Red wolf

This article is about the species Canis rufus. For other uses, see Red wolf (disambiguation).
Red wolf
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: C. rufus
Binomial name
Canis rufus
(Audubon & Bachman, 1851)[2]
Subspecies
  • † C. r. floridanis
  • † C. r. gregoryi
  • C. r. rufus
Modern range of North American Canis
Synonyms
  • Canis lupus rufus
  • Canis latrans × Canis lupus[3]

The red wolf (Canis rufus[4]/Canis lupus rufus[5]), also known as the Florida wolf or Mississippi Valley wolf[6] is a canid of unresolved taxonomic identity native to the eastern United States.[7][8] It is generally, morphologically, an intermediate between the coyote and gray wolf, and is of a reddish, tawny color.[9][10]

Red wolves were likely 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.[8] 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 captively bred population, which was established in the Point Defiance Zoo & 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 in order to proceed with restoration efforts. In 1987, the captive animals were released into North Carolina's Alligator River National Wildlife Refuge, with a second release taking place two years later in the Great Smoky Mountains National Park.[11]

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.[12] 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 with fragmentation of their social structure from hunting.[13] 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,[4] Although this 2012 review was not universally accepted among relevant authorities,[14] 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.[8][15] Finally, a 2015 genetics study, using the most comprehensive mitochondrial DNA data, Y-chromosome data and genome-wide 127,235 SNP 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".[16] 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,[8] a possibility considered by some researchers.[13]

Taxonomy

Skulls of North American Canis, with red wolf in the center.

The taxonomic reference Mammal Species of the World (2005) does not recognize Canis rufus, however NCBI/Genbank does list it.[17]

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.[18] 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.[19] 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.[20] Nevertheless, some in the scientific community considered it a subspecies of the gray wolf[21] or a hybrid of the gray wolf and the coyote.[22][23]

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.[24][25][26] 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).[18] 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,[27][28] and MSW3[5] listed them both in 2005 as subspecies of the gray wolf.

Comparative image of the red wolf and coyote

In May 2011, an analysis of red wolf, eastern wolf, gray wolf, and dog genomes suggested that the red wolf was 76–80 percent coyote and only 20–24 percent gray wolf, suggesting that the red wolf is actually much more coyote in origin than the eastern wolf. This study analyzed 48,000 SNP and found no evidence for a unique eastern wolf or red wolf species.[12] 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.[29] 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.[30][31] 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.[13] 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.[32] 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.[13] 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.[33] 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.[4]

In a pair of 2012 reports, scientists critical of the May 2011 paper outlined three main points of criticism.[13][32] 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, it remains unclear whether loci important to red wolf genetic variation were actually identified and analyzed (for example, nuclear DNA was not compared in the SNP analysis). 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.[4]

When considered as a full species, three subspecies of red wolf were originally recognized by Goldman;[34] two of these subspecies are extinct. The Florida black wolf (Canis rufus floridanus) (Maine to Florida)[28] has been extinct since 1930 and Gregory's wolf (Canis rufus gregoryi) (south-central United States)[28] was declared functionally extinct in the wild by 1980. The Texas red wolf (Canis rufus rufus), the third surviving 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 in eastern 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.[35] The subspecies designations are essentially moot since two are extinct but the genetic evidence for the three subspecies appears to have been unconvincing anyway.[4]

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 or not if 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 on transferred to the Wildlife Science Center of Forest Lake in Minnesota where their behaviors are now being studied.[36] While this research is still ongoing, the debate over the eastern wolf's taxonomy remained unsettled. Moreover, 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 wolves 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.[14] 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.[8][15]

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.[37][38] Between 150,000–300,000 years ago, the North American branch evolved into the red wolf, eastern wolf and the coyote.[39] Another wolf-like branch migrated to Eurasia and evolved into the gray wolf, which later migrated to North America.[38] It is thought that its original distribution included much of eastern North America, where red wolves were found from Maine south to Florida and in south central US westward to Texas. Records of bounty payments to Wappinger Indians in New York in the middle 18th century confirm its range at least that far north;[40] it's possible that it could have extended as far as extreme eastern Canada.[41]

Physical description and behavior

Melanistic individual at the Audubon Park, New Orleans (1931)

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.[42] Adults measure 136–160 cm (53.5–63 in) in length, and weigh 23–39 kg (50-85 lbs).[42] Its pelage is typically more reddish and sparsely furred than the coyote's and gray wolf's, though melanistic individuals do occur.[9] Its fur is generally tawny to grayish in color, with light markings around the lips and eyes.[10] Like the eastern wolf,[43] 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.[9]

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.[44][45] 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,[44] and reach full size at the age of one year, becoming sexually mature two years later.[10]

Prior to its extinction in the wild, the red wolf's diet consisted of nutria, rabbits and rodents.[18] In contrast, the red wolves from the restored population rely on white-tailed deer, raccoon, nutria and rabbits.[46][47] 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.[48]

Range and habitat

Historical range of the red wolf.

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.[49] But 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.[50] Since 1987, red wolves have been released into northeastern North Carolina where they roam 1.7 million acres.[51] 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.[51] 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.[50] 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.[52] 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 utilized a large suite of habitat types at one time. The last naturally occurring population utilized coastal prairie marshes, swamps and agricultural fields used to grow rice and cotton. However, this environment probably does not typify preferred red wolf habitat. There is evidence that the species was found in highest numbers in the once extensive bottom land river forests and swamps of the southeastern United States. Red wolves re-introduced into northeastern North Carolina have utilized habitat types ranging from agricultural lands to forest/wetland mosaics characterized by an over story 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.[53]

History and conservation

Discovery and persecution

Audubon's depiction of the species (1851).

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.[54] Its reddish color was seen as symbolizing war, as well as all things sacred and beloved.[55] Cherokees generally avoided killing red wolves, as it was believed that such an act would bring about the vengeance of the killed animals' pack-mates.[56]

The red wolf was first described by William Bartram, who encountered it in Florida and wrote of it in his Travels (1791), noting that it was smaller and lighter in color than the gray wolves of Canada and Pennsylvania.[53] 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.[2] 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.[6] However, by 1962, red wolf sightings became increasingly rare, and few tracks or scats were found.[11]

Captive breeding and reintroduction

FWS worker with red wolf pups, August 2002

Formal efforts backed by the 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 southwestern Louisiana and southeastern Texas from 1973 to 1980 by the US Fish and Wildlife Service.[57][58] Measurements, vocalization analyses, and skull X-rays were used to distinguish red wolves from coyotes and red wolf-coyote hybrids.[57] 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 captive breeding 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.[58] These 14 were so closely related that they had the genetic effect of being only eight individuals.

In Dec. 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.[59] The first experimental trans location lasted for 11 days, during which a mated pair of red wolves were monitored day and night with remote telemetry. A second experimental trans location was tried in 1978 with a different mated pair, and they were allowed to remain on the island for close to nine months.[59] 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.[60]

In September 1987, four male-female pairs of red wolves were released in Alligator River National Wildlife Refuge 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).[61]

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, Florida.

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 US Fish and Wildlife Service estimated that there were 300 red wolves remaining in the world, with 207 of those in captivity.[62]

According to the latest Red Wolf Recovery Program First Quarter Report (October–December 2010), the US Fish and Wildlife Service estimates that there are currently 110-130 red wolves in the Red Wolf Recovery Area in North Carolina, however, since not all of the newly bred in the wild red wolves have radio collars, they can only confirm a total of 70 "known" individuals, 26 packs, 11 breeding pairs, and 9 additional individuals not associated with a pack.[51]

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.[51]

Over 30 facilities participate in the red wolf Species Survival Plan and oversee the breeding and reintroduction of over 150 wolves.[63]

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 5-county restoration area in eastern North Carolina.[64] A 2014 court-approved settlement agreement was reached that banned nighttime hunting of coyotes and requires permitting and reporting coyote hunting.[64] In response to the settlement, the North Carolina Wildlife Resources Commission adopted a resolution requesting the United States Fish and Wildlife Service to remove all wild red wolves from private lands, terminate recovery efforts, and declare red wolves extinct in the wild.[65] This resolution came in the wake of a 2014 programmatic review of the red wolf conservation program conducted by The Wildlife Management Institute.[66][67][68] 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.[68] The report stated that the United States Fish and Wildlife Service needed to update its red wolf recovery plan, thoroughly evaluate its strategy for preventing coyote hybridization and increase its public outreach.[69] Since the programmatic review, the United States Fish and Wildlife Service 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.[70] Since then, the wild population has decreased from 100-115 red wolves to 50-65.[71]

In 2014, the United States Fish and Wildlife Service issued the first take permit for a red wolf to a private landowner.[72] Since then, the United States Fish and Wildlife Service issued several other take permits to landowners in the 5-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.[73][74] In response, the Southern Environmental Law Center filed a lawsuit against the United States Fish and Wildlife Service for violating the Endangered Species Act.[75]

References

  1. ↑ Kelly BT, Beyer A & Phillips MK (2008). Canis rufus. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 2008-10-23. Database entry includes justification for why this species is critically endangered.
  2. 1 2 Audubon, J. and J.Bachman.1851.The quadrupeds of North America, Volume 2.New York, NY.p.334.
  3. ↑ VonHolt, BM et al
  4. 1 2 3 4 5 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.
  5. 1 2 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.
  6. 1 2 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.
  7. ↑ Reich, D.E.; Wayne, R.K.; Goldstein, D.B. (1999). "Genetic evidence for a recent origin by hybridization of red wolves". Molecular Ecology 8: 139–144. doi:10.1046/j.1365-294x.1999.00514.x.
  8. 1 2 3 4 5 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: 722–724. doi:10.3390/ani3030722. Retrieved 2015-08-16.
  9. 1 2 3 Paradiso, J. L.; Nowak, R. M. (1972). "Canis rufus" (PDF). Mammalian Species 22: 1–4. doi:10.2307/3503948.
  10. 1 2 3 Woodward, D. W. (1980), The Red Wolf, FWS
  11. 1 2 Hendry, D. (2007). "Red Wolf Restoration: A 20-Year Journey". International Wolf Center 17: 4.
  12. 1 2 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. PMID 21566151.
  13. 1 2 3 4 5 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.
  14. 1 2 Dumbacher, J., Review of Proposed Rule Regarding Status of the Wolf Under the Endangered Species Act, NCEAS (January 2014)
  15. 1 2 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.
  16. ↑ 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: 1–4. doi:10.1098/rsbl.2015.0303. Retrieved 2015-08-16.
  17. ↑ "Canis rufus".
  18. 1 2 3 Atkins, D. (1971). "Evolution of the cerebellum in the genus Canis". J. Mammal 52: 96–97. doi:10.2307/1378435.
  19. ↑ Paradiso, J. (1968). "Canids recently collected in east Texas, with comments on the taxonomy of the red wolf". Am. Midl. Nat. 80: 529–34. doi:10.2307/2423543.
  20. ↑ Ferrell; et al. (1980). "Biochemical markers in species endangered by introgression: The red wolf". Biochem. Genet 18: 39–49. doi:10.1007/bf00504358.
  21. ↑ 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.
  22. ↑ Mech, L.1970. The wolf: The ecology and behavior of an endangered species. Natural History Press, Garden City, NY.
  23. ↑ Wayne, R. (1991). "Mitochondrial DNA analysis supports extensive hybridization of the endangered red wolf (Canis rufus)". Nature 351: 565–68. doi:10.1038/351565a0.
  24. ↑ Phillips, M.; Henry, V. (1992). "Comments on red wolf taxonomy". Conservation Biology 6: 596–599. doi:10.1046/j.1523-1739.1992.06040596.x.
  25. ↑ Nowak, R. (1992). "The red wolf is not a hybrid". Conservation Biology 6: 593–595. doi:10.1046/j.1523-1739.1992.06040593.x.
  26. ↑ 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.
  27. ↑ Koblmuller, S.; Nord, M.; Wayne, R. K.; Leonard, J. (2009). "Origin and Status of the Great Lakes wolf". Molecular Ecology 18: 2313–2326. doi:10.1111/j.1365-294x.2009.04176.x.
  28. 1 2 3 Nowak, R. M. (2002). "The original status of Wolves in Eastern North America". Southeastern Naturalist 1: 95–130. doi:10.1656/1528-7092(2002)001[0095:tosowi]2.0.co;2.
  29. ↑ T. DeLene Beeland (2013). The Secret World of Red Wolves. Chapel Hill, NC: University of North Carolina Press. ISBN 9781469601991.
  30. ↑ "US Fish and Wildlife Species Profile". USFWS. Retrieved 2013-07-08.
  31. ↑ Esch, Mary. "Study: Eastern wolves are hybrids with coyotes". AP News. Retrieved 1 June 2011.
  32. 1 2 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. Retrieved 2013-07-01.
  33. ↑ Stronen, Astrid V.; Nathalie Tessier, Hélène Jolicoeur, Paul C. Paquet, Michel Hénault, Mario Villemure, Brent R. Patterson, Tim Sallows, Gloria Goulet and 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.
  34. ↑ Goldman EA (1937). "The wolves of North America". Journal of Mammalogy 18: 37–45. doi:10.2307/1374306.
  35. ↑ "Red Wolf Recovery Program". U.S. Fish and Wildlife Program. Retrieved 2013-07-02.
  36. ↑ http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0088861
  37. ↑ W. W. Dalquest and G. E. Schultz 1992. Ice Age mammals of northwestern Texas
  38. 1 2 B. Kurten and E. Anderson. 1980. Pleistocene mammals of North America 1-442
  39. ↑ 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: 2156–2166. doi:10.1139/cjz-78-12-2156.
  40. ↑ J. Michael Smith: The Highland King Nimhammaw and the Native Indian Proprietors of Land in Dutchess County, NY: 1712-1765. Hudson River Valley Review
  41. ↑ Wolves of Algonquin Provincial Park
  42. 1 2 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.
  43. ↑ 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
  44. 1 2 Hinton, Joseph W.; Chamberlain, Michael J. (2010-01-01). "Space and Habitat Use by a Red Wolf Pack and Their Pups During Pup-Rearing". The Journal of Wildlife Management 74 (1): 55–58. doi:10.2193/2008-583. ISSN 1937-2817.
  45. ↑ 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.
  46. ↑ 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.
  47. ↑ 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.
  48. ↑ Powell, W. S. (1973). Creatures of North Carolina from Roanoke Island to Purgatory Mountain. North Carolina Historical Review, 50 (2 ), 121-168.
  49. ↑ U.S. Fish and Wildlife Service.1997. Endangered Red Wolves. http://library.fws.gov/Pubs4/endangered_red_wolves.pdf:p7.
  50. 1 2 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]2.0.co;2.
  51. 1 2 3 4 "Current Red Wolf Facts," found on the Red Wolf Recovery web page, http://www.fws.gov/redwolf/index.html, accessed on July 5, 2011.
  52. ↑ National Park Service. "Mammals". Retrieved 2014-08-06.
  53. 1 2 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.
  54. ↑ Camuto, C. (2000), Another Country: Journeying Toward the Cherokee Mountains, University of Georgia Press, ISBN 0820322377
  55. ↑ Panther-Yates, D. N. (2013), Cherokee Clans: An Informal History, Panther`s Lodge Publishers, p. 7.
  56. ↑ Lopez, B. H. (1978), Of Wolves and Men, J. M. Dent and Sons Limited, p. 109, ISBN 0-7432-4936-4
  57. 1 2 Carley,C.1975.Activities and findings of the red wolf recovery program from late 1973 to July 1, 1975. U.S. Fish and Wildlife Service, Albuquerque,NM.
  58. 1 2 McCarley,H. and J. Carley. 1979. Recent changes in distribution and status of red wolves (Canis rufus) Endangered Species Report no.4. U.S. Fish and Wildlife Service, Albuquerque, NM.
  59. 1 2 Carley, Curtis J. 1979. "Report on the Successful Trans location 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.
  60. ↑ U.S. Fish and Wildlife Service Cape Romain NWR, red wolf web page
  61. ↑ USFWS.2010. Red Wolf Recovery Program, 1st Quarter Report, October–December 2010, Manteco, NC.
  62. ↑ U.S. Fish and Wildlife Service. 2007. Red Wolf (Canis rufus) 5-Year Status Review: Summary and Evaluation.
  63. ↑ http://www.pdza.org/page.php?id=296
  64. 1 2 "Protection of Red Wolves | Animal Welfare Institute". awionline.org. Retrieved 2015-12-26.
  65. ↑ "N.C. Wildlife Resources Commission > News > News Article". www.ncwildlife.org. Retrieved 2015-12-26.
  66. ↑ "WMI to Coordinate Comprehensive Review and Evaluation of the Red Wolf Recovery Program". www.wildlifemanagementinstitute.org. Retrieved 2015-12-26.
  67. ↑ 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)
  68. 1 2 "Red Wolf Recovery Program". www.fws.gov. Retrieved 2015-12-26.
  69. ↑ "Wildlife Management Institute Releases New Report on Red Wolf Recovery Program". Defenders of Wildlife. Retrieved 2015-12-26.
  70. ↑ "Service Halts Red Wolf Reintroductions Pending Examination of Recovery Program". www.fws.gov. Retrieved 2015-12-26.
  71. ↑ "2015 Brings No Conclusions On Red Wolf Recovery Program In Eastern NC". wfae.org. Retrieved 2015-12-26.
  72. ↑ "USFWS Grants Landowner Permit to Kill Critically Endangered Red Wolf | Wolf Conservation Center". nywolf.org. Retrieved 2015-12-26.
  73. ↑ "How management rule allows certain red wolf killings". newsobserver. Retrieved 2015-12-26.
  74. ↑ "Sierra Weaver: No defense for death of red wolf". newsobserver. Retrieved 2015-12-26.
  75. ↑ "U.S. Fish and Wildlife faces lawsuit over red wolf program". newsobserver. Retrieved 2015-12-26.

Further reading

Wikispecies has information related to: Canis rufus
Wikimedia Commons has media related to:

External links

This article is issued from Wikipedia - version of the Monday, February 08, 2016. The text is available under the Creative Commons Attribution/Share Alike but additional terms may apply for the media files.