Canis lupus rufus

History

Red wolves were once distributed throughout the southeastern and south-central United States from the Atlantic Ocean to central Texas, southeastern Oklahoma and southwestern Illinois in the west, and in the north from the Ohio River Valley, northern Pennsylvania, southern New York, and extreme southern Ontario in Canada south to the Gulf of Mexico. The red wolf was nearly driven to extinction by the mid-1900s due to aggressive predator-control programs, habitat destruction, and extensive hybridization with coyotes. By the late 1960s, it occurred in small numbers in the Gulf Coast of western Louisiana and eastern Texas.

Fourteen of these survivors were selected to be the founders of a captive-bred population, which was established in the Point Defiance Zoo and Aquarium between 1974 and 1980. After a successful experimental relocation to Bulls Island off the coast of South Carolina in 1978, the red wolf was declared extinct in the wild in 1980 so that restoration efforts could proceed. In 1987, the captive animals were released into the Alligator River National Wildlife Refuge (ARNWR) on the Albemarle Peninsula in North Carolina, with a second unsuccessful release taking place two years later in the Great Smoky Mountains National Park. Of 63 red wolves released from 1987 to 1994, the population rose to as many as 100–120 individuals in 2012, but due to the lack of regulation enforcement by the US Fish and Wildlife Service, the population has declined to 40 individuals in 2018, about 14 in 2019 and 8 as of October 2021. No wild litters were born between 2019 and 2020.

Under pressure from conservation groups, the US Fish and Wildlife Service resumed reintroductions in 2021 and increased protection. In 2022, the first wild litter was born since 2018. As of 2023, there are between 15 and 17 wild red wolves in ARNWR.

Description and behavior

The red wolf's appearance is typical of the genus Canis, and is generally intermediate in size between the coyote and gray wolf, though some specimens may overlap in size with small gray wolves. A study of Canis morphometrics conducted in eastern North Carolina reported that red wolves are morphometrically distinct from coyotes and hybrids. Adults measure 136–165 cm (53.5–65 in) in length, comprising a tail of about 37 cm (14.6 in). Their weight ranges from 20 to 39 kg (44–85 lbs) with males averaging 29 kg (64 lbs) and females 25 kg (55 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, pig, raccoon, rice rats, muskrats, nutria, rabbits and carrion. 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 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. 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.

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.

Extirpation in the wild

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

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.

Captive breeding and reintroduction

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

Measurements, vocalization analyses, and skull X-rays were used to distinguish red wolves from coyotes and red wolf × coyote hybrids. Of the 400 canids captured, only 43 were believed to be red wolves and sent to the breeding facility. The first litters were produced in captivity in May 1977. Some of the pups were determined to be hybrids, and they and their parents were removed from the program. Of the original 43 animals, only 17 were considered pure red wolves and since three were unable to breed, 14 became the breeding stock for the captive-breeding program. These 14 were so closely related that they had the genetic effect of being only eight individuals.

In 1996, the red wolf was listed by the International Union for Conservation of Nature as a critically endangered species.

20th century releases

1976 release in Cape Romain National Wildlife Refuge

In December 1976, two wolves were released onto Cape Romain National Wildlife Refuge's Bulls Island in South Carolina with the intent of testing and honing reintroduction methods. They were not released with the intent of beginning a permanent population on the island. The first experimental translocation lasted for 11 days, during which a mated pair of red wolves was monitored day and night with remote telemetry. A second experimental translocation was tried in 1978 with a different mated pair, and they were allowed to remain on the island for close to nine months. After that, a larger project was executed in 1987 to reintroduce a permanent population of red wolves back to the wild in the Alligator River National Wildlife Refuge (ARNWR) on the eastern coast of North Carolina. Also in 1987, Bulls Island became the first island breeding site. Pups were raised on the island and relocated to North Carolina until 2005.

1986 release in Alligator River National Wildlife Refuge

In September 1987, four male-female pairs of red wolves were released in the 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).

1989 release on Horn Island, Mississippi

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.

1991 release in the Great Smoky Mountains

21st century status

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 five 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.

Existing population

In April and May 2023, two captive male red wolves were paired with two wild female wolves in acclimation pens and were later released into the wild. At the same time, the wild breeding pair that produced a litter of pups the previous year gave birth to a second litter of 5 pups, 2 males and 3 females. A male wolf pup from a captive litter was fostered into the pack, and with this new addition, the family of red wolves, which was named the Milltail pack by FWS, has grown to 13 wild individuals. These six new pups have brought the wild population of red wolves up to 23–25 wild individuals.

In May 2023, two families of red wolves were placed in acclimation pens to be released into the wild in the Pocosin Lakes National Wildlife Refuge in Tyrrell County. One family consisted of a breeding pair and three pups, while the other consisted of a breeding pair, a yearling female, and four young pups that were born in the acclamation pen. In early June 2023, the two families of red wolves were released into the wild to roam through PLNWR. With the addition of these two separate packs, the wild population of red wolves had increased to about 35 wild individuals. In addition to the wild population, there are approximately 270 red wolves in zoos and captive breeding programs across the U.S.

Coyote interbreeding

Interbreeding with the coyote has been recognized as a threat affecting the restoration of red wolves. 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 human-caused mortality, are of concern in the restoration of red wolves. Efforts to reduce the threats are presently [when?] being explored.

By 1999, introgression of coyote genes was recognized as the single greatest threat to wild red wolf recovery and an adaptive management plan which included coyote sterilization has been successful, with coyote genes being reduced by 2015 to less than 4% of the wild red wolf population.

Since the 2014 programmatic review, the USFWS ceased implementing the red wolf adaptive management plan that was responsible for preventing red wolf hybridization with coyotes and allowed the release of captive-born red wolves into the wild population. Since then, the wild population has decreased from 100–115 red wolves to less than 30. Despite the controversy over the red wolf's status as a unique taxon, as well as the USFWS' apparent disinterest towards wolf conservation in the wild, the vast majority of public comments (including NC residents) submitted to the USFWS in 2017 over their new wolf management plan were in favor of the original wild conservation plan.

A 2016 genetic study of canid scats found that despite high coyote density inside the Red Wolf Experimental Population Area (RWEPA), hybridization occurs rarely (4% are hybrids).

Contested killing of re-introduced red wolves

High wolf mortality related to anthropogenic causes appeared to be the main factor limiting wolf dispersal westward from the RWEPA.

In 2012, the Southern Environmental Law Center filed a lawsuit against the North Carolina Wildlife Resources Commission for jeopardizing the existence of the wild red wolf population by allowing nighttime hunting of coyotes in the five-county restoration area in eastern North Carolina. A 2014 court-approved settlement agreement was reached that banned nighttime hunting of coyotes and requires permitting and reporting coyote hunting. In response to the settlement, the North Carolina Wildlife Resources Commission adopted a resolution requesting the USFWS to remove all wild red wolves from private lands, terminate recovery efforts, and declare red wolves extinct in the wild. This resolution came in the wake of a 2014 programmatic review of the red wolf conservation program conducted by The Wildlife Management Institute. The Wildlife Management Institute indicated the reintroduction of the red wolf was an incredible achievement. The report indicated that red wolves could be released and survive in the wild, but that illegal killing of red wolves threatens the long-term persistence of the population. The report stated that the USFWS needed to update its red wolf recovery plan, thoroughly evaluate its strategy for preventing coyote hybridization and increase its public outreach.

In 2014, the USFWS issued the first take permit for a red wolf to a private landowner. Since then, the USFWS issued several other take permits to landowners in the five-county restoration area. During June 2015, a landowner shot and killed a female red wolf after being authorized a take permit, causing a public outcry. In response, the Southern Environmental Law Center filed a lawsuit against the USFWS for violating the Endangered Species Act.

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

In June 2018, the USFWS announced a proposal that would limit the wolves' safe range to only Alligator River National Wildlife Refuge, where only about 35 wolves remain, thus allowing hunting on private land. In November 2018, Chief Judge Terrence W. Boyle found that the USFWS had violated its congressional mandate to protect the red wolf, and ruled that USFWS had no power to give landowners the right to shoot them.

Relationship to humans

Since before European colonization of the Americas, the red wolf has featured prominently in Cherokee spiritual beliefs, where it is known as wa'ya (ᏩᏯ), and is said to be the companion of Kana'ti - the hunter and father of the Aniwaya or Wolf Clan. Traditionally, Cherokee people generally avoid killing red wolves, as such an act is believed to bring about the vengeance of the killed animals' pack-mates.

Taxonomy

The taxonomic status of the red wolf is debated. It has been described as either a species with a distinct lineage, a recent hybrid of the gray wolf and the coyote, an ancient hybrid of the gray wolf and the coyote which warrants species status, or a distinct species that has undergone recent hybridization with the coyote.

The naturalists John James Audubon and John Bachman were the first to suggest that the wolves of the southern United States were different from wolves in its other regions. In 1851, they recorded the "Black American Wolf" as C. l. var. ater that existed in Florida, South Carolina, North Carolina, Kentucky, southern Indiana, southern Missouri, Louisiana, and northern Texas. They also recorded the "Red Texan Wolf" as C. l. var. rufus that existed from northern Arkansas, through Texas, and into Mexico. In 1912, the zoologist Gerrit Smith Miller Jr. noted that the designation ater was unavailable and recorded these wolves as C. l. floridanus.

In 1937, the zoologist Edward Alphonso Goldman proposed a new species of wolf Canis rufus. Three subspecies of red wolf were originally recognized by Goldman, with two of these subspecies now being extinct. The Florida black wolf (Canis rufus floridanus) (Maine to Florida) has been extinct since 1908 and the Texas red wolf (Canis rufus rufus) (south-central United States) was declared extinct by 1970. By the 1970s, the Mississippi Valley red wolf (Canis rufus gregoryi) existed only in the coastal prairies and marshes of extreme southeastern Texas and southwestern Louisiana. These were removed from the wild to form a captive breeding program and reintroduced into eastern North Carolina in 1987.

In 1967, the zoologists Barbara Lawrence and William H. Bossert believed that the case for classifying C. rufus as a species was based too heavily on the small red wolves of central Texas, from where it was known that there existed hybridization with the coyote. They said that if an adequate number of specimens had been included from Florida, then the separation of C. rufus from C. lupus would have been unlikely. The taxonomic reference Catalogue of Life classifies the red wolf as a subspecies of Canis lupus. The mammalogist W. Christopher Wozencraft, writing in Mammal Species of the World (2005), regards the red wolf as a hybrid of the gray wolf and the coyote, but due to its uncertain status compromised by recognizing it as a subspecies of the gray wolf Canis lupus rufus.

Taxonomic debate

When European settlers first arrived to North America, the coyote's range was limited to the western half of the continent. They existed in the arid areas and across the open plains, including the prairie regions of the midwestern states. Early explorers found some in Indiana and Wisconsin. From the mid-1800s onward, coyotes began expanding beyond their original range.

The taxonomic debate regarding North American wolves can be summarised as follows:

There are two prevailing evolutionary models for North American Canis:

(i) a two-species model

that identifies grey wolves (C. lupus) and (western) coyotes (Canis latrans) as distinct species that gave rise to various hybrids, including the Great Lakes-boreal wolf (also known as Great Lakes wolf), the eastern coyote (also known as Coywolf / brush wolf / tweed wolf), the red wolf, and the eastern (Algonquin) wolf;

and

(ii) a three-species model

that identifies the grey wolf, western coyote, and eastern wolf (C. lycaon) as distinct species, where Great Lakes-boreal wolves are the product of grey wolf × eastern wolf hybridization, eastern coyotes are the result of eastern wolf × western coyote hybridization, and red wolves are considered historically the same species as the eastern wolf, although their contemporary genetic signature has diverged owing to a bottleneck associated with captive breeding.

The paleontologist Ronald M. Nowak notes that the oldest fossil remains of the red wolf are 10,000 years old and were found in Florida near Melbourne, Brevard County, Withlacoochee River, Citrus County, and Devil's Den Cave, Levy County. He notes that there are only a few, but questionable, fossil remains of the gray wolf found in the southeastern states. He proposes that following the extinction of the dire wolf, the coyote appears to have been displaced from the southeastern US by the red wolf until the last century, when the extirpation of wolves allowed the coyote to expand its range. He also proposes that the ancestor of all North American and Eurasian wolves was C. mosbachensis, which lived in the Middle Pleistocene 700,000–300,000 years ago.