Global Range: (5000-20,000 square km (about 2000-8000 square miles)) Delineation of the range of this wolf is hampered by a paucity of specimens and debate over the taxonomic status of the species (seeNowak 2002). Data presented by Nowak (2002) indicate a range extending from Maine to Florida and eastward (south of the Great Lakes) to Illinois, Oklahoma, and Texas, based on pre-1918 complete skulls (fragmentary archaeological and paleontological material dating back 10,000 years do not change this much). Basically, if the red wold is regarded as a valid species, its range historically was essentially confined to the southeastern United States.
Assuming the traditional view that the red wolf is a valid species and that it still exists in unhybridized form, the species was, until recent reintroductions, extinct in the wild since early 1980s (or mid-1970s, Rennie 1991). Formerly it was believed to have occurred from central Texas eastward to the coasts of Florida and Georgia and north to North Carolina, and along the Mississippi River Valley north to southern Illinois, and occasionally in Mexico. The last remnant population along Texas/Louisiana coast was rendered functionally extinct due to hybridization with coyote.
A reintroduced population now occurs in an area of roughly 6,900 square kilometers in northeastern North Carolina (reintroduction in Great Smoky Mountains National Park failed and has been terminated; Federal Register, 8 October 1998). Propagation populations currently exist on two islands: Bulls Island, Cape Romain National Wildlife Refuge, South Carolina; and St. Vincent National Wildlife Refuge, Florida. Other red wolves exist in many captive-breeding facilities.
Formerly the range of red wolves included most habitats of the southeastern United States, however this species range was reduced in historic times to extreme southeastern Texas and southwestern Louisiana. Presently, red wolves are being reintroduced into areas of their historical range--Alligator River in North Carolina, and the Great Smoky Mountains of Tennessee and North Carolina (Wilson & Ruff 1999, Nowak 1991).
Biogeographic Regions: nearctic (Native )
- Wilson, D., S. Ruff. 1999. The Smithsonian Book of North American Mammals. Washington: Smithsonian Institution Press.
- Nowak, R., J. Paradiso. 1991. Canis rufus: Mammalian Species No. 22. The American Society of Mammologists.
U.S.A. (SE U.S.A., west to central TX)
endemic to a single nation
Regularity: Regularly occurring
Type of Residency: Year-round
Red wolves are distinguished from their nearest relative, Canis lupus, by their smaller size, relatively narrower proportions, longer legs and ears, and shorter fur. Red wolves have a total length between 1000 and 1300 mm, tail length of from 300 to 420 mm, and shoulder height of 660 to 790 mm. Among red wolves, males average 10 percent larger than females. Red wolves usually have upperparts that are a mixture of cinnamon, tawny, and gray or black, while the back is normally blackish. The muzzle and limbs are tawny and the tail is tipped with black. In winter, the reddish element of the pelage is dominant. An annual molt takes place in the summer (Wilson & Ruff 1999, Nowak 1991).
Range mass: 20 to 40 kg.
Average mass: 23.5 kg.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: male larger
Length: 165 cm
Weight: 40900 grams
Size in North America
Range: "1,300-1,700 mm "
Range: 20-38 kg males; 18-30 kg females
Coyote is smaller (total length 105-132 cm), with a narrower nose pad and hind foot. Gray wolf tends to be larger (total length 137-206 cm). However, see taxonomy comments.
Comments: Suitable habitat for this habitat generalist includes upland and lowland forests, shrublands, and coastal prairies and marshes; areas with heavy vegetative cover. Young are born in a den in a hollow log, in a burrow, or in similar secluded sites.
Habitat and Ecology
Before becoming endangered, red wolves inhabited mountains, lowland forests, and wetlands. Presently, red wolves survive mainly as small relict and reintroduced populations in inaccessible swampland and mountainous terrain (Wilson & Ruff 1999, Nowak 1991).
Habitat Regions: temperate ; terrestrial
Terrestrial Biomes: forest ; scrub forest
Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.
Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: No. No populations of this species make annual migrations of over 200 km.
Home ranges variously reported as 65 to 130 square kilometers (Riley and McBride 1975), 117 (males) and 78 square kilometers (females, Carley 1979); 100-200 sq km mentioned by Lowman 1975; varies with conditions.
Comments: Opportunistic. Diet consists of a variety of invertebrates and vertebrates (rabbits, rodents, deer, birds, etc.). Particularly favors marsh rabbits (SYLVILAGUS AQUATICUS), nutria (MYOCASTOR COYPUS), and carrion. Not considered a threat to livestock (does not hunt in packs), but may prey on unattended young calves, pigs, and barnyard fowl (Matthews and Moseley 1990).
Rodents, ungulates, and other small mammals are the main prey of red wolves. The dominant prey species include raccoons, white-tailed deer, swamp rabbits, cottontail rabbits, pigs, rice rats, nutria, and muskrats. Red wolves will also eat carrion. They typically hunt in a particular area for 7 to 10 days, then switch to a different range (Wilson & Ruff 1999, Nowak 1991).
Animal Foods: birds; mammals; carrion
Plant Foods: fruit
Primary Diet: carnivore (Eats terrestrial vertebrates)
Red wolves are important as top predators in the ecosystems in which they live.
Red wolves are primarily preyed on by other canids, including conspecifics from other packs, gray wolves, and coyote as a result of agonistic interactions over territories. Young red wolves may also be taken by other large predators such as alligator, large raptors, and bobcats.
Known prey organisms
Number of Occurrences
Note: For many non-migratory species, occurrences are roughly equivalent to populations.
Estimated Number of Occurrences: 1 - 5
Comments: This species occurs in the wild in one major location (plus two islands that serve as propagation areas).
1 - 250 individuals
Comments: As of 2005, there were about 100 red wolves in the wild on 6,900 square kilometers in northeastern North Carolina. The number of reproductive individuals probably is not greater than 50. The captive population included around 165 wolves.
More social than coyote but less so than gray wolf; typically travels and forages in small family groups or alone. Formerly density probably not more than 1 per 2 sq miles.
Life History and Behavior
Communication and Perception
Red wolves communicate with conspecifics through a complex suite of behavioral, tactile, chemical, and auditory signals. Body language, pheromones, and vocalizations all serve to communicate about social and reproductive status and mood. Social bonding is often acheived through touch. Home ranges are delimited using scent marks.
Communication Channels: visual ; tactile ; acoustic ; chemical
Other Communication Modes: pheromones ; scent marks
Perception Channels: visual ; acoustic
Comments: Primarily nocturnal.
Most individuals live to about 4 years, though one captive individual was recorded at 14 years old (Wilson & Ruff 1999, Nowak 1991).
Status: captivity: 14 (high) years.
Status: wild: 4 years.
Status: wild: 16.0 years.
Status: captivity: 20.0 years.
Status: captivity: 14.0 years.
Mates in January-February. Gestation lasts 60-63 days. Litter of 3-12 (average 6-7) is born in March-May. One litter per year. Sexually mature in 3 years.
The dominant male and female pair are solely able to reproduce within a pack. Other pack members assist in raising young and obtaining food for lactating females.
Mating System: monogamous ; cooperative breeder
Breeding season extends from January to March. The gestation period is 60-63 days, with average litters of 3-6 pups occurring in the spring. However, litters of up to 12 pups can occur.
Breeding season: Breeding occurs between January and March.
Range number of offspring: 1 to 12.
Average number of offspring: 3-6.
Range gestation period: 60 to 62 days.
Average age at sexual or reproductive maturity (female): 1 years.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous
Average number of offspring: 5.
Both males and females participate in rearing the young in the den, as well as other pack members. The young are cared for, nursed, and sheperded through their first year of life.
Parental Investment: altricial ; male parental care ; female parental care ; post-independence association with parents; extended period of juvenile learning
- Wilson, D., S. Ruff. 1999. The Smithsonian Book of North American Mammals. Washington: Smithsonian Institution Press.
- Nowak, R., J. Paradiso. 1991. Canis rufus: Mammalian Species No. 22. The American Society of Mammologists.
National NatureServe Conservation Status
Rounded National Status Rank: N1 - Critically Imperiled
NatureServe Conservation Status
Rounded Global Status Rank: G1 - Critically Imperiled
Reasons: Exterminated from formerly large range in the southeastern U.S.; now exists in one major population in northeastern North Carolina, plus a couple islands used for propagation; intensively monitored and managed; attempted reintroduction in Great Smoky Mountains National Park was unsuccessful.
IUCN Red List Assessment
Red List Category
Red List Criteria
- 2004Critically Endangered
- 1996Critically Endangered
- 1994Endangered(Groombridge 1994)
- 1990Endangered(IUCN 1990)
- 1988Endangered(IUCN Conservation Monitoring Centre 1988)
- 1986Endangered(IUCN Conservation Monitoring Centre 1986)
- 1982Endangered(Thornback and Jenkins 1982)
Date Listed: 03/11/1967
Lead Region: Southeast Region (Region 4)
Where Listed: except where EXPN
Status: Experimental Population, Non-Essential
Date Listed: 11/19/1986
Lead Region: Southeast Region (Region 4)
Where Listed: U.S.A. (portions of NC and TN)
Population location: Entire, except where listed as experimental populations below
Listing status: E
Population location: U.S.A. (portions of NC and TN_see 17.84(c)(9))
Listing status: EXPN
For most current information and documents related to the conservation status and management of Canis rufus , see its USFWS Species Profile
Red wolves have been blamed for depredations on livestock and game. As a result, humans, mainly ranchers, farmers, and government trappers, steadily eliminated populations of red wolves. In 1967, red wolves were listed as endangered and the U.S. Fish and Wildlife Service engaged in a salvage effort to protect remaining populations. Fourteen remaining red wolves were placed in a captive-breeding facility; they have become the founders of the present red wolf population. Currently, 200+ red wolves exist, and reintroductions are occurring in a few areas, including North Carolina and the Great Smoky Mountains.
US Federal List: endangered
CITES: appendix i; no special status
IUCN Red List of Threatened Species: critically endangered
Global Short Term Trend: Relatively stable (=10% change)
Comments: USFWS (1990) categorized the status as "improving," but range has not expanded sigificantly beyond the reintroduction areas.
Global Long Term Trend: Decline of >90%
Comments: Assuming an historical range throughout southeastern North America (e.g., Nowak 2002), the extent of occurrence, area of occupancy, and abundance of this species have undergone a drastic decline over the long term.
Extinct in the Wild by 1980, the Red Wolf was reintroduced by the United States Fish and Wildlife Service (USFWS) in 1987 into eastern North Carolina. The Red Wolf is now common within the reintroduction area of roughly 6,000 kmÂ². However, the species' abundance outside the reintroduction area is unknown.
Degree of Threat: Medium
Comments: Historical decline was due to human persecution (trapping, poisoning), habitat loss, and hybridization with coyote. Attempted reintroduction in Great Smoky Mountians National Park failed, probably due to parovirus and other common canine diseases, internal and external parasites, poor nutrition caused by low food availability, and predation (Federal Register, 8 October 1998). Hybridization with coyotes is an ongoing threat in northeastern North Carolina.
It has been estimated that the Red Wolf population in North Carolina can sustain only one hybrid litter out of every 59 litters (1.7%) to maintain 90% of its genetic diversity for the next 100 years (Kelly et al. 1999). However, prior to learning of this acceptable introgression rate, the introgression rate noted in the reintroduced population was minimally 15% (Kelly et al. 1999) or approximately 900% more than the population can sustain to maintain 90% of its genetic diversity for 100 years. If such levels of hybridization continued beyond 1999, non-hybridized Red Wolves could disappear within 12?24 years (3?6 generations). An adaptive management plan designed to test whether hybridization can be reduced to acceptable levels was initiated in 1999 (Kelly 2000). Initial results from this plan suggest that the intensive management specified in the plan may be effective in reducing introgression rates to acceptable levels (B. Fazio pers. comm.).
In the absence of hybridization, recovery of the Red Wolf and subsequent removal of the species from the U.S. Endangered Species List is deemed possible. It is noteworthy that similar hybridization has been observed in the population of suspected Red Wolf-type wolves in Algonquin Provincial Park, Ontario, Canada. If these wolves are ultimately shown to be Red Wolf-type wolves, this will enhance the conservation status of the species and nearly triple the known number of Red Wolf-type wolves surviving in the wild.
Human induced mortality (vehicles and gunshot) can be significant. However, the threat this mortality represents to the population is unclear. Most vehicle deaths occurred early in the reintroduction and were likely due to naive animals. Nonetheless, the overall impact of these mortality factors will depend on the proportion of the losses attributable to the breeding segment of the population (effective population (Ne) and what proportion of the overall population is lost due to these human factors (both N and Ne)).
Management Requirements: See USFWS (Federal Register, 13 April 1995) for revision of the special rule for nonessential experimental populations in North Carolina and Tennessee.
Global Protection: Few (1-3) occurrences appropriately protected and managed
Comments: A population exists in northeastern North Carolina on three national wildlife refuges, a Department of Defense bombing range, state-owned lands, and private property, spanning a total of 6,100 square kilometers.
Needs: Application of the experimental population designation has been useful in getting support for introductions as part of the recovery process (Parker and Phillips 1991).
See 1989 recovery plan.
The only free-ranging population of Red Wolves exists in north-eastern North Carolina in an area comprised of 60% private land and 40% public land. This area contains three national wildlife refuges (Alligator River NWR, Pocosin Lakes NWR, and Mattamuskeet NWR) which provide important protection to the wolves. Red Wolves or a very closely related taxon may also occupy Algonquin Provincial Park, Ontario, Canada.
A very active recovery programme for the Red Wolf has been in existence since the mid-1970s (USFWS 1990; Phillips et al. 2003), with some measures from as early as the mid-1960s (USFWS, unpubl.). By 1976, a captive breeding programme was established using 17 animals captured in Texas and Louisiana (Carley 1975; USFWS 1990). Of these, 14 became the founders of the current captive breeding programme. In 1977, the first pups were born in the captive programme, and by 1985, the captive population had grown to 65 individuals in six zoological facilities (Parker 1986).
With the species reasonably secure in captivity, the USFWS began reintroducing Red Wolves at the Alligator River National Wildlife Refuge in north-eastern North Carolina in 1987. As of September 2002, 102 animals have been released with a minimum of 281 descendants produced in the wild since 1987. As of September 2002, there is a minimum population of 66 wild Red Wolves in north-eastern North Carolina, with a total wild population believed to be at least 100 individuals. Likewise, at this same time, there is a minimum population of 17 hybrid canids present in north-eastern North Carolina. The 17 known hybrids are sterilized and radio-collared (USFWS, unpubl.).
During 1991 a second reintroduction project was initiated at the Great Smoky Mountains National Park, Tennessee (Lucash et al. 1999). Thirty-seven Red Wolves were released from 1992 to 1998. Of these, 26 either died or were recaptured after straying onto private lands outside the Park (Henry 1998). Moreover, only five of the 32 pups known to have been born in the wild survived but were removed from the wild during their first year (USFWS, unpubl.). Biologists suspect that disease, predation, malnutrition, and parasites contributed to the high rate of pup mortality (USFWS, unpubl.). Primarily because of the poor survival of wild-born offspring, the USFWS terminated the Tennessee restoration effort in 1998 (Henry 1998).
Occurrence in captivity
As of September 2002, there are approximately 175 Red Wolves in captivity at 33 facilities throughout the United States and Canada (USFWS, unpubl.). The purpose of the captive population is to safeguard the genetic integrity of the species and to provide animals for reintroduction. In addition, there are propagation projects on two small islands off the South Atlantic and Gulf Coasts of the U.S. which, through reintroduction of known breeding individuals and capture of their offspring, provide wild-born pups for release into mainland reintroduction projects (USFWS 1990).
Relevance to Humans and Ecosystems
Economic Importance for Humans: Negative
Red wolves were long thought by the public to be a serious threat to livestock. This threat has been grossly exaggerated, though they may occasionally kill domestic animals (Fox 1975).
Economic Importance for Humans: Positive
Red wolves eat many rodents, thus helping to control the populations of these pests (Fox 1975).
Positive Impacts: controls pest population
The red wolf (Canis rufus, formerly Canis lupus rufus), also known as the Florida wolf or Mississippi Valley wolf is a canid native to the Southeastern United States. It is generally, morphologically, an intermediate between the coyote and gray wolf, and is of a reddish, tawny color.
Originally distributed throughout the Southeastern US, 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.
The red wolf's taxonomic status has been a subject of controversy. A 2011 genetic study indicated that it may be a hybrid species between gray wolves and coyotes. Re-analysis of this study coupled with a broader contextual analysis including behavioral, morphological and additional genetic information led to arguments that the red wolf is an independent species but has suffered from significant introgression of coyote genes likely due to decimation of red wolf packs with fragmentation of their social structure from hunting. A comprehensive review in October 2012 concluded that the red wolf is a distinct species which diverged from the coyote alongside the closely related eastern wolf 150,000-300,000 years ago.
The taxonomy of the red wolf has been debated since before efforts began in 1973 to save it from extinction. In 1971, Atkins and Dillon conducted a study on the brains of canids and confirmed the basal characteristics of the red wolf. Many studies throughout the 1970s focused on the morphology of the red wolf came to the conclusion that the red wolf is a distinct species. In 1980, a unique allele was found in Canis specimens from within the red wolf range, supporting the conclusion that the red wolf is a distinct species. Nevertheless, some in the scientific community considered it a subspecies of the gray wolf or a hybrid of the gray wolf and the coyote.
In 1992, the USFWS conducted an exhaustive review of the literature, including their own, and concluded that the red wolf is either a separate species unto itself or a subspecies of the gray wolf. Many agency reports, books and web pages list the red wolf as Canis rufus but genetic research re-opened the debate about the taxonomy of both the red wolf and Canada's eastern wolf (Canis lupus lycaon). Wilson et al. (2000) concluded that the eastern wolf and red wolf should be considered as sister taxa due to a shared common ancestor going back 150,000–300,000 years. In addition, Wilson et al. further stated that they should be recognized as distinct species from other North American canids, and not as subspecies of the gray wolf (Canis lupus). However, these conclusions were disputed, and MSW3 listed them both in 2005 as subspecies of the gray wolf.
In May 2011, an analysis of red wolf, Eastern wolf, gray wolf, and dog genomes suggested that the red wolf was 76–80 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. 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. 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. In 2012, re-analysis of the 2011 SNP study argued that the original SNP study suffered from insufficient sampling and noted that gray wolves do not mate with coyotes. Another Y-chromosome genetic study in 2012 also argued that the eastern wolf and red wolf are not hybrids but rather are a distinct species from the gray wolf, although eastern and red wolves do intermix with coyotes. The same authors have argued that the 2011 SNP study finding that red wolves are not an independent species is flawed and that historical hunting and culling of wolves, leading to invasion of coyotes into eastern North America, has led to introgression of coyote mitochondrial and nuclear DNA into fragmented, decimated eastern wolf packs. They and other authors have postulated that large populations of eastern and red wolves with intact social/pack structures are less likely to interbreed with coyotes. The controversy over the red wolf's species status was the subject of a comprehensive review of the 2011 and 2012 genetics studies, which concluded that there are three separate species of wolf in North America, the red wolf, eastern wolf and gray wolf.
In a pair of 2012 reports, scientists critical of the May 2011 paper outlined three main points of criticism. First, the 2011 paper relied on mtDNA SNPs derived from boxer and poodle genomes and used these to extrapolate inference about genetic variation within wild canids across the globe. While it is true that many SNPs were examined, 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.
When considered as a full species, three subspecies of red wolf were originally recognized by Goldman; two of these subspecies are extinct. The Florida black wolf (Canis rufus floridanus) (Maine to Florida) has been extinct since 1930 and Gregory's wolf (Canis rufus gregoryi) (south-central United States) was declared functionally extinct in the wild by 1980. The Texas red wolf (Canis rufus rufus), the third 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. The subspecies designations are essentially moot since two are extinct but the genetic evidence for the three subspecies appears to have been unconvincing anyway.
In 2013, an experiment which produced hybrids of coyotes and northwestern gray wolves in captivity using artificial insemination contributed more information to the controversy surrounding the eastern wolf's taxonomy. The purpose of this project was to determine whether 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. While this research is still ongoing, the debate over the eastern wolf's taxonomy remains unsettled. Moreover, in 2014, the research 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 USF&WS to commission a peer review of it, known as NCAES (2014). This peer review concluded unanimously that the Chambers review "is not accepted as consensus scientific opinion or best available science..."
Fossil and historic record
Paleontological evidence has suggested an origin of the red wolf line 1–2 Ma, branching from a wolf-coyote ancestor, which itself appeared about 4.9 Ma. Between 150,000–300,000 years ago, the North American branch evolved into the red wolf, eastern wolf and the coyote. Another wolf-like branch migrated to Eurasia and evolved into the gray wolf, which later migrated to North America. It is thought that its original distribution included much of eastern North America, where red wolves were found from Maine south to 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; it's possible that it could have extended as far as extreme eastern Canada.
Physical description and behavior
The red wolf's appearance is typical of the genus Canis, and is generally intermediate in size between the coyote and gray wolf, though some specimens may overlap in size with small gray wolves. Adults measure 136–160 cm in length, and weigh 40-90 lbs. Its pelage is typically more reddish and sparsely furred than the coyote's and gray wolf's, though melanistic individuals do occur. Its fur is generally tawny to grayish in color, with light markings around the lips and eyes. Like the eastern wolf, the red wolf has been compared by some authors to the greyhound in general form, owing to its relatively long and slender limbs. The ears are also proportionately larger than the coyote's and gray wolf's. The skull is typically narrow, with a long and slender rostrum, a small braincase and a well developed sagittal crest. Its cerebellum is unlike that of 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 basal members of its genus.
The red wolf is more sociable than the coyote, but less so than the gray wolf. It mates in January–February, with an average of 6-7 pups being born in March, April and May. It is monogamous, with both parents participating the rearing of young. Denning sites include hollow tree trunks, along stream banks and the abandoned earths of other animals. By the age of six weeks, the pups distance themselves from the den, and reach full size at the age of one year, becoming sexually mature two years later.
Prior to its extinction in the wild, the red wolf's diet consisted of nutria, rabbits and rodents. In contrast, the red wolves from the restored population rely on white-tailed deer, raccoon, nutria and rabbits. It should be noted, however, that white-tailed deer were largely absent from the last wild refuge of red wolves on the Gulf Coast between 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.
Range and habitat
The originally-recognized red wolf range extended throughout the Southeast, from the Atlantic and Gulf Coasts, north to the Ohio River Valley and central Pennsylvania, and west to Central Texas and southeastern Missouri. 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. Since 1987, red wolves have been released into northeastern North Carolina where they roam 1.7 million acres. These lands span five counties (Dare, Hyde, Tyrrell, Washington and Beaufort) and include three national wildlife refuges, a U.S. Air Force bombing range, and private land. The red wolf recovery program is unique for a large carnivore reintroduction in that more than half of the land used for reintroduction lies on private property. Approximately 680,000 acres (2,800 km2) are federal and state lands, and 1,002,000 acres (4,050 km2) are private lands. Beginning in 1991, red wolves were also released into the Great Smoky Mountains National Park in eastern Tennessee. However, due to exposure to environmental disease (parvovirus), parasites and competition (with coyotes as well as intraspecific aggression), the red wolf was unable to successfully establish a wild population in the park. Low prey density was also a problem, forcing the wolves to leave the park boundaries in pursuit of food in lower elevations. In 1998, the FWS removed the remaining red wolves in the Great Smoky Mountains National Park, relocating them to Alligator River National Wildlife Refuge in eastern North Carolina. Other red wolves have been released on the coastal islands in Florida, Mississippi, and South Carolina as part of the captive breeding management plan. St. Vincent Island in 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.
History and conservation
Discovery and persecution
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. Its reddish color was seen as symbolizing war, as well as all things sacred and beloved. 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.
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. John James Audubon was the first to describe the red wolf in detail, giving it the trinomial name of Canis lupus rufus in 1851. He described it as being more fox-like than the gray wolf, but retaining the same "sneaking, cowardly, yet ferocious disposition". Audubon also recounted that red wolves occasionally fed on the battlefield corpses of fallen soldiers during the Mexican–American War. The red wolf's full specific rank was first given by Edward Goldman in 1937, who examined a number of skulls and noted that the red wolf's skull and dentition differed from those of gray wolves, and closely approached those of coyotes. He wrote that by the time of writing, the species may have become extinct in the lower Sonoran zone, where it was heavily persecuted by ranchmen. Stanley P. Young noted in 1940 that the red wolf was still common in eastern Texas, where more than 800 had been caught in 1939, and further cast doubt over the prospect of fully exterminating them, due to their habit of living concealed in thickets. However, by 1962, red wolf sightings became increasingly rare, and few tracks or scats were found.
Captive breeding and reintroduction
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. 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 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. 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. 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. After that, a larger project was executed in 1987 to reintroduce a permanent population of red wolves back to the wild in the Alligator River National Wildlife Refuge (ARNWR) on the eastern coast of North Carolina. Also in 1987, Bulls Island became the first island breeding site. Pups were raised on the island and relocated to North Carolina until 2005.
In September 1987, four male-female pairs of red wolves were released in 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).
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.
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.
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.
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Names and Taxonomy
Comments: Based on recent genetic studies, Wilson et al. (2000) concluded that the eastern timber wolf (Canis lupus lycaon) and the red wolf (Canis rufus) are sister taxa and are best considered to be conspecific. Additionally, Wilson et al. found that these two taxa form a North American lineage with the coyote (Canis latrans) that is distinct from that of the gray wolf (Canis lupus), which is Eurasian in origin. Wilson et al. (2000, 2003) proposed that the eastern timber wolf (Canis lycaon) be recognized as a species distinct from the gray wolf (C. lupus). In contrast, Nowak (2002) presented an analysis of cranial morphology of recent and Pleistocene Canis and concluded that Canis rufus is a valid species and that lycaon may be a hybrid between Canis rufus and western Canis lupus.
In a recent checklist of North American mammals, Baker et al. (2003) accepted C. lycaon and C. lupus as separate species as proposed by Wilson et al. (2000, 2003). Without explanation they retained the red wolf (Canis rufus) as a third North American wolf species. Wozencraft (in Wilson and Reeder 2005) recognized one Northern Hemisphere wolf species (Canis lupus) and listed rufus and lycaon as subspecies, noting that rufus appears to be a hybrid. In view of the unstable taxonomy of North American wolves and pending further information, this database retains the traditional arrangement of two wolf species in North America (C. lupus and C. rufus), with lycaon treated as a subspecies of C. lupus.
Genetic data (see Wayne and Jenks 1991, Wayne 1992, Wayne et al. 1998, Reich et al. 1999) indicate that existing populations of what have been called red wolves have no unique genetic characteristics and most likely are a product of hybridization between Canis lupus and C. latrans. Generally, however, the red wolf has been accepted as a valid species since the early 1970s (Paradiso and Nowak 1971). See Nowak (1992, 2002) for a response advocating that the red wolf is not a hybrid. Phillips and Henry (1992) reviewed various characters and concluded that Canis rufus is a valid species or at the very least a subspecies of Canis lupus. See also Dowling et al. (1992) for additional discussion of the taxonomic status of the red wolf.
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