Red wolf

From Wikipedia, the free encyclopedia
  (Redirected from Red Wolf)
Jump to: navigation, search
Red wolf
Temporal range: Holocene 100,000 years ago – present[1]
Red wolf (4531335218).jpg
Red wolf showing typical coloration
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: C. rufus
Binomial name
Canis rufus
(Audubon & Bachman, 1851)[3]
  • C. r. floridanis
  • C. r. gregoryi
  • C. r. rufus
  • Canis lupus rufus[3][4]
  • C. lupus × C. latrans[5]

The red wolf (Canis rufus[6] or Canis lupus rufus),[7] also known as the Florida black wolf or Mississippi Valley wolf,[8] is a canid native to the southeastern United States of unresolved taxonomic identity.[9][10] Morphologically it is intermediate between the coyote and gray wolf, and is of a reddish, tawny color.[11][12] The Red Wolf is a federally listed endangered species of the United States and is protected by law.[13] It has been listed by IUCN as a critically endangered species since 1996.[2] It is considered the rarest species of wolf and is one of the five most endangered species of canid in the world.[14]

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 Pennsylvania and southern New York south to the Gulf of Mexico.[10] 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 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.[15] Of 63 red wolves released from 1987–1994,[16] the population rose to as many as 100–120 individuals in 2012, but has declined to 40 individuals in 2018.[17]

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.[18] 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.[19] 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,[6] Although this 2012 review was not universally accepted among relevant authorities,[20] 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.[10][21] 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 are "a distinct remnant entity of a historical wolf that most likely existed throughout the eastern United States".[22] 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),[10] a possibility considered by some researchers.[22] 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.[23][24]


Comparative image of the red wolf (C. rufus) & the coyote (C. latrans)

The taxonomy of the red wolf is not clear, with this wolf being regarded as either a species with a distinct lineage,[25] a recent hybrid of the gray wolf and the coyote,[4] an ancient hybrid of the gray wolf and the coyote which warrants species status,[26] or a distinct species that has undergone recent hybridization with the coyote.[27][28]

The mammalogist W. Christopher Wozencraft, writing in Mammal Species of the World (2005), regards the red wolf as a hybrid of the gray wolf and the coyote, but due to its uncertain status compromised by recognizing it as a subspecies of the grey wolf Canis lupus rufus.[4] However, the genetic authority NCBI/Genbank lists the red wolf as the species Canis rufus.[29] The red wolf has been listed by the International Union for Conservation of Nature as a critically endangered and unique species since 1996.[2] The red wolf is also federally listed as a unique and endangered species of the United States, and is protected by law.[13]

Three subspecies of red wolf were originally recognized by Goldman[30] with two of these subspecies now being extinct. The Florida black wolf (Canis rufus floridanus) (Maine to Florida) has been extinct since 1908 and Gregory's wolf (Canis rufus gregoryi) (south-central United States)[1] was declared functionally extinct in the wild by 1980. The Texas red wolf (Canis rufus rufus) was also functionally extinct in the wild by 1980, although that status was changed to endangered when captive-bred red wolves from Texas were reintroduced in eastern North Carolina in 1987.[31]

Taxonomic debate[edit]

The taxonomic status of the red wolf is confused and disagreed. There are four points of view, with the red wolf described as: (1) a distinct species; (2) a subspecies of gray wolf; (3) a descendant of a gray wolf x coyote hybrid and should not be recognized as a separate species; (4) a distinct species that was recently hybridized with the coyote to such an extent that it should no longer be recognized as a separate species.[32]

Fossil evidence[edit]

The paleontologist Ronald M. Nowak notes that the oldest fossil remains of the red wolf are 10,000 years old and were found in Florida near Melbourne, Brevard County, Withlacoochee River, Citrus County, and Devil's Den Cave, Levy County. He notes that there are only a few, but questionable, fossil remains of the gray wolf found in the southeastern states. He proposes that following the extinction of the dire wolf, the coyote appears to have been displaced from the southeast of the US by the red wolf until last century, when the extirpation of wolves allowed the coyote to expand its range. He also proposes that the ancestor of all North American and Eurasian wolves was C. mosbachensis, which lived in the Middle Pleistocene 700,000–300,000 years ago.[1] C. mosbachensis was a wolf that once lived across Eurasia before going extinct. It was smaller than most North American wolf populations and smaller than C. rufus, and has been described as being similar in size to the small Indian wolf, Canis lupus pallipes. He further proposes that C. mosbachensis invaded North America where it became isolated by the later glaciation and there gave rise to C. rufus. In Eurasia, C. mosbachensis evolved into C. lupus, which later invaded North America.[25]:242

Morphological evidence[edit]

Audubon's depiction of the species (1851)
Skulls of North American Canis, with red wolf in the center

In 1771 the English naturalist Mark Catesby referred to Florida and the Carolinas when he wrote that "The Wolves in America are like those of Europe, in shape and colour, but are somewhat smaller." They were described as being more timid and less voracious.[33] In 1791 the American naturalist William Bartram wrote in his book Travels about a wolf which he had encountered in Florida that was larger than a dog, but was black in contrast to the larger yellow-brown wolves of Pennsylvania and Canada.[34][35] In 1851 the naturalists John James Audubon and John Bachman described the "Red Texan Wolf" in detail. They noted that it could be found in Florida and other southeastern states, but it differed from other North American wolves and named it Canis lupus rufus. It was described as being more fox-like than the gray wolf, but retaining the same "sneaking, cowardly, yet ferocious disposition".[3]

In 1905 the mammologist Vernon Bailey referred to the "Texan Red Wolf" with the first use of the name Canis rufus.[36] In 1937 the zoologist Edward Goldman undertook a morphological study of southeastern wolf specimens. He noted that their skulls and dentition differed from those of gray wolves and closely approached those of coyotes. He identified the specimens as all belonging to the one species which he referred to as Canis rufus.[30][37] Goldman then examined a large number of southeastern wolf specimens and identified three subspecies, noting that their colors ranged from black, gray, and cinnamon-buff.[37]

It is difficult to distinguish the red wolf from a red wolf x coyote hybrid.[35] During the 1960s, two studies of the skull morphology of wild Canis in the southeastern states found them to belong to the red wolf, the coyote, or many variations in between. The conclusion was that there has been recent massive hybridization with the coyote.[38][39] In contrast, another 1960s study of Canis morphology concluded that the red wolf, eastern wolf, and domestic dog were closer to the gray wolf than the coyote, while still remaining clearly distinctive from each other. The study regarded these 3 canines as subspecies of the gray wolf. However, the study noted that "red wolf" specimens taken from the edge of their range which they shared with the coyote could not be attributed to any one species because the cranial variation was very wide. The study proposed further research to ascertain if hybridization had occurred.[40][41]

In the late 19th Century, sheep farmers in Kerr County, Texas stated that the coyotes in the region were larger than normal coyotes, and they believed that they were a gray wolf and coyote cross.[42] In 1970, the wolf mammalogist L. David Mech proposed that the red wolf was a hybrid of the gray wolf and coyote, and suggested that it should be taxonomically recognized as C. lupus x C. latrans.[5] However, a 1971 study compared the cerebellum within the brain of 6 Canis species and found that the cerebellum of the red wolf indicated a distinct species, was closest to that of the gray wolf, but in contrast indicated some characteristics that were more primitive than those found in any of the other Canis species.[43] In 2014, a three dimensional morphometrics study of Canis species accepted only 6 red wolf specimens for analysis from those on offer due to the impact of hybridization on the others.[27]

Mitochondrial DNA evidence[edit]

In 1980, a unique allele was found in Canis specimens from within the red wolf range, supporting the conclusion that the red wolf is a distinct species.[44] Nevertheless, some in the scientific community considered it a hybrid of the gray wolf and the coyote[45] or a subspecies of the gray wolf. 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.[46][47][48] 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).[49]

In 2000, an mDNA study indciated that between 150,000–300,000 years ago, the North American branch evolved into the red wolf, eastern wolf, and the coyote. The study proposed that the eastern wolf and red wolf should be considered as sister taxa due to a shared common ancestor, and that these should be recognized as distinct species from other North American canids, and not as subspecies of the gray wolf (Canis lupus). [50] However, these conclusions were disputed,[48][51] and MSW3[7] 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.[18] 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 inaccurate.[52] The 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.[53][54] 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.[22] 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.[55] 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.[56] 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.[57] 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.[6]

In a pair of 2012 reports, scientists critical of the May 2011 paper outlined three main points of criticism.[22][55] 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 wolf.[6]

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.[58] 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.[16] 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 choice/assortative mating.

High wolf mortality related to anthropogenic causes appeared to be the main factor limiting wolf dispersal westward from the RWEPA.[59] High anthropogenic wolf mortality similarly limits expansion of eastern wolves outside of protected areas in south-eastern Canada.[60]

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.[20] 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 Eurasia.[10][21]

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.[61] Coyote–wolf hybrids may have occupied the southeastern United States for a long time, filling an important niche as a large predator.[61][62] 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).[63]

Whole-genome evidence[edit]

Red wolf in forest

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.[24][64] The red wolf may possess some genomic elements that were unique to gray wolf and coyote lineages from the American South.[24] 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,[65] and red wolf fossil specimens dating back 10,000 years ago.[1] 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."[24]

In September 2016, the USFWS announced a program of changes to the red wolf recovery program[66] 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.[67]

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.[26] This was rebutted by the authors of the earlier study.[68]

Physical description and behavior[edit]

Red wolf

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 study of Canis morphometrics conducted in eastern North Carolina reported that red wolves are morphometrically distinct from coyotes and hybrids.[69] Adults measure 136–160 cm (53.5–63 in) in length, and weigh 23–39 kg (50-85 lbs).[69] Its pelage is typically more reddish and sparsely furred than the coyote's and gray wolf's, though melanistic individuals do occur.[11] Its fur is generally tawny to grayish in color, with light markings around the lips and eyes.[12] Like the eastern wolf,[70] 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 other Canis species, being closer in form to that of canids of the Vulpes and Urocyon genera, thus indicating that the red wolf is one of the more plesiomorphic members of its genus.[11]

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.[71][72] 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,[71] and reach full size at the age of one year, becoming sexually mature two years later.[12]

Using long-term data on red wolf individuals of known pedigree, it was found that inbreeding among first-degree relatives was rare.[73] 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.[73] 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.[74]

Prior to its extinction in the wild, the red wolf's diet consisted of rabbits, rodents, and nutria (an introduced species).[75] In contrast, the red wolves from the restored population rely on white-tailed deer, raccoon, nutria and rabbits.[76][77] 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 Texas and 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 deer.[78]

Range and habitat[edit]

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 the Ohio River Valley and central Pennsylvania, and west to Central Texas and southeastern Missouri.[79] 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 Texas.[1]

Since 1987, red wolves have been released into northeastern North Carolina, where they roam 1.7 million acres.[80] 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.[80] 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 in eastern Tennessee.[81] 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 in eastern North Carolina.[82] 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 Florida is 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.[35]

Extirpation in the wild[edit]

Melanistic individual at the 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.[83] Cherokees generally avoided killing red wolves, as such an act was believed to bring about the vengeance of the killed animals' pack-mates.[84]

In 1940 the biologist Stanley P. Young noted that the red wolf was still common in eastern Texas, where more than 800 had been caught in 1939 because of their attacks on livestock. He did not believe that they could be exterminated because of their habit of living concealed in thickets.[85] In 1962 a study of skull morphology of wild Canis in the states of Arkansas, Louisiana, Oklahoma, and Texas indicated that the red wolf existed in only a few populations due to hybridization with the coyote. The explanation was that either the red wolf could not adapt to changes to its environment due to human land-use along with its accompanying influx of competing coyotes from the west, or that the red wolf was being hybridized out of existence by the coyote.[38]

Captive breeding and reintroduction[edit]

USFWS worker with red wolf pups, August 2002

After the passage of the Endangered Species Act in 1973, formal efforts backed by the U.S. Fish and Wildlife Service began to save the red wolf from extinction, when a captive-breeding program was established at the Point Defiance Zoological Gardens, Tacoma, Washington. Four hundred animals were captured from southwestern Louisiana and southeastern Texas from 1973 to 1980 by the USFWS.[86][87] 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.[88] 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.[89] 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.[89] 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.[90]

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 (6,900 km2).[91]

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 park.

In 2007, the USFWS estimated that 300 red wolves remained in the world, with 207 of those in captivity.[92]

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 explored.[80]

Over 30 facilities participate in the red wolf Species Survival Plan and oversee the breeding and reintroduction of over 150 wolves.[93]

In 2012, the Southern Environmental Law Center filed a lawsuit against the 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.[94] A 2014 court-approved settlement agreement was reached that banned nighttime hunting of coyotes and requires permitting and reporting coyote hunting.[94] In response to the settlement, 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.[95] This resolution came in the wake of a 2014 programmatic review of the red wolf conservation program conducted by The Wildlife Management Institute.[96][97][98] 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.[98] 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.[99] 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.[100] Since then, the wild population has decreased from 100–115 red wolves to 50–65.[101] 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.[102]

In 2014, the USFWS issued the first take permit for a red wolf to a private landowner.[103] 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.[104][105] In response, the Southern Environmental Law Center filed a lawsuit against the USFWS for violating the Endangered Species Act.[106]

By 2016, the red wolf population of North Carolina had declined to 45-60 wolves. The largest cause of this decline was gunshot.[107]



  1. ^ a b c d e Nowak, Ronald M. (2002). "The Original Status of Wolves in Eastern North America". Southeastern Naturalist. 1 (2): 95–130. doi:10.1656/1528-7092(2002)001[0095:TOSOWI]2.0.CO;2. 
  2. ^ a b c Kelly BT, Beyer A & Phillips MK (2008). "Canis rufus". The IUCN Red List of Threatened Species. IUCN. 2008: e.T3747A10057394. doi:10.2305/IUCN.UK.2008.RLTS.T3747A10057394.en. Retrieved 12 January 2018. 
  3. ^ a b c Audubon, J. and Bachman J. (1851) [The Vivaparous Quadrupeds of North America], Volume 2. New York, NY. p.240.
  4. ^ a b c Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 575–577. ISBN 978-0-8018-8221-0. OCLC 62265494.  url=
  5. ^ a b Mech, L.1970. The wolf: The ecology and behavior of an endangered species. Natural History Press, Garden City, NY. pp. 22-25, 285, 331.
  6. ^ a b c d 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. 
  7. ^ a b Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  8. ^ 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.
  9. ^ 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. PMID 9919703. 
  10. ^ 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 4494459Freely accessible. PMID 26479530. Retrieved 2015-08-16. 
  11. ^ a b c Paradiso, J. L.; Nowak, R. M. (1972). "Canis rufus" (PDF). Mammalian Species. 22 (22): 1–4. doi:10.2307/3503948. JSTOR 3503948. Archived from the original (PDF) on 2014-04-16. 
  12. ^ a b c Woodward, D. W. (1980), The Red Wolf, FWS
  13. ^ a b "Red wolf (Canis rufus)". U.S. Fish & Wildlife Service: Environmental Conservation Online System. 
  14. ^ "The 5 Most Endangered Canine Species",, John R. Platt, May 9, 2013,
  15. ^ Hendry, D. (2007). "Red Wolf Restoration: A 20-Year Journey". International Wolf Center. 17: 4. 
  16. ^ 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. 
  17. ^ "Government: Wild red wolf population could soon be wiped out". Retrieved 2018-04-25. 
  18. ^ 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 3149496Freely accessible. PMID 21566151. 
  19. ^ 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 Canis evolution in North America" (PDF). Biological Conservation. 155: 186–192. doi:10.1016/j.biocon.2012.05.017. Retrieved 2013-07-01. 
  20. ^ a b Dumbacher, J., Review of Proposed Rule Regarding Status of the Wolf Under the Endangered Species Act, NCEAS (January 2014)
  21. ^ 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. 
  22. ^ 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 4528444Freely accessible. PMID 26156129. Retrieved 2015-08-16. 
  23. ^ Morell, V. (2016). "How do you save a wolf that's not really a wolf?". Science. doi:10.1126/science.aag0699. 
  24. ^ 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. doi:10.1126/sciadv.1501714. 
  25. ^ a b R.M. Nowak (2003). "Chapter 9 - Wolf evolution and taxonomy". In Mech, L. David; Boitani, Luigi. Wolves: Behaviour, Ecology and Conservation. University of Chicago Press. pp. 239–258. ISBN 978-0-226-51696-7. 
  26. ^ a b 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 5462499Freely accessible. PMID 28630899. 
  27. ^ a b Schmitt, E; Wallace, S (2014). "Shape Change and Variation in the Cranial Morphology of Wild Canids (Canis lupus, Canis latrans, Canis rufus) Compared to Domestic Dogs (Canis familiaris) Using Geometric Morphometrics". International Journal of Osteoarchaeology. 24: 42. doi:10.1002/oa.1306. 
  28. ^ Fredrickson RJ, Hedrick PW. 2006. Dynamics of Hybridization and Introgression in Red Wolves and Coyotes. Conservation Biology 20(4):1272–1283
  29. ^ "Canis rufus". 
  30. ^ a b Goldman E (1937). "The wolves of North America". Journal of Mammalogy. 18: 37–45. doi:10.2307/1374306. JSTOR 1374306. 
  31. ^ "Red Wolf Recovery Program". U.S. Fish and Wildlife Program. Retrieved 2013-07-02. 
  32. ^ Sheldon, Jennifer W. (2004). "Ch.3-Genus Canis". Wild Dogs: The Natural History of the Non-domestic Canidae. The Blackburn Press. p. 54. ISBN 978-1930665842. 
  33. ^ Catesby, M (1771) "The natural history of Carolina, Florida, and the Bahama Islands". Royal Society London, 2 vols. p.xxvi
  34. ^ Bartram, W. (1791) "Travels through North and South Carolina, Georgia, East and West Florida, the Cherokee Country, the Extensive Territories of the Muscogulges or Creek Confederacy, and the Country of the Chactaws. Containing an Account of the Soil and Natural Productions of Those Regions; Together with Observations on the Manners of the Indians", James & Johnson Philadelphia, p197. The second edition that was printed in London in 1794 can be found here
  35. ^ a b c Phillips, M.; Henry, V.; Kelly, B. (2003). "ch.11-Restoration of the Red Wolf". In Mech, L. D.; Boitani, L. Wolves, Behavior, Ecology and Conservation. University of Chicago Press. pp. 272–274. ISBN 0226516970. 
  36. ^ Bailey, V. (1905) "Biological survey of Texas : Life zones, with characteristic species of mammals, birds, reptiles, and plants. Reptiles, with notes of distribution. Mammals, with notes on distribution, habits and economic importance", North American fauna; no. 25. p.174
  37. ^ a b Young, Stanley P.; Goldman, Edward A. (1944). The Wolves of North America. 2. Dover Publications, New York. pp. 413–477. ISBN 0486211932. 
  38. ^ a b McCarley, H. (1962) The Taxonomic Status of Wild Canis (Canidae) in the South Central United States, The Southwestern Naturalist, Vol. 7, No. 3/4 (Dec. 15, 1962), pp. 227-235
  39. ^ 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. 
  40. ^ Lawrence, B. and Bossert, W. (May, 1967) "Multiple Character Analysis of Canis lupus, latrans, and familiaris, with a Discussion of the Relationships of Canis niger", American Zoologist Vol. 7, No. 2 , pp. 223-232, Oxford University Press
  41. ^ 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.
  42. ^ Nowak, Ronald M. (1979). "North American Quaternary Canis". 6. Monograph of the Museum of Natural History, University of Kansas: 39. doi:10.5962/bhl.title.4072. ISBN 978-0-89338-007-6. Retrieved 1 May 2018. 
  43. ^ Atkins, D. (1971). "Evolution of the cerebellum in the genus Canis". J. Mammal. 52: 96–97. doi:10.2307/1378435. JSTOR 1378435. 
  44. ^ 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. 
  45. ^ 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. doi:10.1038/351565a0. 
  46. ^ Phillips, M.; Henry, V. (1992). "Comments on red wolf taxonomy". Conservation Biology. 6 (4): 596–599. doi:10.1046/j.1523-1739.1992.06040596.x. 
  47. ^ Nowak, R. (1992). "The red wolf is not a hybrid". Conservation Biology. 6 (4): 593–595. doi:10.1046/j.1523-1739.1992.06040593.x. 
  48. ^ 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.
  49. ^ 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. doi:10.1139/z00-158. 
  50. ^ 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. doi:10.1139/cjz-78-12-2156. 
  51. ^ Koblmuller, S.; Nord, M.; Wayne, R. K.; Leonard, J. (2009). "Origin and Status of the Great Lakes wolf" (PDF). Molecular Ecology. 18 (11): 2313–2326. doi:10.1111/j.1365-294x.2009.04176.x. PMID 19366404. 
  52. ^ T. DeLene Beeland (2013). The Secret World of Red Wolves. Chapel Hill, NC: University of North Carolina Press. ISBN 978-1-4696-0199-1. 
  53. ^ "US Fish and Wildlife Species Profile". USFWS. Retrieved 2013-07-08. 
  54. ^ Esch, Mary. "Study: Eastern wolves are hybrids with coyotes". AP News. Retrieved 1 June 2011. 
  55. ^ 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 3488682Freely accessible. PMID 23139890. Retrieved 2013-07-01. 
  56. ^ 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 3297175Freely accessible. PMID 22408723. Retrieved 2013-07-01. 
  57. ^ 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 3488665Freely accessible. PMID 23139873. Retrieved 2013-07-01. 
  58. ^ 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 3934856Freely accessible. PMID 24586418 – via PLoS Journals. 
  59. ^ 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 4908465Freely accessible. PMID 27330555. Retrieved July 20, 2016. 
  60. ^ 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. 
  61. ^ 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 4885236Freely accessible. PMID 26774058. 
  62. ^ Roy, Michael S.; Geffen, Eli; Smith, Deborah; Wayne, Robert K. (1996). "Molecular Genetics of Pre-1940 Red Wolves". Conservation Biology. 10 (5): 1413–1424. doi:10.1046/j.1523-1739.1996.10051413.x. 
  63. ^ 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 4908465Freely accessible. PMID 27330555. Retrieved July 20, 2016. 
  64. ^ Morell, Virginia (2016). "How do you save a wolf that's not really a wolf?". Science. doi:10.1126/science.aag0699. 
  65. ^ 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. 
  66. ^ 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.
  67. ^ 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.
  68. ^ 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 5462503Freely accessible. PMID 28630935. 
  69. ^ a b Hinton, Joseph W.; Chamberlain, Michael J. (2014-08-22). "Morphometrics of Canis taxa in eastern North Carolina". Journal of Mammalogy. 95 (4): 855–861. doi:10.1644/13-MAMM-A-202. ISSN 0022-2372. 
  70. ^ 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 
  71. ^ 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. ISSN 1937-2817. 
  72. ^ 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 3061142Freely accessible. PMID 20961897. 
  73. ^ 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): 1186–1194. doi:10.1093/beheco/ars099. 
  74. ^ Charlesworth D, Willis JH (2009). "The genetics of inbreeding depression". Nat. Rev. Genet. 10 (11): 783–96. doi:10.1038/nrg2664. PMID 19834483. 
  75. ^ Shaw, J.1975. Ecology, behavior and systematic of the red wolf (Canis rufus). Ph.D. dissertation, Yale University, New Haven, CT.
  76. ^ 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. 
  77. ^ 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. 
  78. ^ Powell, W. S. (1973). Creatures of North Carolina from Roanoke Island to Purgatory Mountain. North Carolina Historical Review, 50 (2 ), 121-168.
  79. ^ U.S. Fish and Wildlife Service.1997. Endangered Red Wolves.
  80. ^ a b c "Current Red Wolf Facts," found on the Red Wolf Recovery web page,, accessed on July 5, 2011.
  81. ^ U.S. Fish and Wildlife Service.1997. Endangered Red Wolves.
  82. ^ National Park Service. "Mammals". Retrieved 2014-08-06. 
  83. ^ Camuto, C. (2000), Another Country: Journeying Toward the Cherokee Mountains, University of Georgia Press, ISBN 0-8203-2237-7
  84. ^ Lopez, B. H. (1978), Of Wolves and Men, J. M. Dent and Sons Limited, p. 109, ISBN 0-7432-4936-4
  85. ^ Young, S. P. (1940) Review of "North American Big Game", Journal of Mammology vol. 21, pp 96-98
  86. ^ 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. 
  87. ^ 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. 
  88. ^ Red Wolf Recovery/Species Survival Plan (Report). Atlanta, GA: U.S. Fish and Wildlife Service. 1990. 
  89. ^ 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.
  90. ^ U.S. Fish and Wildlife Service Cape Romain NWR, red wolf web page
  91. ^ USFWS.2010. Red Wolf Recovery Program, 1st Quarter Report, October–December 2010, Manteco, NC.
  92. ^ U.S. Fish and Wildlife Service. 2007. Red Wolf (Canis rufus) 5-Year Status Review: Summary and Evaluation.
  93. ^ "Archived copy". Archived from the original on 2011-05-24. Retrieved 2011-09-29. 
  94. ^ a b "Protection of Red Wolves | Animal Welfare Institute". Retrieved 2015-12-26. 
  95. ^ "N.C. Wildlife Resources Commission > News > News Article". Retrieved 2015-12-26. 
  96. ^ "WMI to Coordinate Comprehensive Review and Evaluation of the Red Wolf Recovery Program". Retrieved 2015-12-26. 
  97. ^ Resolution Requesting that the United States Fish and Wildlife Service Declare the Red Wolf (Canis rufus) Extinct in the Wild and Terminate the Red Wolf Reintroduction Program in Beaufort, Dare, Hyde, Tyrrell, and Washington Counties, North Carolina (January 29, 2015)
  98. ^ a b "Red Wolf Recovery Program". Retrieved 2015-12-26. 
  99. ^ "Wildlife Management Institute Releases New Report on Red Wolf Recovery Program". Defenders of Wildlife. Retrieved 2015-12-26. 
  100. ^ "Service Halts Red Wolf Reintroductions Pending Examination of Recovery Program". Retrieved 2015-12-26. 
  101. ^ "2015 Brings No Conclusions On Red Wolf Recovery Program In Eastern NC". Retrieved 2015-12-26. 
  102. ^ "Public Overwhelmingly Supports Protecting Wild Red Wolves". Retrieved 2017-09-25. 
  103. ^ "USFWS Grants Landowner Permit to Kill Critically Endangered Red Wolf | Wolf Conservation Center". Archived from the original on 2015-09-11. Retrieved 2015-12-26. 
  104. ^ "How management rule allows certain red wolf killings". newsobserver. Retrieved 2015-12-26. 
  105. ^ "Sierra Weaver: No defense for death of red wolf". newsobserver. Retrieved 2015-12-26. 
  106. ^ "U.S. Fish and Wildlife faces lawsuit over red wolf program". newsobserver. Retrieved 2015-12-26. 
  107. ^ 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. doi:10.1002/jwmg.21206. 

Further reading[edit]

  • 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 Carolina Press.
  • 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. doi:10.3996/nafa.77.0001. 
  • ^  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 4494459Freely accessible. 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. doi:10.1656/1528-7092(2002)001[0095:tosowi];2. 
  • 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 Press.
  • 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 Chicago Press.
  • ^  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 4528444Freely accessible. PMID 26156129. 
  • ^  Silverstein, A., Silverstein, V. B. & Silverstein, R. A. (1994). "The Red wolf: endangered in America.". Brookfield: Conn. Millbrook Press.
  • ^  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. 

External links[edit]