 23, 2017
Source: University of Nebraska-Lincoln
As wolf populations plummeted, the eastern coyote assumed the role of apex predator in forests along the Atlantic Coast. New research, however, shows that the eastern coyote is no match for the wolf. While the eastern coyote can bring down moose and other large prey, it prefers to attack smaller animals and to scavenge.

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John Benson, assistant professor of vertebrate ecology, University of Nebraska-Lincoln.
Credit: Photo courtesy John Benson

It may have replaced the dwindling eastern wolf atop many food chains, but the eastern coyote lacks the chops to become the big-game hunter of an ecosystem, new research led by a University of Nebraska-Lincoln ecologist shows.

Eastern wolves once roamed forests along the Atlantic coast, preying on moose, white-tailed deer and other hoofed mammals collectively known as ungulates. As the wolf population plummeted via the rifle and the trap, however, the eastern coyote inherited the status of apex predator in those habitats.

But a study from John Benson and colleagues provides evidence that the eastern coyote hunts moose and other large prey far less frequently than does the eastern wolf -- instead preferring to attack smaller game or scavenge human leftovers.

The findings help resolve long-standing questions about whether eastern coyotes have filled the ecological niche left vacant when the eastern wolf became threatened, Benson said.

"Wolves rely on large prey to survive," said Benson, assistant professor of vertebrate ecology who conducted the research as a doctoral student at Trent University. "But the smaller size of coyotes appears to give them dietary flexibility to survive on a wider variety of food and prey sizes, making them less predictable predators of large prey.

"Having a top predator that preys consistently on large animals like deer and moose may be an important part of maintaining stable predator-prey dynamics and healthy, naturally functioning ecosystems."

After GPS-tracking 10 packs of eastern wolves and analyzing their kill sites in Ontario, the team estimated that the wolves consumed 54 percent of their ungulate meat from moose and 46 percent from white-tailed deer. By contrast, eight packs of eastern coyote ancestry that occupied separate but neighboring territories got just 11 percent of their ungulate meat from moose and 89 percent from deer.

The eastern wolf weighs between 50 and 65 pounds; the eastern coyote typically hits 40 to 50. Though the extra weight gives eastern wolves a greater chance of killing a moose -- or at least surviving the encounter -- it also demands the greater caloric intake that moose and other meaty prey can provide.

Because wolves need to feed on large prey, their populations tend to rise and fall together, Benson said. Wolves may kill many moose during a winter, for instance, depleting their numbers. With fewer moose available, the wolf population declines, boosting the moose population, which in turn boosts the wolf population, and so on.

Yet the buffet-style menu of the eastern coyote means that its numbers can remain steady or even rise without large prey if alternative food is abundant. This opportunistic diet, Benson said, might also be driving erratic population swings among those lower on the food chain.

"It's important to understand the role that wolves play in ecosystems and to not assume that smaller predators ... perform the same ecological functions," Benson said. "If coyotes start hammering white-tailed deer, and deer start to decline, then (coyotes) can just eat rabbits or squirrels or garbage but continue to prey on deer, too. So we think that could be a destabilizing element.

"There are some areas where you've got way too many white-tailed deer in the east, and then you've got other areas where hunters are concerned because the deer are declining. That speaks to the fact that coyotes are an unpredictable predator."

The study is timely: Canada recently designated the eastern wolf as threatened, with the vast majority of eastern wolves living protected in Ontario's Algonquin Provincial Park.

Human-caused mortality has limited efforts to expand the population beyond Algonquin Park, Benson said, which is made worse by the fact that wolves there are likely naïve to the dangers posed by humans. Another issue: Eastern wolves readily breed with eastern coyotes in the wild, making it difficult to maintain a pure lineage.

"Is there a way to get them to expand numerically and geographically outside of the park? We're not sure at this point," said Benson, who provides advice to a team now developing a recovery plan. "One thing that can be managed is human-caused mortality, so if we can provide additional protection, that should put them on equal demographic footing.

"It's an incredibly challenging situation that is complicated by the interactions of these wolves with coyotes and humans. If the park stays the same, there's no immediate reason that they would go extinct. However, we wouldn't want to go forward with that as our only plan because it offers little chance for expansion."

Though large-scale reintroduction across eastern North America will probably not occur soon, Benson said the study emphasizes the value of preserving delicate predator-prey balances that ecosystems have calibrated over millennia.

"Our work suggests that there's an ecological role that wolves play that won't be played by other animals," he said. "That's probably a role that's worth conserving on landscapes, even outside protected areas. If we're interested in restoring landscapes to a more natural, functioning ecosystem, this would be an important part of that."

Story Source: University of Nebraska-Lincoln. "Big-game jitters: Coyotes no match for wolves' hunting prowess: Eastern coyote lacks the chops to replace wolves in the ecosystem." ScienceDaily. (accessed March 24, 2017).

Journal Reference:
John F. Benson, Karen M. Loveless, Linda Y. Rutledge, Brent R. Patterson. Ungulate predation and ecological roles of wolves and coyotes in eastern North America.Ecological Applications, 2017; DOI: 10.1002/eap.1499

Understanding the ecological roles of species that influence ecosystem processes is a central goal of ecology and conservation biology. Eastern coyotes (Canis latrans) have ascended to the role of apex predator across much of eastern North America since the extirpation of wolves (Canis spp.) and there has been considerable confusion regarding their ability to prey on ungulates and their ecological niche relative to wolves. Eastern wolves (C. lycaon) are thought to have been the historical top predator in eastern deciduous forests and have previously been characterized as deer specialists that are inefficient predators of moose because of their smaller size relative to gray wolves (C. lupus). We investigated intrinsic and extrinsic influences on per capita kill rates of white-tailed deer (Odocoileus virginianus) and moose (Alces alces) during winter by sympatric packs of eastern coyotes, eastern wolves, and admixed canids in Ontario, Canada to clarify the predatory ability and ecological roles of the different canid top predators of eastern North America. Eastern coyote ancestry within packs negatively influenced per capita total ungulate (deer and moose combined) and moose kill rates. Furthermore, canids in packs dominated by eastern coyote ancestry consumed significantly less ungulate biomass and more anthropogenic food than packs dominated by wolf ancestry. Similar to gray wolves in previous studies, eastern wolves preyed on deer where they were available. However, in areas were deer were scarce, eastern wolves killed moose at rates similar to those previously documented for gray wolves at comparable moose densities across North America. Eastern coyotes are effective deer predators, but their dietary flexibility and low kill rates on moose suggest they have not replaced the ecological role of wolves in eastern North America.;jsessionid=15E68E7F4EC5C84CFE4CFAF8C373DE13.f03t03 
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