HOME

TheInfoList



OR:

The eastern wolf (''Canis lycaon'' or ''Canis lupus lycaon'' or ''Canis rufus lycaon'') also known as the timber wolf, Algonquin wolf or eastern timber wolf, is a canine of debated taxonomy native to the
Great Lakes region The Great Lakes region of North America is a binational Canadian–American region that includes portions of the eight U.S. states of Illinois, Indiana, Michigan, Minnesota, New York, Ohio, Pennsylvania and Wisconsin along with the Canadian p ...
and southeastern Canada. It is considered to be either a unique
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
of
gray wolf The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...
or
red wolf The red wolf (''Canis rufus'') is a canine native to the southeastern United States. Its size is intermediate between the coyote (''Canis latrans'') and gray wolf (''Canis lupus''). The red wolf's taxonomic classification as being a separate s ...
or a separate species from both. Many studies have found the eastern wolf to be the product of ancient and recent genetic admixture between the gray wolf and the
coyote The coyote (''Canis latrans'') is a species of canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological nich ...
, while other studies have found some or all populations of the eastern wolf, as well as coyotes, originally separated from a common ancestor with the wolf over 1 million years ago and that these populations of the eastern wolf may be the same species as or a closely related species to the red wolf (''Canis lupus rufus'' or ''Canis rufus'') of the
Southeastern United States The Southeastern United States, also referred to as the American Southeast or simply the Southeast, is a geographical List of regions in the United States, region of the United States. It is located broadly on the eastern portion of the south ...
. Regardless of its status, it is regarded as unique and therefore worthy of conservation with Canada citing the population in eastern Canada (also known as the "Algonquin wolf") as being the eastern wolf population subject to protection. There are two forms, the larger being referred to as the Great Lakes wolf, which is generally found in Minnesota, Wisconsin, the Upper Peninsula of Michigan, southeastern Manitoba and northern Ontario, and the smaller being the Algonquin wolf, which inhabits eastern Canada, specifically central and eastern Ontario and southwestern Quebec, with some overlapping and mixing of the two types in the southern portions of northeastern and northwestern Ontario. The eastern wolf's morphology is midway between that of the gray wolf and the coyote. The fur is typically of a grizzled grayish-brown color mixed with cinnamon. The nape, shoulder and tail region are a mix of black and gray, with the flanks and chest being
rufous Rufous () is a color that may be described as reddish-brown or brownish-red, as of rust or oxidised iron. The first recorded use of ''rufous'' as a color name in English was in 1782. However, the color is also recorded earlier in 1527 as a dia ...
or creamy. It primarily preys on
white-tailed deer The white-tailed deer (''Odocoileus virginianus''), also known as the whitetail or Virginia deer, is a medium-sized deer native to North America, Central America, and South America as far south as Peru and Bolivia. It has also been introduced t ...
, but may occasionally hunt
moose The moose (in North America) or elk (in Eurasia) (''Alces alces'') is a member of the New World deer subfamily and is the only species in the genus ''Alces''. It is the largest and heaviest extant species in the deer family. Most adult ma ...
and
beaver Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers a ...
s.Theberge, J. B. & M. T. Theberge (2004).
The wolves of Algonquin Park, a 12 Year Ecological Study
'', Department of Geography, Publication Series Number 56, University of Waterloo, Waterloo, Ontario
In the third edition of '' Mammal Species of the World'' published in 2005, the mammalogist W. Christopher Wozencraft listed the eastern wolf as a gray wolf subspecies, which supports its earlier classification based on morphology in three studies. This taxonomic classification has since been debated, with proposals based on DNA analyses that includes a gray wolf
ecotype In evolutionary ecology, an ecotype,Greek: ''οίκος'' = home and ''τύπος'' = type, coined by Göte Turesson in 1922 sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a species ...
, a gray wolf with genetic
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...
from the coyote, a gray wolf/coyote hybrid, a gray wolf/red wolf hybrid, the same species as the red wolf, or a separate species (''Canis lycaon'') closely related to the red wolf. Commencing in 2016, two studies using
whole genome sequencing Whole genome sequencing (WGS), also known as full genome sequencing, complete genome sequencing, or entire genome sequencing, is the process of determining the entirety, or nearly the entirety, of the DNA sequence of an organism's genome at a ...
indicate that North American gray wolves and wolf-like canids were the result of ancient and complex gray wolf and coyote mixing, with the Great Lakes wolf possessing 25% coyote ancestry and the Algonquin wolf possessing 40% coyote ancestry. In the US, a bill is before Congress to remove protections under the
Endangered Species Act of 1973 The Endangered Species Act of 1973 (ESA or "The Act"; 16 U.S.C. § 1531 et seq.) is the primary law in the United States for protecting imperiled species. Designed to protect critically imperiled species from extinction as a "consequence of ec ...
for the gray wolf populations located in the western Great Lakes region. In Canada, the eastern wolf is listed as ''Canis lupus lycaon'' under the '' Species At Risk Act 2002, Schedule 1'' - List of Wildlife at Risk. In 2015, the
Committee on the Status of Endangered Wildlife in Canada The Committee on the Status of Endangered Wildlife in Canada (COSEWIC, French: Comité sur la situation des espèces en péril au Canada, COSEPAC) is an independent committee of wildlife experts and scientists whose "raison d'être is to identify s ...
recognized the eastern wolf in central and eastern Ontario and southwestern Quebec as ''Canis''
cf. The abbreviation ''cf.'' (short for the la, confer/conferatur, both meaning "compare") is used in writing to refer the reader to other material to make a comparison with the topic being discussed. Style guides recommend that ''cf.'' be used onl ...
''lycaon'' (''Canis'' species believed to be ''lycaon'') and a threatened species worthy of conservation. The main threat to this wolf is human hunting and trapping outside of the protected areas, which leads to genetic introgression with the
eastern coyote An eastern coyote in the snow near the West Virginia–Virginia state line">Virginia.html" ;"title="West Virginia–Virginia">West Virginia–Virginia state line The eastern coyote is a wild North American canine hybrid with both coyote and wolf ...
due to a lack of mates. Further human development immediately outside of the protected areas and the negative public perception of wolves are expected to inhibit any further expansion of their range. In 2016, the Committee on the Status of Species at Risk in Ontario recognized the Algonquin wolf as a ''Canis sp.'' (''Canis'' species) differentiated from the hybrid Great Lakes wolves which it found were the result of "hybridization and backcrossing among Eastern Wolf (''Canis lycaon'') (aka ''C. lupus lycaon''), Gray Wolf (''C. lupus''), and Coyote (''C. latrans'')".


Taxonomy

The first published name of a taxon belonging to the genus ''Canis'' from North America is ''Canis lycaon''. It was published in 1775 by the German naturalist Johann Schreber, who had based it on the earlier description and illustration of one specimen that was thought to have been captured near Quebec. It was later reclassified as a subspecies of gray wolf by Edward Goldman. In the third edition of '' Mammal Species of the World'' published in 2005, the mammalogist W. Christopher Wozencraft listed the eastern wolf as a gray wolf subspecies, which supports its earlier classification based on morphology in three studies. This taxonomic classification has since been debated. In 2021, the
American Society of Mammalogists The American Society of Mammalogists (ASM) was founded in 1919. Its primary purpose is to encourage the study of mammals, and professions studying them. There are over 4,500 members of this society, and they are primarily professional scientists w ...
considered the eastern wolf as its own species (''Canis lycaon'').


Taxonomic debate

When European settlers first arrived to North America, the
coyote The coyote (''Canis latrans'') is a species of canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological nich ...
's range was limited to the western half of the continent. They existed in the arid areas and across the open plains, including the prairie regions of the midwestern states. Early explorers found some in Indiana and Wisconsin. From the mid-1800s coyotes began expanding beyond their original range. The taxonomic debate regarding North American wolves can be summarised as follows: The evolutionary biologist Robert K. Wayne, whose team is involved in an ongoing scientific debate with the team led by Linda K. Rutledge, describes the difference between these two evolutionary models: "In a way, it is all semantics. They call it a species, we call it an ecotype."


Archaeological evidence

Some of the earliest ''Canis lupus'' specimen were discovered at Cripple Creek Sump,
Fairbanks Fairbanks is a home rule city and the borough seat of the Fairbanks North Star Borough in the U.S. state of Alaska. Fairbanks is the largest city in the Interior region of Alaska and the second largest in the state. The 2020 Census put the p ...
, Alaska, in strata dated 810,000 years old. The dental measurements of the specimens clearly match historical ''Canis lycaon'' specimens from Minnesota.


Genetic evidence

Mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
(mDNA) passes along the maternal line and can date back thousands of years. In 1991, a study of the
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
(mDNA) sequences of wolves and coyotes from across North America found that the wolves of the Minnesota, Ontario and Quebec regions possessed coyote genotypes. The study proposes that dispersing male gray wolves were mating with coyote females in deforested areas bordering wolf territory. The distribution of coyote genotypes within wolves matched the phenotypic differences between these wolves found in an earlier study, with the larger Great Lakes wolf found in Minnesota, the smaller Algonquin (
Provincial Park Ischigualasto Provincial Park A provincial park (or territorial park) is a park administered by one of the provinces of a country, as opposed to a national park. They are similar to state parks in other countries. They are typically open to t ...
) type found in central Ontario, and the smallest and more coyote-like tweed wolf or eastern coyote type occupying sections of southeastern Ontario and southern Quebec. In 2000, a study looked at red wolves and eastern wolves from both eastern Canada and Minnesota. The study agreed that these two wolves readily hybridize with the coyote. The study used 8  microsatellites (genetic markers taken from across the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
of a specimen). The phylogenetic tree produced from the genetic sequences showed a close relationship among the red wolves and the eastern wolves from Algonquin Park, southern Quebec, and Minnesota such that they all clustered together. These then clustered next closer with the coyote and away from the gray wolf. A further analysis using mDNA sequences indicated the presence of coyote in both of these two wolves, and that these two wolves had diverged from the coyote 150,000–300,000 years ago. No gray wolf sequences were detected in the samples. The study proposed that these findings are inconsistent with the two wolves being subspecies of the gray wolf, that red wolves and eastern wolves (eastern Canadian and Minnesota) evolved in North America after having diverged from the coyote, and therefore they are more likely to hybridize with coyotes. In 2009, a study of eastern Canadian wolves – which was referred to as the "Great Lakes" wolf in this study – using microsatellites, mDNA, and the paternally-inherited yDNA markers found that the eastern Canadian wolf was a unique
ecotype In evolutionary ecology, an ecotype,Greek: ''οίκος'' = home and ''τύπος'' = type, coined by Göte Turesson in 1922 sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a species ...
of the gray wolf that had undergone recent hybridization with other gray wolves and coyotes. It could find no evidence to support the findings of the earlier 2000 study regarding the eastern Canadian wolf. The study did not include the red wolf. This study was quickly rebutted on the grounds that it had misinterpreted the findings of earlier studies that it relied upon, nor did it provide a definition for a number of the terms that it used, such as "ecotype". In 2011, a study compared the genetic sequences of 48,000 
single nucleotide polymorphisms In genetics, a single-nucleotide polymorphism (SNP ; plural SNPs ) is a germline substitution of a single nucleotide at a specific position in the genome. Although certain definitions require the substitution to be present in a sufficiently lar ...
(
mutations In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
) taken from the genomes of canids from around the world. The comparison indicated that the red wolf was about 76% coyote and 24% gray wolf with hybridization having occurred 287–430 years ago. The eastern wolf – which was referred to as the "Great Lakes" wolf in this study – was 58% gray wolf and 42% coyote with hybridization having occurred 546–963 years ago. The study rejected the theory of a common ancestry for the red and eastern wolves. However the next year, a study reviewed a subset of the 2011 study's
Single-nucleotide polymorphism In genetics, a single-nucleotide polymorphism (SNP ; plural SNPs ) is a germline substitution of a single nucleotide at a specific position in the genome. Although certain definitions require the substitution to be present in a sufficiently ...
(SNP) data and proposed that its methodology had skewed the results and that the eastern wolf is not a hybrid but a separate species. The 2012 study proposed that there are 3 true canis species in North America - the gray wolf, the western coyote, and red wolf/eastern wolf with the eastern wolf represented by the Algonquin wolf, with the Great Lakes wolf being a hydrid of the eastern wolf and the gray wolf, and the eastern coyote being a hybrid of the western coyote and the eastern (Algonquin) wolf. Also in 2011, a scientific
literature review A literature review is an overview of the previously published works on a topic. The term can refer to a full scholarly paper or a section of a scholarly work such as a book, or an article. Either way, a literature review is supposed to provid ...
was undertaken to help assess the taxonomy of North American wolves. One of the findings proposed was that the eastern wolf, whose range includes eastern Canada and the Upper Peninsula of Michigan plus Wisconsin and Minnesota is supported as a separate species by morphological and genetic data. Genetic data supports a close relationship between the eastern and red wolves, but not close enough to support these as one species. It was "likely" that these were the separate descendants of a common ancestor shared with coyotes. This review was published in 2012. Another study of both mDNA and yDNA in wolves and coyotes by the same authors indicates that the eastern wolf is genetically divergent from the gray wolf and is a North American evolved species with a long-standing history. The study could not dismiss the possibility of the eastern wolf having evolved from an ancient hybridization of gray wolf and coyote in the Late Pleistocene or Early Holocene. Another study by the same authors found that eastern wolf mDNA genetic diversity had been lost after their culling in the early 1960s, leading to the invasion of coyotes into their territory and introgression of coyote mDNA. In 2014, the
National Center for Ecological Analysis and Synthesis The National Center for Ecological Analysis and Synthesis (NCEAS) is a research center at the University of California, Santa Barbara, in Santa Barbara, California. Better known by its acronym, NCEAS (pronounced “n-seas”) opened in May 1995. Fu ...
was invited by the
United States Fish and Wildlife Service The United States Fish and Wildlife Service (USFWS or FWS) is an agency within the United States Department of the Interior dedicated to the management of fish, wildlife, and natural habitats. The mission of the agency is "working with othe ...
to provide an independent review of its proposed rule relating to gray wolves. The Center's panel findings were that the proposed rule was heavily dependent upon the analysis contained in a scientific literature review conducted in 2011 (Chambers et al.), that this work was not universally accepted and that the issue was "not settled", and that the rule does not represent the "best available science". Also in 2014, an experiment to hybridize a captive western gray wolf and a captive western coyote was successful, and therefore possible. The study did not assess the likelihood of such hybridization in the wild. In 2015, the
Committee on the Status of Endangered Wildlife in Canada The Committee on the Status of Endangered Wildlife in Canada (COSEWIC, French: Comité sur la situation des espèces en péril au Canada, COSEPAC) is an independent committee of wildlife experts and scientists whose "raison d'être is to identify s ...
changed its designation of the eastern wolf from ''Canis lupus lycaon'' to ''Canis cf. lycaon'' (''Canis'' species believed to be ''lycaon'') and a species at risk. Later that year, a study compared the
DNA sequences A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequences are us ...
using 127,000 single-nucleotide polymorphisms (mutations) of wolves and coyotes, but did not include red wolves and used Algonquin wolves as the representative eastern wolf, not wolves from the western Great Lakes states (usually referred to as Great Lakes wolves). The study indicated that Algonquin wolves were a distinct genomic cluster, even distinct from the Great Lakes states' wolves, which it found were actually hybrids of the gray wolf and the Algonquin wolf. The study's results did not exclude a possibility that the Great Lakes states' wolf (the gray wolf x eastern wolf hybrid (''C. l. lycaon'')) historically inhabited southern Ontario, southern Quebec and the northeastern United States alongside the Algonquin wolf, as there is evidence to suggest both inhabited those areas. In 2016, a study of mDNA once again indicated the Eastern wolf as a coyote–wolf hybrid. In 2018, a study looked at the
y-chromosome The Y chromosome is one of two sex chromosomes ( allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or ...
male lineage of canines. The unexpected finding was that the one Great Lakes wolf specimen included in this study showed a high degree of
genetic divergence Genetic divergence is the process in which two or more populations of an ancestral species accumulate independent genetic changes ( mutations) through time, often leading to reproductive isolation and continued mutation even after the populations ha ...
. Previous studies propose the Great Lakes wolf to be an ancient ecotype of the gray wolf that had experienced genetic
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...
from other types of gray wolves and coyotes. The study called for further research into the Y-chromosomes of coyotes and wolves to ascertain if this is where this unique genetic male lineage may have originated from.


Genomic evidence

In 2016, a whole-genome DNA study proposed, based on the assumptions made, that all of the North American wolves and coyotes diverged from a common ancestor less than 6,000–117,000 years ago, including the coyote diverging from Eurasian wolf about 51,000 years ago (which matches other studies indicating that the extant wolf came into being around this time), the red wolf diverging from the coyote between 55,000–117,000 years ago, and the eastern wolf (Great Lakes region and Algonquin) wolf diverging from the coyote 27,000–32,000 years ago, and asserts that these do not qualify as an ancient divergences that justify them being considered unique species. The study also indicated that all North America wolves have a significant amount of coyote ancestry and all coyotes some degree of wolf ancestry, and that the
red wolf The red wolf (''Canis rufus'') is a canine native to the southeastern United States. Its size is intermediate between the coyote (''Canis latrans'') and gray wolf (''Canis lupus''). The red wolf's taxonomic classification as being a separate s ...
and eastern wolf are highly admixed with different proportions of gray wolf and coyote ancestry. The study found that coyote ancestry was highest in red wolves from the southeast of the United States and lowest among the Great Lakes wolves. The study also determined how unique each type of canids alleles were compared to Eurasian wolves, all of which had no coyote ancestry. It found the following proportion of unique alleles: coyotes 5.13% unique; red wolf 4.41%; Algonquin wolves 3.82%; Great Lakes wolves 3.61%; and gray wolves 3.3%. They asserted that the amount of unique alleles in all wolves was lower than expected and does not support an ancient (greater than 250,000 years) unique ancestry for any of the species. The authors contended that the proportion of unique alleles and ratio of wolf / coyote ancestry findings matched the south to north disappearance of the wolf due to European colonization since the 18th century and the resulting loss of habitat. Bounties led to the extirpation of wolves initially in the southeast, and as the wolf population declined wolf–coyote admixture increased. Later, this process occurred in the Great Lakes region and then eastern Canada with the influx of coyotes replacing wolves, followed by the expansion of coyotes and their hybrids. The Great Lakes and Algonquin wolves largely reflect lineages that have descendants in the modern wolf and coyote populations, but also reflect a distinct gray wolf ecotype which may have descendants in the modern wolf populations. As a result of these findings, the
American Society of Mammalogists The American Society of Mammalogists (ASM) was founded in 1919. Its primary purpose is to encourage the study of mammals, and professions studying them. There are over 4,500 members of this society, and they are primarily professional scientists w ...
recognizes ''Canis lycaon'' as its own species. The proposed timing of the wolf/coyote divergence conflicts with the finding of a coyote-like specimen in strata dated to 1 million years before present. In 2017 a group of canid researchers challenged the 2016 whole-genome DNA study's finding that the red wolf and the eastern wolf were the result of recent coyote–gray wolf hybridization. The group asserts the three-year generation time used to calculate the divergence periods between different species was lower than empirical estimates of 4.7 years. The group also found deficiencies in the previous study's selection of specimens (two representative coyotes were from areas where recent coyote and gray wolf mixing with eastern wolves is known to have occurred), the lack of certainty in the ancestry of the selected Algonquin wolves, and the grouping of Great Lakes and Algonquin wolves together as eastern wolves, despite opposing genetic evidence. As well, they asserted the 2016 study ignored the fact that there is no evidence of hybridization between coyotes and gray wolves. The group also questioned the conclusions of genetic differentiation analysis in the study stating that results showing Great Lakes, Algonquin and red wolves, plus eastern coyotes differentiated from gray and Eurasian wolves were actually more consistent with an ancient hybridization or a distinct cladogenic origin for the red and Algonquin wolves than of a recent hybrid origin. The group further asserted that the levels of unique alleles for red and Algonquin wolves found the 2017 study were high enough to reveal a high degree of evolutionary distinctness. Therefore, the group argues that both the red wolf and the eastern wolf remain genetically distinct North American taxa. This was rebutted by the authors of the earlier study.


Wolf genome

Genetic studies relating to wolves or dogs have inferred phylogenetic relationships based on the only reference genome available: that of the dog breed called the Boxer. In 2017, the first reference genome of the wolf ''Canis lupus lupus'' was mapped to aid future research. In 2018, a study looked at the genomic structure and admixture of North American wolves, wolf-like canids, and coyotes using specimens from across their entire range that mapped the largest dataset of nuclear genome sequences and compared these against the wolf reference genome. The study supports the findings of previous studies that North American gray wolves and wolf-like canids were the result of complex gray wolf and coyote mixing. A polar wolf from Greenland and a coyote from Mexico represented the purest specimens. The coyotes from Alaska, California, Alabama, and Quebec show almost no wolf ancestry. Coyotes from Missouri, Illinois, and Florida exhibit 5–10% wolf ancestry. There was 40%:60% wolf to coyote ancestry in red wolves, 60%:40% in eastern wolves, and 75%:25% in the Great Lakes wolves. There was 10% coyote ancestry in Mexican wolves, 5% in Pacific Coast and Yellowstone wolves, and less than 3% in Canadian archipelago wolves. The study indicates that the genomic ancestry of red, eastern and Great Lakes wolves were the result of admixture between modern gray wolves and modern coyotes. This was then followed by development into local populations. Individuals within each group showed consistent levels of coyote to wolf inheritance, indicating that this was the result of relatively ancient admixture. The eastern wolf as found in Algonquin Provincial Park is genetically closely related to the Great Lakes wolf as found in Minnesota and Isle Royale National Park in Michigan. If a third canid had been involved in the admixture of the North American wolf-like canids, then its genetic signature would have been found in coyotes and wolves, which it has not. Later in 2018, a study based on a much smaller sample of 65,000 SNPs found that although the eastern wolf carries regional gray wolf and coyote
alleles An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chrom ...
(gene variants), it also exhibits some alleles that are unique and therefore worthy of conservation.


Ancestor

The software that is currently used to conduct whole genome analysis for evidence of hybridization does not distinguish between past and present hybridization. In 2021, an mDNA analysis of modern and extinct North American wolf-like canines indicates that the extinct Late Pleistocene Beringian wolf was the ancestor of the southern wolf
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
, which includes the Mexican wolf and the Great Plains wolf. The Mexican wolf is the most ancestral of the gray wolves that live in North America today. The modern coyote appeared around 10,000 years ago. The most genetically basal coyote mDNA clade pre-dates the
Last Glacial Maximum The Last Glacial Maximum (LGM), also referred to as the Late Glacial Maximum, was the most recent time during the Last Glacial Period that ice sheets were at their greatest extent. Ice sheets covered much of Northern North America, Northern Eu ...
and is a haplotype that can only be found in the Eastern wolf. This implies that the large, wolf-like Pleistocene coyote was the ancestor of the Eastern wolf. Further, another ancient haplotype detected in the Eastern wolf can be found only in the Mexican wolf. The authors propose that Pleistocene coyote and Beringian wolf admixture led to the Eastern wolf long before the arrival of the modern coyote and the modern wolf.


Description and ecology


Description

Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
was told that there were two types of wolf living in the Catskill Mountains, one being a lightly-built, greyhound-like animal that pursued deer, and the other being a bulkier, shorter-legged wolf. The eastern wolf's fur is typically of a grizzled grayish-brown coloration, mixed with cinnamon. The flanks and chest are rufous or creamy, while the nape, shoulder and tail region are a mix of black and gray. Unlike gray wolves, eastern wolves rarely produce
melanistic The term melanism refers to black pigment and is derived from the gr, μελανός. Melanism is the increased development of the dark-colored pigment melanin in the skin or hair. Pseudomelanism, also called abundism, is another variant of pi ...
individuals. The first documented all-black eastern wolf was found to have been an eastern wolf–gray wolf hybrid. Like the red wolf, the eastern wolf is intermediate in size between the coyote and gray wolf, with females weighing on average and males . Like the gray wolf, its average lifespan is 3–4 years, with a maximum of 15 years. Their physical sizes that sets them intermediate between gray wolves and coyotes are actually believed to be more related to their adaptations to an environment with predominately medium-sized prey (similar to the case with the Mexican wolf in the southwestern US) rather than their close relationship to red wolves and coyotes.


Ecology and behaviour

The eastern wolf primarily targets small to medium-sized prey items like
white-tailed deer The white-tailed deer (''Odocoileus virginianus''), also known as the whitetail or Virginia deer, is a medium-sized deer native to North America, Central America, and South America as far south as Peru and Bolivia. It has also been introduced t ...
and
beaver Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers a ...
s, unlike the gray wolf, which can effectively hunt large
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
s like
caribou Reindeer (in North American English, known as caribou if wild and ''reindeer'' if domesticated) are deer in the genus ''Rangifer''. For the last few decades, reindeer were assigned to one species, ''Rangifer tarandus'', with about 10 subspe ...
, elk,
moose The moose (in North America) or elk (in Eurasia) (''Alces alces'') is a member of the New World deer subfamily and is the only species in the genus ''Alces''. It is the largest and heaviest extant species in the deer family. Most adult ma ...
and
bison Bison are large bovines in the genus ''Bison'' (Greek: "wild ox" (bison)) within the tribe Bovini. Two extant and numerous extinct species are recognised. Of the two surviving species, the American bison, ''B. bison'', found only in North A ...
. Despite being carnivores, packs in
Voyageurs National Park Voyageurs National Park is an American national park in northern Minnesota near the city of International Falls established in 1975. The park's name commemorates the ''voyageurs''—French-Canadian fur traders who were the first European settle ...
forage for
blueberries Blueberries are a widely distributed and widespread group of perennial flowering plants with blue or purple berries. They are classified in the section ''Cyanococcus'' within the genus ''Vaccinium''. ''Vaccinium'' also includes cranberries, b ...
in much of July and August, when the berries are in season. Packs carefully avoid each other; only lone wolves sometimes enter other packs' territories. The average territory ranges between 110–185 km², and the earliest age of dispersal for young eastern wolves is 15 weeks, much earlier than gray wolves.


Distribution

The past range of the eastern wolf included southern Quebec, most of Ontario, the Great Lakes states, New York State and New England. Today, the Great Lakes wolf is generally found in the northern halves of Minnesota and Wisconsin, the Upper Peninsula of Michigan, southeastern Manitoba and northern Ontario, and the Algonquin wolf inhabits central and eastern Ontario as well as southwestern Quebec north of the St. Lawrence River. Algonquin wolves are particularly concentrated in Algonquin Provincial Park and other nearby protected areas, such as Killarney, Kawartha Highlands and Queen Elizabeth II Wildlands Provincial Parks and recent surveys also reveal small numbers of Algonquin wolves in the southern areas of northeastern Ontario and northwestern Ontario as far west as the Lake of the Woods near the border with Manitoba, where there is some mixing with Great Lakes wolves, and into southcentral Ontario, where there is some mixing with eastern coyotes. There are some reports of eastern wolf sightings and of wolves being shot by hunters in Quebec south of the St. Lawrence River, New Brunswick, New York State, northern Vermont and Maine. For example, DNA results of a canid killed near
Cherry Valley, New York Cherry Valley is a town in Otsego County, New York, United States. According to the 2020 US census, the village of Cherry Valley had a population of 487. However, the town has a much higher population. Within the town of Cherry Valley is a vill ...
in 2021 initially pointed to it being an eastern coyote, but a recent statement by the
New York State Department of Environmental Conservation The New York State Department of Environmental Conservation (informally referred to as NYSDEC, DEC, EnCon or NYSENCON) is a department of New York state government. The department guides and regulates the conservation, improvement, and protection ...
(DEC) confirms the animal was a wolf, with most of its ancestry matching wolves in Michigan; the DEC also has not confirmed or denied a breeding population within the state. Certain proponents of wolf recolonization state that wolves are already established in New York and New England, and have naturally dispersed from Canada by crossing the frozen St. Lawrence River.


History, hybridization and conservation

Mitochondrial DNA indicates that before the arrival of Europeans, eastern wolves may have numbered at 64,500 to 90,200 individuals. In 1942 it was believed that, before European settlement, the wolf had ranged throughout the wooded and open areas of eastern North America from what is now southern Quebec westward to the Great Plains and towards the Southeastern Woodlands (the southern extent was uncertain, but was believed to around what is now Kentucky, Tennessee and North Carolina).Glover, A. (1942)
''Extinct and vanishing mammals of the western hemisphere, with the marine species of all the oceans''
American Committee for International Wild Life Protection, pp. 210–217.
The region's indigenous human populations did not fear eastern wolves, though they did occasionally catch them in traps, and their bones occur in native shell heaps. Early European settlers often kept their livestock on eastern wolf-free outer islands, though animals kept in pasture on the mainland were vulnerable, to the point that a campaign against eastern wolves was launched in the early years of the
Plymouth Plymouth () is a port city status in the United Kingdom, city and unitary authority in South West England. It is located on the south coast of Devon, approximately south-west of Exeter and south-west of London. It is bordered by Cornwall to ...
and Massachusetts Bay Colonies, in which both settlers and natives participated. A
bounty Bounty or bounties commonly refers to: * Bounty (reward), an amount of money or other reward offered by an organization for a specific task done with a person or thing Bounty or bounties may also refer to: Geography * Bounty, Saskatchewan, a g ...
system was put into effect, offering higher rewards for adult wolves, with their heads exposed on hooks in meetinghouses. Nevertheless, wolves were still plentiful enough in New England in the early 18th century to warrant the settlers of
Cape Cod Cape Cod is a peninsula extending into the Atlantic Ocean from the southeastern corner of mainland Massachusetts, in the northeastern United States. Its historic, maritime character and ample beaches attract heavy tourism during the summer mont ...
discussing the building of a high fence between
Sandwich A sandwich is a food typically consisting of vegetables, sliced cheese or meat, placed on or between slices of bread, or more generally any dish wherein bread serves as a container or wrapper for another food type. The sandwich began as a po ...
and Wareham to keep them out of grazing lands. The scheme failed, though the settlers continued to utilize wolf pits, a wolf trapping technique learned from the region's indigenous peoples. Eastern wolf numbers declined noticeably shortly before and after the
American Revolution The American Revolution was an ideological and political revolution that occurred in British America between 1765 and 1791. The Americans in the Thirteen Colonies formed independent states that defeated the British in the American Revoluti ...
, particularly in
Connecticut Connecticut () is the southernmost state in the New England region of the Northeastern United States. It is bordered by Rhode Island to the east, Massachusetts to the north, New York (state), New York to the west, and Long Island Sound to the ...
, where the wolf bounty was repealed in 1774. Eastern wolf numbers, however, were still high enough to cause concern in the more sparsely populated areas of southern
New Hampshire New Hampshire is a U.S. state, state in the New England region of the northeastern United States. It is bordered by Massachusetts to the south, Vermont to the west, Maine and the Gulf of Maine to the east, and the Canadian province of Quebec t ...
and
Maine Maine () is a state in the New England and Northeastern regions of the United States. It borders New Hampshire to the west, the Gulf of Maine to the southeast, and the Canadian provinces of New Brunswick and Quebec to the northeast and nor ...
, with wolf hunting becoming a regular occupation among settlers and natives alike. By the early 19th century, few eastern wolves remained in southern New Hampshire and
Vermont Vermont () is a U.S. state, state in the northeast New England region of the United States. Vermont is bordered by the states of Massachusetts to the south, New Hampshire to the east, and New York (state), New York to the west, and the Provin ...
. Prior to the establishment of
Algonquin Provincial Park Algonquin Provincial Park is a provincial park located between Georgian Bay and the Ottawa River in Ontario, Canada, mostly within the Unorganized South Part of Nipissing District. Established in 1893, it is the oldest provincial park in Can ...
in 1893, the eastern wolf was common in central
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central Ca ...
and the Algonquin Highlands. It continued to persist throughout the late 1800s, despite extensive logging and efforts by park rangers to eliminate it, largely due to the sustaining influence of plentiful prey items like deer and beaver. By the mid-1900s, there were as many as 55 eastern wolf packs in the park, with an average of 49 wolves being killed annually between 1909–1958, until they were given official protection by the
Ontario government The government of Ontario (french: Gouvernement de l'Ontario) is the body responsible for the administration of the Canadian province of Ontario. A constitutional monarchy, the Crown—represented in the province by the lieutenant governor� ...
in 1959, by which time the eastern wolf population in and around the park had been reduced to 500–1,000 individuals.Algonquin Wolf Advisory Group (2000).
The Wolves of Algonquin Provincial Park: A Report to the Minister of Natural Resources
', Algonquin Wolf Advisory Group, Ontario
Nevertheless, between 1964–1965, 36% of the park's wolf population was culled by researchers trying to understand the reproduction and age structure of the population. This cull coincided with the expansion of coyotes into the park, and lead to an increase in eastern wolf–coyote hybridization. Introgression of gray wolf genes into the eastern wolf population also occurred across northern and eastern Ontario, into
Manitoba Manitoba ( ) is a Provinces and territories of Canada, province of Canada at the Centre of Canada, longitudinal centre of the country. It is Canada's Population of Canada by province and territory, fifth-most populous province, with a population o ...
and
Quebec Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirte ...
, as well as into the western Great Lakes states of
Minnesota Minnesota () is a state in the upper midwestern region of the United States. It is the 12th largest U.S. state in area and the 22nd most populous, with over 5.75 million residents. Minnesota is home to western prairies, now given over t ...
,
Wisconsin Wisconsin () is a state in the upper Midwestern United States. Wisconsin is the 25th-largest state by total area and the 20th-most populous. It is bordered by Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake M ...
, and
Michigan Michigan () is a state in the Great Lakes region of the upper Midwestern United States. With a population of nearly 10.12 million and an area of nearly , Michigan is the 10th-largest state by population, the 11th-largest by area, and t ...
. Despite protection within the park boundaries, there was a population decline in the east side of the park between 1987–1999, with an estimated number of 30 packs by 2000. This decline exceeded annual recruitment, and was attributed to human-caused mortality, which mostly occurred when dispersing animals left the park in search of deer during the winter months, and when pack ranges overlapped with park boundaries. By 2001, protection was extended to eastern wolves occurring in the outskirts of the park. By 2012, the genetic composition of the park's eastern wolves was roughly restored to what it was in the mid-1960s than in the 1980s–1990s, when the majority of wolves had large amounts of coyote DNA. In 2013, an experiment which produced hybrids of coyotes and northwestern gray wolves in captivity using artificial insemination contributed more information to the controversy surrounding the eastern wolf's taxonomy. The purpose of this project was to determine whether the female western coyotes are capable of bearing hybrid western gray wolf-coyote pups, as well as to test the hybrid theory surrounding the origin of the eastern wolves. The resulting six hybrids produced in this captive artificial breeding were later transferred to the Wildlife Science Center of Forest Lake in Minnesota, where their behaviors were studied.


Relationships with humans


In Algonquin folklore

The wolf is prominently portrayed in Algonquin mythology, where it is referred to as ''ma-hei-gan'' or ''nah-poo-tee'' in the
Algonquian languages The Algonquian languages ( or ; also Algonkian) are a subfamily of indigenous American languages that include most languages in the Algic language family. The name of the Algonquian language family is distinguished from the orthographically simi ...
. It is the spirit brother of the Algonquian folk hero Nanabozho, and assisted him in several of his adventures, including thwarting the plots of the malicious anamakqui spirits and assisting him in recreating the world after a worldwide flood.Young, E. R. (1903).
Algonquin Indian Tales
', New York : Eaton & Mains; Cincinnati, Jennings & Pye.


Howling

Since the discovery in 1963 that eastern wolves answered human imitations of their howls, Algonquin Provincial Park began it
Public Wolf Howls attraction
where as many as 2,500 visitors are led on expeditions into areas where eastern wolves were sighted the night before and listen to them answering the park staff's imitation howls. By 2000, 85 Public Wolf Howls had been held, with over 110,000 people having participated. The park considers the attraction as the cornerstone of its wolf education program and credits it with changing public attitudes towards wolves in Ontario.


Attacks on humans

Since the early 1970s, there have been several incidents of bold or aggressive behavior towards humans in Algonquin Provincial Park. Between 1987 and 1996, there were four instances of wolves biting people. The most serious case occurred in 1998 when a male wolf that had been long noted to be unafraid of humans stalked a couple walking their four-year-old daughter in September that year, losing interest when the family took refuge in a trailer. Two days later, the wolf attacked a 19-month-old boy, causing several puncture wounds on his chest and back before being driven off by campers. After the animal was killed later that day, it was found to not be rabid.McNay, Mark E. (2002)
A Case History of Wolf–Human Encounters in Alaska and Canada
''Alaska Department of Fish and Game Wildlife Technical Bulletin''. Retrieved on 2013-10-09.


References


Further reading

*
Ministry of Natural Resources and Forestry The Northern Development, Mines, Natural Resources and Forestry is a government ministry of the Canadian province of Ontario that is responsible for Ontario's provincial parks, forests, fisheries, wildlife, mineral aggregates and the Crown lands ...
. (2018).
DRAFT Recovery Strategy for the Algonquin Wolf in Ontario
'. * Pimlott, D. H., Shannon, J. A. & Kolenosky, G. B. (1969). "The ecology of the timber wolf in Algonquin Provincial Park, Ontario", Research Report (''Wildlife''), no. 87. Ontario, Department of Lands and Forests, Toronto, Ontario * Rutledge, L.
The Eastern Wolf: What we do and do not know...
''Wolf Steward'' (April 2010) * Theberge, J. & Theberge, M. (2013). ''Wolf Country: Eleven Years Tracking the Algonquin Wolves'', Random House LLC


External links



Wolf and Coyote DNA Bank @ Trent University
Eastern wolf survey
{{Taxonbar, from=Q677746 Wolf, Eastern Wolves Canid hybrids Controversial mammal taxa Wolves in the United States Mammals described in 1775