Red Wolf

## Description

The red wolf (Canis rufus ) is a canine native to the southeastern United States. Its size is intermediate between the coyote (Canis latrans ) and gray wolf (Canis lupus ).

The red wolf's taxonomic classification as being a separate species, a subspecies of the gray wolf *Canis lupus rufus*, or a coywolf (a genetic admixture of wolf and coyote) has been contentious for nearly a century. Because of this, it is sometimes excluded from endangered species lists, despite its critically low numbers. Under the Endangered Species Act of 1973, the U.S. Fish and Wildlife Service currently recognizes the red wolf as an endangered species and grants protected status. Since 1996, the IUCN has listed the red wolf as a Critically Endangered species, however it is not listed in the CITES Appendices of endangered species.

## Appearance

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. 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. 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".

In 1905, the mammalogist Vernon Bailey referred to the "Texan Red Wolf" with the first use of the name Canis rufus. 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. 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.

It is difficult to distinguish the red wolf from a red wolf × coyote hybrid. 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. 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.

In 1971, a study of the skulls of C. rufus, C. lupus and C. latrans indicated that C. rufus was distinguishable by being in size and shape midway between the gray wolf and the coyote. A re-examination of museum canine skulls collected from central Texas between 1915 and 1918 showed variations spanning from C. rufus through to C. latrans. The study proposes that by 1930 due to human habitat modification, the red wolf had disappeared from this region and had been replaced by a hybrid swarm. By 1969, this hybrid swarm was moving eastwards into eastern Texas and Louisiana.

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. In 1970, the wolf mammalogist L. David Mech proposed that the red wolf was a hybrid of the gray wolf and coyote. However, a 1971 study compared the cerebellum within the brain of six *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. In 2014, a three-dimensional morphometrics study of *Canis* species accepted only six red wolf specimens for analysis from those on offer, due to the impact of hybridization on the others.

## Distribution

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.

idence 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.

### Reintroduced habitat

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 took away 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.

## Behavior

The red wolf's appearance is typical of the genus Canis, and is generally intermediate in size between the coyote and gray wolf, though some specimens may overlap in size with small gray wolves. A study of Canis morphometrics conducted in eastern North Carolina reported that red wolves are morphometrically distinct from coyotes and hybrids. Adults measure 136–160 cm (53.5–63 in) in length, and weigh 23–39 kg (50-85 lbs). Its pelage is typically more reddish and sparsely furred than the coyote's and gray wolf's, though melanistic individuals do occur. Its fur is generally tawny to grayish in color, with light markings around the lips and eyes. 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.

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 in the rearing of young. Denning sites include hollow tree trunks, along stream banks and the abandoned earths of other animals. By the age of six weeks, the pups distance themselves from the den, and reach full size at the age of one year, becoming sexually mature two years later.

Using long-term data on red wolf individuals of known pedigree, it was found that inbreeding among first-degree relatives was rare. A likely mechanism for avoidance of inbreeding is independent dispersal trajectories from the natal pack. Many of the young wolves spend time alone or in small non-breeding packs composed of unrelated individuals. The union of two unrelated individuals in a new home range is the predominant pattern of breeding pair formation. Inbreeding is avoided because it results in progeny with reduced fitness (inbreeding depression) that is predominantly caused by the homozygous expression of recessive deleterious alleles.

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

## Population

### Population number

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

In 2007, the USFWS estimated that 300 red wolves remained in the world, with 207 of those in captivity. By late 2020, the number of wild individuals had shrunk to only about 7 radio-collared and a dozen uncollared individuals, with no wild pups born since 2018. This decline has been linked to shooting and poisoning of wolves by landowners, and suspended conservation efforts by the USFWS.

A 2019 analysis by the Center for Biological Diversity of available habitat throughout the red wolf's former range found that over 20,000 square miles of public land across 5 sites had viable habitat for red wolves to be reintroduced to in the future. These sites were chosen based on prey levels, isolation from coyotes and human development, and connectivity with other sites. These sites include: the Apalachicola and Osceola National Forests along with the Okefenokee National Wildlife Refuge and nearby protected lands; numerous national parks and national forests in the Appalachian Mountains including the Monongahela, George Washington & Jefferson, Cherokee, Pisgah, Nantahala, Chattahoochee, and Talladega National Forests along with Shenandoah National Park and the lower elevations of Great Smoky Mountains National Park; Croatoan National Forest and Hofmann Forest on the North Carolina coast, and the Ozark, Ouatchita, and Mark Twain National Forests in the central United States.

In late 2018, two canids that are largely coyote were found on Galveston Island, Texas with red wolf alleles (gene expressions) left from a ghost population of red wolves. Since these alleles are from a different population from the red wolves in the North Carolina captive breeding program, there has been a proposal to selectively cross-breed the Galveston Island coyotes into the captive red wolf population. Another study published around the same time analyzing canid scat and hair samples in southwestern Louisiana found genetic evidence of red wolf ancestry in about 55% of sampled canids, with one such individual having between 78 and 100% red wolf ancestry, suggesting the possibility of more red wolf genes in the wild that may not be present in the captive population.

From 2015 to 2019, there were no red wolves released into the wild. But in March 2020, the FWS released a new breeding pair of red wolves, including a young male red wolf from St. Vincent Island, Florida into the Alligator River National Wildlife Refuge. The pair were unsuccessful at producing a litter of pups in the wild. On March 1, 2021, two male red wolves from Florida were paired with two female wild red wolves from eastern North Carolina and released into the wild. One of the male wolves was killed by a car shortly after being released into the wild. On April 30 and May 1, four adult red wolves were released into the wild and four red wolf pups were fostered by a wild female red wolf. In addition to the eight released wolves, the total number of red wolves living in the wild amount to nearly thirty wild individuals, including a dozen other wolves not wearing radio collars.

A study published in 2020 reported camera traps recorded "the presence of a large canid possessing wolf-like characters" in northeast Texas and later hair samples and tracks from the area indicated the presence of red wolves.

By fall of 2021, a total of six red wolves had been killed, including the four adults that had been released in the spring. Three of the released adults had been killed in vehicle collisions, two had died from unknown cases, and the fourth released adult had been shot by a landowner who feared the wolf was attempting to get his chickens. These losses dropped the number of wolves in the wild down to about 20 wild individuals. In the winter of 2021-2022, the Fish and Wildlife Services selected nine captive adult red wolves to be released into the wild. A family of five red wolves were released into the Pocosin Lakes National Wildlife Refuge, while two new breeding pairs of adult wolves were released into the Alligator River National Wildlife Refuge. The release of these new wolves brought the number of wild red wolves in eastern North Carolina up to less than 30 wild individuals.

On April 22, 2022, one of the breeding pairs of adult red wolves produced a litter of six wolf pups, four females and two males. This new litter of red wolf pups became the first litter born in the wild since 2018.