The red wolf (
Canis rufus or
Canis lupus rufus), also known as
Florida black wolf
Florida black wolf or Mississippi Valley wolf, is a canid of
unresolved taxonomic identity native to the eastern United
States. It is generally, morphologically, an intermediate
between the coyote and gray wolf, and is of a reddish, tawny
color. The Red Wolf is a federally listed endangered species of
the United States, and is protected by law. It has been listed by
IUCN as a critically endangered species since 1996. It is
considered the rarest species of wolf and is one of the five most
endangered species of wolf in the world.
Red wolves may have been the first New World wolf species encountered
by European colonists, and were originally distributed throughout the
eastern United States from the Atlantic Ocean to central Texas, and in
the north from the Ohio River Valley, northern
southern New York south to the Gulf of Mexico. The red wolf was
nearly driven to extinction by the mid-1900s due to aggressive
predator-control programs, habitat destruction, and extensive
hybridization with coyotes. By the late 1960s, it occurred in small
numbers in the Gulf Coast of western
Louisiana and eastern Texas.
Fourteen of these survivors were selected to be the founders of a
captive-bred population, which was established in the Point Defiance
Zoo and Aquarium between 1974 and 1980. After a successful
experimental relocation to Bulls Island off the coast of South
Carolina in 1978, the red wolf was declared extinct in the wild in
1980 to proceed with restoration efforts. In 1987, the captive animals
were released into the
Alligator River National Wildlife Refuge
Alligator River National Wildlife Refuge on the
Albemarle Peninsula in North Carolina, with a second release, since
reversed, taking place two years later in the Great Smoky Mountains
National Park. Of 63 red wolves released from 1987–1994, the
population rose to as many as 100–120 individuals in 2012, but has
declined to 50–75 individuals in 2015.
The red wolf's taxonomic status has been a subject of controversy. A
2011 genetic study indicated that it may be a hybrid species between
gray wolves and coyotes. Re-analysis of this study, coupled with a
broader contextual analysis including behavioral, morphological and
additional genetic information, led to arguments that the red wolf is
an independent species, but has suffered from significant
introgression of coyote genes likely due to decimation of red wolf
packs and fragmentation of their social structure due to hunting.
A comprehensive review in October 2012 concluded that the red wolf is
a distinct species which diverged from the coyote alongside the
closely related eastern wolf 150,000–300,000 years ago, Although
this 2012 review was not universally accepted among relevant
authorities, two subsequent reviews of updated research in 2013
and 2014 suggest that the red wolf was once a species distinct from
the gray wolf and coyote. A 2015 genetics study, using the most
comprehensive mitochondrial DNA data, Y-chromosome data and
genome-wide 127,235 single nucleotide polymorphism data, concluded
that "the most parsimonious explanation" is that eastern wolves in
Algonquin Provincial Park
Algonquin Provincial Park are "a distinct remnant entity of a
historical wolf that most likely existed throughout the eastern United
States". This view is supported by the idea that the coyote and
gray wolf did not historically range into the eastern United States,
with current academic debate on red wolf taxonomy shifting to a new
question: whether the eastern wolf and red wolf are conspecific
(belong to the same species), a possibility considered by some
researchers. In contrast, a 2016 study of 28 sequenced canid
genomes concluded that red and eastern wolves have sequences that can
be explained as resulting from gray wolf-coyote hybridization.
1.1 Whole-genome analysis
1.2 Fossil and historic record
2 Physical description and behavior
3 Range and habitat
3.1 History and extirpation in the wild
3.2 Captive breeding and reintroduction
6 Further reading
7 External links
Comparative image of the red wolf (C. rufus) & the coyote (C.
latrans). A 2016 genetic study of canid scats found that despite high
coyote density inside the Red Wolf Experimental Population Area
(RWEPA), hybridization occurs rarely (4% are hybrids).
Skulls of North American Canis, with red wolf in the center
Genbank lists the red wolf as a completely separate species
from the grey wolf. Contrary to what is sometimes asserted, the
Species of the World (2005) does not conclude that
is not a wolf, rather it defines the red wolf as a subspecies of the
grey wolf. The red wolf has been listed by the International
Union for Conservation of Nature as a critically endangered and unique
species since 1996.
The red wolf is also federally listed as a unique and endangered
species of the United States, and is protected by law.
The taxonomy of the red wolf has been debated since before efforts
began in 1973 to save it from extinction. In 1971, Atkins and Dillon
conducted a study on the brains of canids and confirmed the basal
characteristics of the red wolf. Many studies throughout the 1970s
focused on the morphology of the red wolf came to the conclusion that
the red wolf is a distinct species. In 1980, a unique allele was
Canis specimens from within the red wolf range, supporting
the conclusion that the red wolf is a distinct species.
Nevertheless, some in the scientific community considered it a
subspecies of the gray wolf or a hybrid of the gray wolf and the
In 1992, the USFWS conducted an exhaustive review of the literature,
including their own, and concluded that the red wolf is either a
separate species unto itself or a subspecies of the gray
wolf. Many agency reports, books and web pages list the
red wolf as
Canis rufus but genetic research re-opened the debate
about the taxonomy of both the red wolf and Canada's eastern wolf
Canis lupus lycaon). Wilson et al. (2000) concluded that the
eastern wolf and red wolf should be considered as sister taxa due to a
shared common ancestor going back 150,000–300,000 years. In
addition, Wilson et al. further stated that they should be recognized
as distinct species from other North American canids, and not as
subspecies of the gray wolf (
Canis lupus). However, these conclusions
were disputed, and MSW3 listed them both in 2005 as
subspecies of the gray wolf.
In May 2011, an analysis of red wolf, eastern wolf, gray wolf, and dog
genomes suggested that the red wolf was 76–80% coyote and only
20–24% gray wolf, suggesting that the red wolf is actually much more
coyote in origin than the eastern wolf. This study analyzed 48,000
single nucleotide polymorphisms (SNPs) and found no evidence for a
unique eastern wolf or red wolf species. However, X-ray analysis
of the 16 red wolf specimens used in the SNP study were later shown to
be wolf-coyote hybrids via cranial morphometric analysis, rendering
the finding that the red wolf was a gray wolf-coyote hybrid
U.S. Fish and Wildlife Service
U.S. Fish and Wildlife Service (USFWS) still
considers the red wolf a valid species (
Canis rufus) and plans to make
no changes to its recovery program. In 2012, re-analysis of
the 2011 SNP study argued that the original SNP study suffered from
insufficient sampling, and noted that gray wolves do not mate with
coyotes. Another Y-chromosome genetic study in 2012 also argued
that the eastern wolf and red wolf are not hybrids, but rather are a
distinct species from the gray wolf, although eastern and red wolves
do intermix with coyotes. The same authors have argued that the
2011 SNP study finding that red wolves are not an independent species
is flawed and that historical hunting and culling of wolves, leading
to invasion of coyotes into eastern North America, has led to
introgression of coyote mitochondrial and nuclear DNA into fragmented,
decimated eastern wolf packs. They and other authors have
postulated that large populations of eastern and red wolves with
intact social/pack structures are less likely to interbreed with
coyotes. The controversy over the red wolf's species status was
the subject of a comprehensive review of the 2011 and 2012 genetics
studies, which concluded that there are three separate species of wolf
in North America: the red wolf, eastern wolf, and gray wolf.
In a pair of 2012 reports, scientists critical of the May 2011 paper
outlined three main points of criticism. First, the 2011 paper
relied on mtDNA SNPs derived from boxer and poodle genomes and used
these to extrapolate inference about genetic variation within wild
canids across the globe. While it is true that many SNPs were
examined, whether loci important to red wolf genetic variation were
actually identified and analyzed (for example, nuclear DNA was not
compared in the SNP analysis) remains unclear. Second, the study
sampled modern red wolf specimens, and not historic red wolf specimens
from prior to 1900 (when extensive hybridization with coyotes is known
to have taken place), which obfuscates the reliability of the study's
findings. This is important because using historic red wolf genetic
material would have created a baseline genetic profile for the species
against which to test the modern captive-bred specimens. (It is common
knowledge that the captive-bred red wolves are likely slightly
hybridized, but this is a separate issue from interpreting their
species origin as due to hybridization.) Third, the authors lumped
eastern wolf specimens (which critics from Trent University warn are
of unverified origin) with other Great Lakes wolf specimens, and did
not test them separately, which again obfuscated any genetic
differences that may have been present. The controversy over the
eastern wolf's origins is not considered by the scientific community
to be laid to rest, although it may be synonymous with the red
When considered as a full species, three subspecies of red wolf were
originally recognized by Goldman; two of these subspecies are
Florida black wolf
Florida black wolf (
Canis rufus floridanus) (
Florida) has been extinct since 1908 and
Gregory's wolf (Canis
rufus gregoryi) (south-central United States) was declared
functionally extinct in the wild by 1980. The
Texas red wolf (Canis
rufus rufus), the third subspecies, was also functionally extinct in
the wild by 1980, although that status was changed to critically
endangered when captive-bred red wolves from
Texas were reintroduced
North Carolina in 1987. The current status of the
"non-essential/experimental" population in
North Carolina is
endangered and the population numbers around 100 wild
animals. The subspecies designations are essentially moot since
two are extinct but the genetic evidence for the three subspecies
appears to have been unconvincing anyway.
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 and red wolves by comparing them
to both. 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 are now being
studied. However, by 1999, introgression of coyote genes was
recognized as the single greatest threat to wild red wolf recovery and
an adaptive management plan which included coyote sterilization has
been successful, with coyote genes being reduced by 2015 to < 4% of
the wild red wolf population. A 2016 genetic study of canid scats
found that despite high coyote density inside the Red Wolf
Experimental Population Area (RWEPA), hybridization occurs rarely (4%
are hybrids) as a result of the management plan and red wolf mate
High wolf mortality related to anthropogenic causes appeared to be the
main factor limiting wolf dispersal westward from the RWEPA. High
anthropogenic wolf mortality similarly limits expansion of eastern
wolves outside of protected areas in south-eastern Canada.
In 2014, the review of Chambers et al. (2012) which suggested the
eastern wolf should be listed either as a distinct species closely
related to the red wolf or conspecific with the latter became
controversial, forcing the USFWS to commission a peer review of it,
known as NCAES (2014), which took issue with the review. However,
more recent reviews suggest the evidence has "tilted towards a North
American canid assemblage composed of the eastern wolf, red wolf, and
coyote as distinct taxa ... that descended from a common ancestral
canid of North American origin" before arrival of the gray wolf from
Red wolf in forest
In early 2016, an mDNA analysis of 3 ancient (300–1900 years old)
wolf-like samples from the south-eastern United States found that they
grouped with the coyote clade, although their teeth were wolf-like.
The study proposed that the specimens were either coyotes and this
would mean that coyotes had occupied this region continuously rather
than intermittently, a North American evolved red wolf lineage related
to coyotes, or an ancient coyote–wolf hybrid. Ancient hybridization
between wolves and coyotes would likely have been due to natural
events or early human activities, not landscape changes associated
with European colonization because of the age of these samples.
Coyote–wolf hybrids may have occupied the southeastern United States
for a long time, filling an important niche as a large
In July 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.
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 and Great Lakes region wolf are
highly admixed with different proportions of gray wolf and coyote
ancestry. One test indicated a wolf/coyote divergence time of 51,000
years before present that matched other studies indicating that the
extant wolf came into being around this time. Another test indicated
that the red wolf diverged from the coyote between 55,000–117,000
years before present and the Great Lakes region wolf 32,000 years
before present. Other tests and modelling showed various divergence
ranges and the conclusion was a range of less than 6,000 and 117,000
years before present. The study found that coyote ancestry was highest
in red wolves from the southeast of the United States and lowest among
the Great Lakes region wolves.
The theory proposed was that this pattern matched the south-to-north
disappearance of the wolf due to European colonization and its
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 with the influx of coyotes replacing wolves,
followed by the expansion of coyotes and their hybrids across the
wider region. The red wolf may possess some genomic elements
that were unique to gray wolf and coyote lineages from the American
South. 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, and red wolf fossil specimens dating
back 10,000 years ago. The study concluded by stating that because
of the extirpation of gray wolves in the American Southeast, "the
reintroduced population of red wolves in eastern
North Carolina is
doomed to genetic swamping by coyotes without the extensive management
of hybrids as is currently practiced by the USFWS."
In September 2016, the USFWS announced a program of changes to the red
wolf recovery program and "will begin implementing a series of
actions based on the best and latest scientific information". The
service will secure the captive population which is regarded as not
sustainable, determine new sites for additional experimental wild
populations, revise the application of the existing experimental
population rule in North Carolina, and complete a comprehensive
Species Status Assessment.
In 2017 a group of canid researchers challenged the recent finding
that the red wolf and the eastern wolf were the result of recent
coyote-wolf hybridization. The group highlight that no testing had
been undertaken to ascertain the time period that hybridization had
occurred and that, by the previous study's own figures, the
hybridization could not have occurred recently but supports a much
more ancient hybridization. The group found deficiencies in the
previous study's selection of specimens and the findings drawn from
the different techniques used. 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
Fossil and historic record
Paleontological evidence has suggested an origin of the red wolf line
1–2 Ma, branching from a wolf-coyote ancestor, which itself appeared
about 4.9 Ma. Between 150,000–300,000 years ago, the North
American branch evolved into the red wolf, eastern wolf, and the
coyote. Another wolf-like branch migrated to
Eurasia and evolved
into the gray wolf, which later migrated to North America. It is
thought that its original distribution included much of eastern North
America, where red wolves were found from
Maine south to
in the south-central US westward to Texas. Records of bounty payments
Wappinger Indians in New York in the middle 18th century confirm
its range at least that far north and it is possible that it could
have extended as far as extreme eastern Canada.
Physical description and behavior
The red wolf's appearance is typical of the genus Canis, and is
generally intermediate in size between the coyote and gray wolf,
though some specimens may overlap in size with small gray wolves. A
Canis morphometrics conducted in eastern North Carolina
reported that red wolves are morphometrically distinct from coyotes
and hybrids. Adults measure 136–160 cm (53.5–63 in)
in length, and weigh 23–39 kg (50-85 lbs). Its pelage
is typically more reddish and sparsely furred than the coyote's and
gray wolf's, though melanistic individuals do occur. Its fur is
generally tawny to grayish in color, with light markings around the
lips and eyes. Like the eastern wolf, the red wolf has been
compared by some authors to the greyhound in general form, owing to
its relatively long and slender limbs. The ears are also
proportionately larger than the coyote's and gray wolf's. The skull is
typically narrow, with a long and slender rostrum, a small braincase
and a well developed sagittal crest. Its cerebellum is unlike that of
Canis species, being closer in form to that of canids of the
Urocyon genera, thus indicating that the red wolf is one of
the more plesiomorphic members of its genus.
The red wolf is more sociable than the coyote, but less so than the
gray wolf. It mates in January–February, with an average of 6-7 pups
being born in March, April, and May. It is monogamous, with both
parents participating the rearing of young. Denning sites
include hollow tree trunks, along stream banks and the abandoned
earths of other animals. By the age of six weeks, the pups distance
themselves from the den, and reach full size at the age of one
year, becoming sexually mature two years later.
Using long-term data on red wolf individuals of known pedigree, it was
found that inbreeding among first-degree relatives was rare. A
likely mechanism for avoidance of inbreeding is independent dispersal
trajectories from the natal pack. Many of the young wolves spend time
alone or in small non-breeding packs composed of unrelated
individuals. The union of two unrelated individuals in a new home
range is the predominant pattern of breeding pair formation.
Inbreeding is avoided because it results in progeny with reduced
fitness (inbreeding depression) that is predominantly caused by the
homozygous expression of recessive deleterious alleles.
Prior to its extinction in the wild, the red wolf's diet consisted of
rabbits, rodents, and nutria (an introduced species). In contrast,
the red wolves from the restored population rely on white-tailed deer,
raccoon, nutria and rabbits. It should be noted, however, that
white-tailed deer were largely absent from the last wild refuge of red
wolves on the Gulf Coast between
Louisiana (where specimens
were trapped from the last wild population for captive breeding),
which likely accounts for the discrepancy in their dietary habits
listed here. Historical accounts of wolves in the southeast by early
explorers such as William Hilton, who sailed along the Cape Fear River
in what is now
North Carolina in 1644, also note that they ate
Range and habitat
Historical range of the red wolf (Canus rufus)
The originally recognized red wolf range extended throughout the
southeastern United States from the Atlantic and Gulf Coasts, north to
Ohio River Valley
Ohio River Valley and central Pennsylvania, and west to Central
Texas and southeastern Missouri. Research into paleontological,
archaeological and historical specimens of red wolves by Ronald Nowak
expanded their known range to include land south of the Saint Lawrence
River in Canada, along the eastern seaboard, and west to Missouri and
mid-Illinois, terminating in the southern latitudes of Central
Since 1987, red wolves have been released into northeastern North
Carolina, where they roam 1.7 million acres. These lands span five
counties (Dare, Hyde, Tyrrell, Washington, and Beaufort) and include
three national wildlife refuges, a U.S. Air Force bombing range, and
private land. The red wolf recovery program is unique for a large
carnivore reintroduction in that more than half of the land used for
reintroduction lies on private property. Approximately 680,000 acres
(2,800 km2) are federal and state lands, and 1,002,000 acres
(4,050 km2) are private lands. Beginning in 1991, red wolves were
also released into the
Great Smoky Mountains National Park
Great Smoky Mountains National Park in eastern
Tennessee. However, due to exposure to environmental disease
(parvovirus), parasites, and competition (with coyotes as well as
intraspecific aggression), the red wolf was unable to successfully
establish a wild population in the park. Low prey density was also a
problem, forcing the wolves to leave the park boundaries in pursuit of
food in lower elevations. In 1998, the FWS removed the remaining red
wolves in the Great Smoky Mountains National Park, relocating them to
Alligator River National Wildlife Refuge
Alligator River National Wildlife Refuge in eastern North
Carolina. Other red wolves have been released on the coastal
islands in Florida, Mississippi, and
South Carolina as part of the
captive breeding management plan. St. Vincent Island in
currently the only active island propagation site.
Given their wide historical distribution, red wolves probably used a
large suite of habitat types at one time. The last naturally occurring
population used coastal prairie marshes, swamps, and agricultural
fields used to grow rice and cotton. However, this environment
probably does not typify preferred red wolf habitat. Some evidence
shows the species was found in highest numbers in the once extensive
bottom-land river forests and swamps of the southeastern United
States. Red wolves reintroduced into northeastern
North Carolina have
used habitat types ranging from agricultural lands to forest/wetland
mosaics characterized by an overstory of pine and an understory of
evergreen shrubs. This suggests that red wolves are habitat
generalists and can thrive in most settings where prey populations are
adequate and persecution by humans is slight.
History and extirpation in the wild
Audubon's depiction of the species (1851)
Melanistic individual at the
Audubon Park, New Orleans
Audubon Park, New Orleans (1931)
Before the arrival of Europeans, the red wolf featured prominently in
Cherokee mythology, where it is known as wa'ya (ᏩᏯ), said to be
the companion of Kana'ti the hunter and father of the Aniwaya or Wolf
Clan. Cherokees generally avoided killing red wolves, as such an
act was believed to bring about the vengeance of the killed animals'
The red wolf was first described by William Bartram, who encountered
Florida and wrote of it in his Travels (1791), noting that it
was smaller and lighter in color than the gray wolves of
John James Audubon
John James Audubon was the first to describe the red
wolf in detail, giving it the trinomial name of
Canis lupus rufus in
1851. He described it as being more fox-like than the gray wolf, but
retaining the same "sneaking, cowardly, yet ferocious disposition".
Audubon also recounted that red wolves occasionally fed on the
battlefield corpses of fallen soldiers during the Mexican–American
War. The red wolf's full specific rank was first given by Edward
Goldman in 1937, who examined a number of skulls and noted that the
red wolf's skull and dentition differed from those of gray wolves, and
closely approached those of coyotes. He wrote that by the time of
writing, the species may have become extinct in the lower Sonoran
zone, where it was heavily persecuted by ranchmen. Stanley P. Young
noted in 1940 that the red wolf was still common in eastern Texas,
where more than 800 had been caught in 1939, and further cast doubt
over the prospect of fully exterminating them, due to their habit of
living concealed in thickets. However, by 1962, red wolf sightings
became increasingly rare, and few tracks or scats were found.
Captive breeding and reintroduction
USFWS worker with red wolf pups, August 2002
Formal efforts backed by the
U.S. Fish and Wildlife Service
U.S. Fish and Wildlife Service began to
save the red wolf from extinction in 1973, when a captive-breeding
program was established at the Point Defiance Zoological Gardens,
Tacoma, Washington. Four hundred animals were captured from
Louisiana and southeastern
Texas from 1973 to 1980 by the
USFWS. Measurements, vocalization analyses, and skull X-rays
were used to distinguish red wolves from coyotes and red wolf-coyote
hybrids. Of the 400 animals captured, only 43 were believed to be red
wolves and sent to the breeding facility. The first litters were
produced in captivity in May 1977. Some of the pups were determined to
be hybrids, and they and their parents were removed from the program.
Of the original 43 animals, only 17 were considered pure red wolves
and since three were unable to breed, 14 became the breeding stock for
the captive-breeding program. These 14 were so closely related
that they had the genetic effect of being only eight individuals.
In December 1976, two wolves were released onto Cape Romain National
Wildlife Refuge's Bulls Island in
South Carolina with the intent of
testing and honing reintroduction methods. They were not released with
the intent of beginning a permanent population on the island. The
first experimental translocation lasted for 11 days, during which a
mated pair of red wolves was monitored day and night with remote
telemetry. A second experimental translocation was tried in 1978 with
a different mated pair, and they were allowed to remain on the island
for close to nine months. After that, a larger project was
executed in 1987 to reintroduce a permanent population of red wolves
back to the wild in the Alligator River National Wildlife Refuge
(ARNWR) on the eastern coast of North Carolina. Also in 1987, Bulls
Island became the first island breeding site. Pups were raised on the
island and relocated to
North Carolina until 2005.
In September 1987, four male-female pairs of red wolves were released
in ARNWR in northeastern
North Carolina and designated as an
experimental population. Since then, the experimental population has
grown and the recovery area expanded to include four national wildlife
refuges, a Department of Defense bombing range, state-owned lands, and
private lands, encompassing about 1,700,000 acres
In 1989, the second island propagation project was initiated with
release of a population on Horn Island off the Mississippi coast. This
population was removed in 1998 because of a likelihood of encounters
with humans. The third island propagation project introduced a
population on St. Vincent Island, Florida, offshore between Cape San
Blas and Apalachicola, Florida, in 1990, and in 1997, the fourth
island propagation program introduced a population to Cape St. George
Island, Florida, south of Apalachicola.
In 1991, two pairs were reintroduced into the Great Smoky Mountains
National Park, where the last known red wolf was killed in 1905.
Despite some early success, the wolves were relocated to North
Carolina in 1998, ending the effort to reintroduce the species to the
In 2007, the USFWS estimated that 300 red wolves remained in the
world, with 207 of those in captivity.
Interbreeding with the coyote has been recognized as a threat
affecting the restoration of red wolves. Currently, adaptive
management efforts are making progress in reducing the threat of
coyotes to the red wolf population in northeastern North Carolina.
Other threats, such as habitat fragmentation, disease, and
anthropogenic mortality, are of concern in the restoration of red
wolves. Efforts to reduce the threats are presently being
Over 30 facilities participate in the red wolf
Species Survival Plan
and oversee the breeding and reintroduction of over 150 wolves.
In 2012, the Southern Environmental Law Center filed a lawsuit against
North Carolina Wildlife Resources Commission for jeopardizing the
existence of the wild red wolf population by allowing nighttime
hunting of coyotes in the five-county restoration area in eastern
North Carolina. A 2014 court-approved settlement agreement was
reached that banned nighttime hunting of coyotes and requires
permitting and reporting coyote hunting. In response to the
North Carolina Wildlife Resources Commission adopted a
resolution requesting the USFWS to remove all wild red wolves from
private lands, terminate recovery efforts, and declare red wolves
extinct in the wild. This resolution came in the wake of a 2014
programmatic review of the red wolf conservation program conducted by
The Wildlife Management Institute. The Wildlife Management
Institute indicated the reintroduction of the red wolf was an
incredible achievement. The report indicated that red wolves could be
released and survive in the wild, but that illegal killing of red
wolves threatens the long-term persistence of the population. The
report stated that the USFWS needed to update its red wolf recovery
plan, thoroughly evaluate its strategy for preventing coyote
hybridization and increase its public outreach. Since the
programmatic review, the USFWS ceased implementing the red wolf
adaptive management plan that was responsible for preventing red wolf
hybridization with coyotes and allowed the release of captive-born red
wolves into the wild population. Since then, the wild population
has decreased from 100–115 red wolves to 50–65. Despite the
controversy over the red wolf's status as a unique taxon as well as
the USFWS' apparent disinterest towards wolf conservation in the wild,
the vast majority of public comments (including NC residents)
submitted to the USFWS in 2017 over their new wolf management plan
were in favor of the original wild conservation plan.
In 2014, the USFWS issued the first take permit for a red wolf to a
private landowner. Since then, the USFWS issued several other take
permits to landowners in the five-county restoration area. During June
2015, a landowner shot and killed a female red wolf after being
authorized a take permit, causing a public outcry. In
response, the Southern Environmental Law Center filed a lawsuit
against the USFWS for violating the Endangered
By 2016, the red wolf population of
North Carolina had declined to
45-60 wolves. The largest cause of this decline was gunshot.
Showing color variation
Red wolf in a breeding program. Less than 100 remain in the wild.
With radio collar
^ a b c Kelly BT, Beyer A & Phillips MK (2008). "
Canis rufus". The
IUCN Red List
IUCN Red List of Threatened Species. IUCN. 2008: e.T3747A10057394.
doi:10.2305/IUCN.UK.2008.RLTS.T3747A10057394.en. Retrieved 12 January
^ a b Audubon, J. and J.Bachman.1851.The quadrupeds of North America,
Volume 2.New York, NY.p.334.
^ VonHolt, BM et al
^ a b c d e Chambers, Steven M.; Fain, Steven R.; Fazio, Bud; Amaral,
Michael (2012). "An account of the taxonomy of North American wolves
from morphological and genetic analyses". North American Fauna. 77:
1–67. doi:10.3996/nafa.77.0001. Retrieved 2013-07-02.
^ a b Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.;
Species of the World: A Taxonomic and Geographic
Reference (3rd ed.). Johns Hopkins University Press.
ISBN 978-0-8018-8221-0. OCLC 62265494.
^ a b 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. 229-233.
^ Reich, D.E.; Wayne, R.K.; Goldstein, D.B. (1999). "Genetic evidence
for a recent origin by hybridization of red wolves". Molecular
Ecology. 8 (1): 139–144. doi:10.1046/j.1365-294x.1999.00514.x.
^ a b c d e Joseph W. Hinton; Michael J. Chamberlain; David R. Rabon
Jr. (August 2013). "Red Wolf (
Canis rufus) Recovery: A Review with
Suggestions for Future Research". Animals. 3 (3): 722–724.
doi:10.3390/ani3030722. PMC 4494459 . PMID 26479530.
^ a b c Paradiso, J. L.; Nowak, R. M. (1972). "
Canis rufus" (PDF).
Mammalian Species. 22 (22): 1–4. doi:10.2307/3503948.
^ a b c Woodward, D. W. (1980), The Red Wolf, FWS
^ a b "
Red wolf (
Canis rufus)". U.S. Fish & Wildlife Service:
Environmental Conservation Online System.
^ "The 5 Most Endangered Canine Species", Sciam.com, John R. Platt,
May 9, 2013,
^ a b Hendry, D. (2007). "Red Wolf Restoration: A 20-Year Journey".
International Wolf Center. 17: 4.
^ a b Eric M. Gese; Fred F. Knowlton; Jennifer R. Adams; Karen Beck;
Todd K. Fuller; Dennis L. Murray; Todd D. Steury; Michael K. Stoskopf;
Will T. Waddell; Lisette P. Waits (2015). "Managing hybridization of a
recovering endangered species: The red wolf
Canis rufus as a case
study" (PDF). Current Zoology. 61 (1): 191–205.
doi:10.1093/czoolo/61.1.191. Retrieved 2016-02-21.
^ Causes of mortality in wild red wolves (
Canis rufus) 2012–2015
(PDF) (Report). U. S. Fish and Wildlife Service. January 11, 2016.
Retrieved February 21, 2016.
^ a b VonHolt, BM; et al. (12 May 2011). "A genome-wide perspective on
the evolutionary history of enigmatic wolf-like canids". Genome Res.
21 (8): 1294–305. doi:10.1101/gr.116301.110. PMC 3149496 .
^ Rutledge, Linda Y.; Wilson, Paul J.; Klütsch, Cornelya F.C.;
Patterson, Brent R.; White, Bradley N. (2012). "Conservation genomics
in perspective: A holistic approach to understanding
in North America" (PDF). Biological Conservation. 155: 186–192.
doi:10.1016/j.biocon.2012.05.017. Retrieved 2013-07-01.
^ a b Dumbacher, J., Review of Proposed Rule Regarding Status of the
Wolf Under the Endangered
Species Act, NCEAS (January 2014)
^ a b A Comprehensive Review and Evaluation of the Red Wolf (Canis
rufus) Recovery Program (PDF) (Report). Wildlife Management Institute,
Inc. November 14, 2014. p. 171. Retrieved 2015-08-16.
^ a b c d L. Y. Rutledge; S. Devillard; J. Q. Boone; P. A. Hohenlohe;
B. N. White (July 2015). "RAD sequencing and genomic simulations
resolve hybrid origins within North American Canis". Biology Letters.
11 (7): 1–4. doi:10.1098/rsbl.2015.0303. PMC 4528444 .
PMID 26156129. Retrieved 2015-08-16.
^ Morell, V. (2016). "How do you save a wolf that's not really a
wolf?". Science. doi:10.1126/science.aag0699.
^ a b c d Vonholdt, B. M.; Cahill, J. A.; Fan, Z.; Gronau, I.;
Robinson, J.; Pollinger, J. P.; Shapiro, B.; Wall, J.; Wayne, R. K.
(2016). "Whole-genome sequence analysis shows that two endemic species
of North American wolf are admixtures of the coyote and gray wolf".
Science Advances. 2 (7): e1501714. Bibcode:2016SciA....2E1714V.
^ Justin H. Bohling, Justin Dellinger, Justin M. McVey, David T. Cobb,
Christopher E. Moorman and Lisette P. Waits (July 2016). "Describing a
developing hybrid zone between red wolves and coyotes in eastern North
Carolina, USA". Evolutionary Applications. 9 (6): 791–804.
doi:10.1111/eva.12388 (inactive 2017-10-11). PMC 4908465 .
PMID 27330555. Retrieved July 20, 2016. CS1 maint: Uses
authors parameter (link)
Species of the World: A Taxonomic and Geographic Reference,
Volume 2 (NOTE: see the very end of the paragraph on page 577).
Species of the World: A Taxonomic and Geographic Reference,
Volume 3, 2005,
^ Atkins, D. (1971). "Evolution of the cerebellum in the genus Canis".
J. Mammal. 52: 96–97. doi:10.2307/1378435. JSTOR 1378435.
^ Paradiso, J. (1968). "Canids recently collected in east Texas, with
comments on the taxonomy of the red wolf". Am. Midl. Nat. 80 (2):
529–34. doi:10.2307/2423543. JSTOR 2423543.
^ Ferrell; et al. (1980). "Biochemical markers in species endangered
by introgression: The red wolf". Biochem. Genet. 18 (1–2): 39–49.
doi:10.1007/bf00504358. PMID 6930264.
^ Lawrence, B. and W. Bossert. 1975. Relationships of North American
Canis shown by a multiple character analysis of selected populations.
P. 73-86 in M.W. Fox, ed., The wild canids: Their systematic,
behavioral ecology, and evolution. Van Nostrand Reinhold, New York.
^ Mech, L.1970. The wolf: The ecology and behavior of an endangered
species. Natural History Press, Garden City, NY.
^ Wayne, R. (1991). "Mitochondrial DNA analysis supports extensive
hybridization of the endangered red wolf (
Canis rufus)". Nature. 351
(6327): 565–68. Bibcode:1991Natur.351..565W.
^ Phillips, M.; Henry, V. (1992). "Comments on red wolf taxonomy".
Conservation Biology. 6 (4): 596–599.
^ Nowak, R. (1992). "The red wolf is not a hybrid". Conservation
Biology. 6 (4): 593–595.
^ a b Nowak et al. (1995). Another look at wolf taxonomy.
pp 375–397 In L.N. Carbyn, S.H. Fritts, and D.R. Seip, eds.
Ecology and conservation of wolves in a changing world. Canadian
Circumpolar Institute, Edmonton, Alberta.
^ Wilson; et al. (2000). "DNA profiles of the eastern Canadian wolf
and the red wolf for a common evolutionary history independent of the
gray wolf". Canadian Journal of Zoology. 78 (12): 2156–2166.
^ Koblmuller, S.; Nord, M.; Wayne, R. K.; Leonard, J. (2009). "Origin
and Status of the Great Lakes wolf" (PDF). Molecular Ecology. 18 (11):
^ T. DeLene Beeland (2013). The Secret World of Red Wolves. Chapel
Hill, NC: University of
North Carolina Press.
^ "US Fish and Wildlife
Species Profile". USFWS. Retrieved
^ Esch, Mary. "Study: Eastern wolves are hybrids with coyotes". AP
News. Retrieved 1 June 2011.
^ a b Wilson, Paul J.; Rutledge, Linda Y.; Wheeldon, Tyler J.;
Patterson, Brent R.; White, Bradley N. (2012). "Y-chromosome evidence
supports widespread signatures of three-species
Canis hybridization in
eastern North America". Ecology and Evolution. 2 (9): 2325–2332.
doi:10.1002/ece3.301. PMC 3488682 . PMID 23139890.
^ Rutledge, Linda Y.; White, Bradley N.; Row, Jeffrey R.; Patterson,
Brent R. (2012). "Intense harvesting of eastern wolves facilitated
hybridization with coyotes". Ecology and Evolution. 2 (1): 19–33.
doi:10.1002/ece3.61. PMC 3297175 . PMID 22408723.
^ Stronen, Astrid V.; Nathalie Tessier; Hélène Jolicoeur; Paul C.
Paquet; Michel Hénault; Mario Villemure; Brent R. Patterson; Tim
Sallows; Gloria Goulet & François-Joseph Lapointe (2012). "Canid
hybridization: contemporary evolution in human-modified landscapes".
Ecology and Evolution. 2 (9): 2128–2140. doi:10.1002/ece3.335.
PMC 3488665 . PMID 23139873. Retrieved 2013-07-01.
^ Goldman E (1937). "The wolves of North America". Journal of
Mammalogy. 18: 37–45. doi:10.2307/1374306. JSTOR 1374306.
^ a b c Nowak, R. M. (2002). "The original status of Wolves in Eastern
North America". Southeastern Naturalist. 1 (2): 95–130.
^ "Red Wolf Recovery Program". U.S. Fish and Wildlife Program.
^ Mech, L. David; Christensen, Bruce W.; Asa, Cheryl S.; Callahan,
Margaret; Young, Julie K. (25 February 2014). "Production of Hybrids
between Western Gray Wolves and Western Coyotes". PLoS ONE. 9 (2):
e88861. Bibcode:2014PLoSO...988861M. doi:10.1371/journal.pone.0088861.
PMC 3934856 . PMID 24586418 – via PLoS Journals.
^ Justin H. Bohling; Justin Dellinger; Justin M. McVey; David T. Cobb;
Christopher E. Moorman & Lisette P. Waits (July 2016). "Describing
a developing hybrid zone between red wolves and coyotes in eastern
North Carolina, USA". Evolutionary Applications. 9 (6): 791–804.
doi:10.1111/eva.12388. PMC 4908465 . PMID 27330555.
Retrieved July 20, 2016.
^ Benson, J.; B. Patterson & P. Mahoney (2014). "A protected area
influences genotype specific survival and the structure of a Canis
hybrid zone". Ecology. 95 (2): 254–264. doi:10.1890/13-0698.1.
PMID 24669720. Retrieved July 20, 2016.
^ a b Brzeski, Kristin E.; Debiasse, Melissa B.; Rabon, David R.;
Chamberlain, Michael J.; Taylor, Sabrina S. (2016). "Mitochondrial DNA
Variation in Southeastern Pre-Columbian Canids" (PDF). Journal of
Heredity. 107 (3): 287–293. doi:10.1093/jhered/esw002.
PMC 4885236 . PMID 26774058.
^ Roy, Michael S.; Geffen, Eli; Smith, Deborah; Wayne, Robert K.
(1996). "Molecular Genetics of Pre-1940 Red Wolves". Conservation
Biology. 10 (5): 1413–1424.
^ Morell, Virginia (2016). "How do you save a wolf that's not really a
wolf?". Science. doi:10.1126/science.aag0699.
^ Wang, Xiaoming; Tedford, Richard H. (2008). Dogs: Their Fossil
Relatives and Evolutionary History. New York: Columbia University
Press. ISBN 978-0-231-13528-3. OCLC 185095648.
^ Nowak, Ronald M. (2002). "The Original Status of Wolves in Eastern
North America". Southeastern Naturalist. 1 (2): 95–130.
^ Science leads Fish and Wildlife Service to significant changes for
red wolf recovery, Red Wolf Program Review, US Fish and Wildlife
Service, 12 September 2016.
^ Science leads Fish and Wildlife Service to significant changes for
red wolf recovery by Jeff Fleming. Media Release, US Fish and Wildlife
Service, 12 September 2016.
^ Paul A. Hohenlohe, Linda Y. Rutledge, Lisette P. Waits, Kimberly R.
Andrews, Jennifer R. Adams, Joseph W. Hinton, Ronald M. Nowak, Brent
R. Patterson, Adrian P. Wydeven, Paul A. Wilson, Brad N. White (2017).
"Comment on "Whole-genome sequence analysis shows two endemic species
of North American wolf are admixtures of the coyote and gray wolf"".
Science Advances. 3 (6): e1602250. doi:10.1126/sciadv.1602250.
PMC 5462499 . PMID 28630899. CS1 maint: Uses authors
^ Vonholdt, Bridgett M.; Cahill, James A.; Gronau, Ilan; Shapiro,
Beth; Wall, Jeff; Wayne, Robert K. (2017). "Response to Hohenloheet
al". Science Advances. 3 (6): e1701233. doi:10.1126/sciadv.1701233.
PMC 5462503 . PMID 28630935.
^ W. W. Dalquest and G. E. Schultz 1992. Ice Age mammals of
^ a b B. Kurten and E. Anderson. 1980. Pleistocene mammals of North
^ Wilson, P.J.; Grewal, S.; Lawford, I.D.; Heal, J.N.M.; Granacki,
A.G.; Pennock, D.; Theberge, J.B.; Theberge, M.T.; Voigt, D.R.;
Waddell, W.; Paquet, P.C.; Goulet, G.; Cluff, D.; White, B.N. (2000).
"DNA profiles of the eastern Canadian wolf and the red wolf provide
evidence for a common evolutionary history independent of the gray
wolf". Canadian Journal of Zoology. 78 (12): 2156–2166.
^ J. Michael Smith: The Highland King Nimhammaw and the Native Indian
Proprietors of Land in Dutchess County, NY: 1712–1765. Hudson River
^ Wolves of Algonquin Provincial Park
^ a b Hinton, Joseph W.; Chamberlain, Michael J. (2014-08-22).
Canis taxa in eastern North Carolina". Journal of
Mammalogy. 95 (4): 855–861. doi:10.1644/13-MAMM-A-202.
^ Darwin, Charles (1859), "On the Origin of
Species by Means of
Natural Selection, or the Preservation of Favoured Races in the
Struggle for Life", Nature (Full image view 1st ed.), London: John
Murray, 5 (121): 92, Bibcode:1872Natur...5..318B,
doi:10.1038/005318a0, retrieved 2011-03-01
^ a b Hinton, Joseph W.; Chamberlain, Michael J. (2010-01-01). "Space
and Habitat Use by a Red Wolf Pack and Their Pups During Pup-Rearing".
Journal of Wildlife Management. 74 (1): 55–58. doi:10.2193/2008-583.
^ Sparkman, Amanda M.; Adams, Jennifer; Beyer, Arthur; Steury, Todd
D.; Waits, Lisette; Murray, Dennis L. (2011-05-07). "Helper effects on
pup lifetime fitness in the cooperatively breeding red wolf (Canis
rufus)". Proceedings of the Royal Society of London B: Biological
Sciences. 278 (1710): 1381–1389. doi:10.1098/rspb.2010.1921.
ISSN 0962-8452. PMC 3061142 . PMID 20961897.
^ a b Sparkman, AM; Adams, JR; Steury, TD; Waits, LP; Murray, DL (July
2012). "Pack social dynamics and inbreeding avoidance in the
cooperatively breeding red wolf". Behavioral Ecology. 23 (6):
^ Charlesworth D, Willis JH (2009). "The genetics of inbreeding
depression". Nat. Rev. Genet. 10 (11): 783–96. doi:10.1038/nrg2664.
^ Shaw, J.1975. Ecology, behavior and systematic of the red wolf
Canis rufus). Ph.D. dissertation, Yale University, New Haven, CT.
^ Dellinger, Justin A.; Ortman, Brian L.; Steury, Todd D.; Bohling,
Justin; Waits, Lisette P. (2011-12-01). "Food Habits of Red Wolves
during Pup-Rearing Season". Southeastern Naturalist. 10 (4):
731–740. doi:10.1656/058.010.0412. ISSN 1528-7092.
^ McVey, Justin M.; Cobb, David T.; Powell, Roger A.; Stoskopf,
Michael K.; Bohling, Justin H.; Waits, Lisette P.; Moorman,
Christopher E. (2013-10-15). "Diets of sympatric red wolves and
coyotes in northeastern North Carolina". Journal of Mammalogy. 94 (5):
1141–1148. doi:10.1644/13-MAMM-A-109.1. ISSN 0022-2372.
^ Powell, W. S. (1973). Creatures of
North Carolina from Roanoke
Island to Purgatory Mountain.
North Carolina Historical Review, 50 (2
^ U.S. Fish and Wildlife Service.1997. Endangered Red Wolves.
^ a b c "Current Red Wolf Facts," found on the Red Wolf Recovery web
page, http://www.fws.gov/redwolf/index.html, accessed on July 5, 2011.
^ U.S. Fish and Wildlife Service.1997. Endangered Red Wolves.
^ National Park Service. "Mammals". Retrieved 2014-08-06.
^ a b Phillips et al. (2003). "Restoration of the Red Wolf" In Wolves,
Behavior, Ecology and Conservation. Edited by Mech, D. and Boitain, L.
University of Chicago Press, University of Chicago Press.
^ Camuto, C. (2000), Another Country: Journeying Toward the Cherokee
Mountains, University of Georgia Press, ISBN 0-8203-2237-7
^ Lopez, B. H. (1978), Of Wolves and Men, J. M. Dent and Sons Limited,
p. 109, ISBN 0-7432-4936-4
^ Carley, C (1975). Activities and findings of the red wolf recovery
program from late 1973 to July 1, 1975 (Report). Albuquerque, NM: U.S.
Fish and Wildlife Service.
^ McCarley, H. & J. Carley (1979). Recent changes in distribution
and status of red wolves (
Canis rufus) Endangered
Species Report no.4
(Report). Albuquerque, NM: U.S. Fish and Wildlife Service.
^ Red Wolf Recovery/
Species Survival Plan (Report). Atlanta, GA: U.S.
Fish and Wildlife Service. 1990.
^ a b Carley, Curtis J. 1979. "Report on the Successful Translocation
Experiment of Red Wolves (
Canis rufus) to Bulls Island, S.C."
Presentation at the Portland Wolf Symposium, Lewis and Clark College,
Portland, Oregon, August 13–17, 1979.
U.S. Fish and Wildlife Service
U.S. Fish and Wildlife Service Cape Romain NWR, red wolf web page
^ USFWS.2010. Red Wolf Recovery Program, 1st Quarter Report,
October–December 2010, Manteco, NC.
^ U.S. Fish and Wildlife Service. 2007. Red Wolf (
Canis rufus) 5-Year
Status Review: Summary and Evaluation.
^ a b "Protection of Red Wolves
Animal Welfare Institute".
awionline.org. Retrieved 2015-12-26.
^ "N.C. Wildlife Resources Commission > News > News Article".
www.ncwildlife.org. Retrieved 2015-12-26.
^ "WMI to Coordinate Comprehensive Review and Evaluation of the Red
Wolf Recovery Program". www.wildlifemanagementinstitute.org. Retrieved
^ Resolution Requesting that the United States Fish and Wildlife
Service Declare the Red Wolf (
Extinct in the Wild and
Terminate the Red Wolf Reintroduction Program in Beaufort, Dare, Hyde,
Tyrrell, and Washington Counties,
North Carolina (January 29, 2015)
^ a b "Red Wolf Recovery Program". www.fws.gov. Retrieved
^ "Wildlife Management Institute Releases New Report on Red Wolf
Recovery Program". Defenders of Wildlife. Retrieved 2015-12-26.
^ "Service Halts Red Wolf Reintroductions Pending Examination of
Recovery Program". www.fws.gov. Retrieved 2015-12-26.
^ "2015 Brings No Conclusions On Red Wolf Recovery Program In Eastern
NC". wfae.org. Retrieved 2015-12-26.
^ "Public Overwhelmingly Supports Protecting Wild Red Wolves".
www.biologicaldiversity.org. Retrieved 2017-09-25.
^ "USFWS Grants Landowner Permit to Kill Critically Endangered Red
Wolf Wolf Conservation Center". nywolf.org. Retrieved
^ "How management rule allows certain red wolf killings".
newsobserver. Retrieved 2015-12-26.
^ "Sierra Weaver: No defense for death of red wolf". newsobserver.
^ "U.S. Fish and Wildlife faces lawsuit over red wolf program".
newsobserver. Retrieved 2015-12-26.
^ Hinton, Joseph W.; White, Gary C.; Rabon, David R.; Chamberlain,
Michael J. (2017). "Survival and population size estimates of the red
wolf". The Journal of Wildlife Management. 81 (3): 417.
Beeland, T. D. (2013). The Secret World of Red Wolves: The Fight to
Save America's Other Wolf. 256 pages. Chapel Hill: University of North
Chambers, S. M.; Fain, S. R.; Fazio, B.; Amaral, M. (2012). "An
account of the taxonomy of North American wolves from morphological
and genetic analyses". North American Fauna. 77: 1–67.
^ R. Nowak, R.M. (1992). "The red wolf is not a hybrid.".
Conservation Biology 6 : 593-595.
Hinton, J. W.; Chamberlain, M. J.; Rabon, D. R. (2013). "Red Wolf
Canis rufus) Recovery: A Review with Suggestions for Future
Research". Animals. 3 (3): 722–744. doi:10.3390/ani3030722.
PMC 4494459 . PMID 26479530.
Nowak, R. M. (1979). North American Quaternary Canis. Unpublished
Monograph, University of Kansas Hays.
Nowak, R. M. (2002). "The original status of wolves in eastern North
America". Southeastern Naturalist. 1 (2): 95–130.
Nowak, R. M. (2003). Chapter 9: Wolf evolution and taxonomy. In D.
Mech & L. Boitani (Eds.), Wolves: Behavior, Ecology, and
Conservation (pp. 239–258). Chicago: University of Chicago
Philips, M. K., Kelly, B., & Henry, G. (2003). Restoration of the
red wolf In D. Mech & L. Boitani (Eds.), "Wolves: Behavior,
Ecology, and Conservation (pp. 272–288). Chicago: University of
^ Roy, M.S., Geffen, E., Smith, D., Ostrander, E.A. & Wayne,
R.K. (1994). "Patterns of differentiation and hybridization in North
American wolflike canids, revealed by analysis of micro satellite
loci.". Molecular Biology and Evolution 11 : 553–570.
^ Roy, M.S., Girman, D.G., Taylor, A.C. & Wayne, R.K.
(1994). "The use of museum specimens to reconstruct the genetic
variability and relationships of extinct populations.". Experientia
50 : 551-557.
L. Y. Rutledge; S. Devillard; J. Q. Boone; P. A. Hohenlohe; B. N.
White (July 2015). "RAD sequencing and genomic simulations resolve
hybrid origins within North American Canis". Biology Letters. 11 (7):
1–4. doi:10.1098/rsbl.2015.0303. PMC 4528444 .
^ Silverstein, A., Silverstein, V. B. & Silverstein, R. A.
(1994). "The Red wolf: endangered in America.". Brookfield: Conn.
^ Wayne, R.K. & Jenks, S.M. (1991). "Mitochondrial DNA
analysis implying extensive hybridization of the endangered red wolf
Canis rufus". Nature 351 : 565-568.
^ Wayne, R.K., Lehman, N., Allard, M.W. & Honeycutt, R.L.
(1992). "Mitochondrial DNA variability of the grey wolf – genetic
consequences of population decline and habitat fragmentation".
Conservation Biology 6: 559-569.
Wildlife Management Institute (2014). A Comprehensive Review and
Evaluation of the Red Wolf (
Canis rufus) Recovery Program. 171 pages.
Wilson, P. J.; Grewal, S.; Lawford, I. D.; Heal, J. N. M.; Granacki,
A. G.; Pennock, D.; et al. (2000). "DNA profiles of the eastern
Canadian wolf and the red wolf provide evidence for a common
evolutionary history independent of the gray wolf". Canadian Journal
of Zoology. 78 (12): 2156–2166. doi:10.1139/cjz-78-12-2156.
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Common genet (G. genetta)
Johnston's genet (G. johnstoni)
Rusty-spotted genet (G. maculata)
Pardine genet (G. pardina)
Aquatic genet (G. piscivora)
King genet (G. poensis)
Servaline genet (G. servalina)
Haussa genet (G. thierryi)
Cape genet (G. tigrina)
Giant forest genet
Giant forest genet (G. victoriae)
African linsang (P. richardsonii)
Leighton's linsang (P. leightoni)
Malabar large-spotted civet
Malabar large-spotted civet (V. civettina)
Large-spotted civet (V. megaspila)
Malayan civet (V. tangalunga)
Large Indian civet
Large Indian civet (V. zibetha)
Small Indian civet
Small Indian civet (V. indica)
Fossa (C. ferox)
Eastern falanouc (E. goudotii)
Western falanouc (E. major)
Malagasy civet (F. fossana)
Ring-tailed mongoose (G. elegans)
Broad-striped Malagasy mongoose
Broad-striped Malagasy mongoose (G. fasciata)
Grandidier's mongoose (G. grandidieri)
Narrow-striped mongoose (M. decemlineata)
Brown-tailed mongoose (S. concolor)
Durrell's vontsira (S. durrelli)
Caniformia (cont. below)
Giant panda (A. melanoleuca)
Sun bear (H. malayanus)
Sloth bear (M. ursinus)
Spectacled bear (T. ornatus)
American black bear
American black bear (U. americanus)
Brown bear (U. arctos)
Polar bear (U. maritimus)
Asian black bear
Asian black bear (U. thibetanus)
Molina's hog-nosed skunk
Molina's hog-nosed skunk (C. chinga)
Humboldt's hog-nosed skunk
Humboldt's hog-nosed skunk (C. humboldtii)
American hog-nosed skunk
American hog-nosed skunk (C. leuconotus)
Striped hog-nosed skunk
Striped hog-nosed skunk (C. semistriatus)
Hooded skunk (M. macroura)
Striped skunk (M. mephitis)
Sunda stink badger
Sunda stink badger (M. javanensis)
Palawan stink badger
Palawan stink badger (M. marchei)
Southern spotted skunk
Southern spotted skunk (S. angustifrons)
Western spotted skunk
Western spotted skunk (S. gracilis)
Eastern spotted skunk
Eastern spotted skunk (S. putorius)
Pygmy spotted skunk
Pygmy spotted skunk (S. pygmaea)
Eastern lowland olingo
Eastern lowland olingo (B. alleni)
Northern olingo (B. gabbii)
Western lowland olingo
Western lowland olingo (B. medius)
Olinguito (B. neblina)
Ring-tailed cat (B. astutus)
Cacomistle (B. sumichrasti)
White-nosed coati (N. narica)
South American coati
South American coati (N. nasua)
Western mountain coati (N. olivacea)
Eastern mountain coati (N. meridensis)
Kinkajou (P. flavus)
Crab-eating raccoon (P. cancrivorus)
Raccoon (P. lotor)
Cozumel raccoon (P. pygmaeus)
Red panda (A. fulgens)
Caniformia (cont. above)
(includes fur seals
and sea lions)
South American fur seal
South American fur seal (A. australis)
Australasian fur seal (A. forsteri)
Galápagos fur seal
Galápagos fur seal (A. galapagoensis)
Antarctic fur seal
Antarctic fur seal (A. gazella)
Juan Fernández fur seal
Juan Fernández fur seal (A. philippii)
Brown fur seal
Brown fur seal (A. pusillus)
Guadalupe fur seal
Guadalupe fur seal (A. townsendi)
Subantarctic fur seal
Subantarctic fur seal (A. tropicalis)
Northern fur seal
Northern fur seal (C. ursinus)
Steller sea lion
Steller sea lion (E. jubatus)
Australian sea lion
Australian sea lion (N. cinerea)
South American sea lion
South American sea lion (O. flavescens)
New Zealand sea lion
New Zealand sea lion (P. hookeri)
California sea lion
California sea lion (Z. californianus)
Galápagos sea lion
Galápagos sea lion (Z. wollebaeki)
Walrus (O. rosmarus)
Hooded seal (C. cristata)
Bearded seal (E. barbatus)
Gray seal (H. grypus)
Ribbon seal (H. fasciata)
Leopard seal (H. leptonyx)
Weddell seal (L. weddellii)
Crabeater seal (L. carcinophagus)
Northern elephant seal
Northern elephant seal (M. angustirostris)
Southern elephant seal
Southern elephant seal (M. leonina)
Mediterranean monk seal
Mediterranean monk seal (M. monachus)
Hawaiian monk seal
Hawaiian monk seal (M. schauinslandi)
Ross seal (O. rossi)
Harp seal (P. groenlandicus)
Spotted seal (P. largha)
Harbor seal (P. vitulina)
Caspian seal (P. caspica)
Ringed seal (P. hispida)
Baikal seal (P. sibirica)
Large family listed below
Large family listed below
Canidae (includes dogs)
Short-eared dog (A. microtis)
Side-striped jackal (C. adustus)
African golden wolf
African golden wolf (C. anthus)
Golden jackal (C. aureus)
Coyote (C. latrans)
Gray wolf (C. lupus)
Black-backed jackal (C. mesomelas)
Red wolf (C. rufus)
Ethiopian wolf (C. simensis)
Crab-eating fox (C. thous)
Maned wolf (C. brachyurus)
Dhole (C. alpinus)
Culpeo (L. culpaeus)
Darwin's fox (L. fulvipes)
South American gray fox
South American gray fox (L. griseus)
Pampas fox (L. gymnocercus)
Sechuran fox (L. sechurae)
Hoary fox (L. vetulus)
African wild dog
African wild dog (L. pictus)
Raccoon dog (N. procyonoides)
Bat-eared fox (O. megalotis)
Bush dog (S. venaticus)
Gray fox (U. cinereoargenteus)
Island fox (U. littoralis)
Bengal fox (V. bengalensis)
Blanford's fox (V. cana)
Cape fox (V. chama)
Corsac fox (V. corsac)
Tibetan sand fox
Tibetan sand fox (V. ferrilata)
Arctic fox (V. lagopus)
Kit fox (V. macrotis)
Pale fox (V. pallida)
Rüppell's fox (V. rueppelli)
Swift fox (V. velox)
Red fox (V. vulpes)
Fennec fox (V. zerda)
African clawless otter
African clawless otter (A. capensis)
Oriental small-clawed otter
Oriental small-clawed otter (A. cinerea)
Sea otter (E. lutris)
Spotted-necked otter (H. maculicollis)
North American river otter
North American river otter (L. canadensis)
Marine otter (L. felina)
Neotropical otter (L. longicaudis)
Southern river otter
Southern river otter (L. provocax)
Eurasian otter (L. lutra)
Hairy-nosed otter (L. sumatrana)
Smooth-coated otter (L. perspicillata)
Giant otter (P. brasiliensis)
Hog badger (A. collaris)
Tayra (E. barbara)
Lesser grison (G. cuja)
Greater grison (G. vittata)
Wolverine (G. gulo)
Saharan striped polecat
Saharan striped polecat (I. libyca)
Striped polecat (I. striatus)
Patagonian weasel (L. patagonicus)
American marten (M. americana)
Yellow-throated marten (M. flavigula)
Beech marten (M. foina)
Nilgiri marten (M. gwatkinsii)
European pine marten
European pine marten (M. martes)
Japanese marten (M. melampus)
Sable (M. zibellina)
Fisher (P. pennanti)
Japanese badger (M. anakuma)
Asian badger (M. leucurus)
European badger (M. meles)
Honey badger (M. capensis)
Bornean ferret-badger (M. everetti)
Chinese ferret-badger (M. moschata)
Javan ferret-badger (M. orientalis)
Burmese ferret-badger (M. personata)
(Weasels and Ferrets)
Amazon weasel (M. africana)
Mountain weasel (M. altaica)
Stoat (M. erminea)
Steppe polecat (M. eversmannii)
Colombian weasel (M. felipei)
Long-tailed weasel (M. frenata)
Japanese weasel (M. itatsi)
Yellow-bellied weasel (M. kathiah)
European mink (M. lutreola)
Indonesian mountain weasel
Indonesian mountain weasel (M. lutreolina)
Black-footed ferret (M. nigripes)
Least weasel (M. nivalis)
Malayan weasel (M. nudipes)
European polecat (M. putorius)
Siberian weasel (M. sibirica)
Back-striped weasel (M. strigidorsa)
Egyptian weasel (M. subpalmata)
American mink (N. vison)
African striped weasel
African striped weasel (P. albinucha)
American badger (T. taxus)
Marbled polecat (V. peregusna)