The red wolf (Canis rufus) is a type of dog that lives in the southeastern United States. It is slightly larger than a coyote (Canis latrans) but smaller than a gray wolf (Canis lupus).

For almost 100 years, scientists have debated whether the red wolf is a separate species or a subspecies of the gray wolf (Canis lupus rufus). Some also believe it is a mix of wolf and coyote genes, sometimes called a "coywolf." Because of this debate, the red wolf is sometimes not included on lists of endangered species, even though there are very few of them left. Under the Endangered Species Act of 1973, the U.S. Fish and Wildlife Service recognizes the red wolf as an endangered species and gives it legal protection. Since 1996, the IUCN has listed the red wolf as Critically Endangered. However, it is not included in the CITES Appendices, which are international lists of endangered species.

History

Red wolves once lived across the southeastern and south-central United States, from the Atlantic Ocean to central Texas, southeastern Oklahoma, and southwestern Illinois in the west, and from the Ohio River Valley, northern Pennsylvania, southern New York, and extreme southern Ontario in Canada in the north, down to the Gulf of Mexico. By the mid-1900s, red wolves were nearly gone because of programs to kill predators, loss of their homes, and mixing with coyotes. By the late 1960s, only a few remained along the Gulf Coast of western Louisiana and eastern Texas.

Fourteen of these wolves were chosen to start a group raised in captivity. This group was created at the Point Defiance Zoo and Aquarium between 1974 and 1980. In 1978, an attempt to move them to Bulls Island near South Carolina was successful. Because of this, red wolves were declared extinct in the wild in 1980 to allow recovery efforts to begin. In 1987, captive wolves were released into the Alligator River National Wildlife Refuge (ARNWR) in North Carolina. Another release in the Great Smoky Mountains National Park two years later did not succeed. Between 1987 and 1994, 63 wolves were released. By 2012, their population reached as many as 100–120 individuals. However, because rules were not followed properly by the US Fish and Wildlife Service, the population dropped to 40 in 2018, 14 in 2019, and 8 by October 2021. No baby wolves were born in the wild between 2019 and 2020.

Because of pressure from groups working to protect animals, the US Fish and Wildlife Service began moving wolves back into the wild again in 2021 and improved their protection. In 2022, the first wild-born babies were born since 2018. As of 2023, there are between 15 and 17 wild red wolves in ARNWR.

Description and behavior

The red wolf looks similar to other members of the Canis genus and is usually between the size of a coyote and a gray wolf. Some red wolves may be about the same size as small gray wolves. A study in eastern North Carolina found that red wolves have body measurements that are different from coyotes and hybrids. Adult red wolves are 136–165 cm (53.5–65 in) long, with a tail about 37 cm (14.6 in). They weigh between 20 and 39 kg (44–85 lbs), with males averaging 29 kg (64 lbs) and females 25 kg (55 lbs). Their fur is usually reddish and less thick than that of coyotes and gray wolves, though some have dark fur. Their coat is generally tawny or grayish, with lighter markings near the mouth and eyes. The red wolf’s body shape is often compared to a greyhound because of its long, slender legs. Its ears are larger than those of coyotes and gray wolves. The skull is narrow, with a long, slender snout, a small braincase, and a strong ridge on the top of the skull. Its cerebellum is more similar to that of foxes and other canid species, suggesting the red wolf is one of the more primitive members of its genus.

Red wolves are more social than coyotes but less social than gray wolves. They mate in January–February, and 6–7 pups are born in March, April, or May. They are monogamous, and both parents help raise the young. Dens are often in hollow tree trunks, near streams, or in abandoned burrows. By six weeks old, pups leave the den, and they reach full size at one year old. They become sexually mature at two years old.

Long-term studies of red wolves with known family backgrounds showed that inbreeding between close relatives was rare. A likely reason is that young wolves leave their birth pack to find new areas. Many young wolves live alone or in small groups with unrelated individuals. New breeding pairs usually form when two unrelated wolves join to claim a new territory. Inbreeding is avoided because it leads to offspring with weaker health, caused by harmful genes becoming dominant.

Before becoming extinct in the wild, red wolves mainly ate rabbits, rodents, and nutria (a non-native species). In restored populations, they eat white-tailed deer, pigs, raccoons, rice rats, muskrats, nutria, rabbits, and carrion. White-tailed deer were not common in the last wild area of red wolves on the Gulf Coast (where the last wild population was captured for breeding), which explains the difference in their diets. Early explorers, like William Hilton, who traveled along the Cape Fear River in North Carolina in 1644, also reported that wolves ate deer.

In Florida, red wolves may be preyed upon by some stages of invasive snakes, including Burmese pythons, reticulated pythons, Southern African rock pythons, Central African rock pythons, boa constrictors, yellow anacondas, Bolivian anacondas, dark-spotted anacondas, and green anacondas.

Range and habitat

The original range of the red wolf covered much of 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 by Ronald Nowak, using ancient bones, historical records, and archaeological findings, showed that red wolves also lived in areas south of the Saint Lawrence River in Canada, along the eastern seaboard, and west to Missouri and mid-Illinois, ending in southern Central Texas.

Red wolves likely lived in many different types of habitats in the past. The last natural population used coastal prairies, marshes, swamps, and fields where rice and cotton were grown. However, these environments may not have been their preferred homes. Evidence suggests that red wolves were most common in the large river bottomlands and swamps of the southeastern United States. Wolves reintroduced to northeastern North Carolina have used a variety of habitats, including farmland and mixed areas of forest and wetlands with pine trees and shrubs. This shows that red wolves can live in many different places if there is enough prey and little human interference.

In 1940, biologist Stanley P. Young noted that red wolves were still common in eastern Texas, where over 800 were caught in 1939 due to attacks on livestock. He believed they were hard to eliminate because they hid in thick vegetation. By 1962, a study of wild Canis skulls in Arkansas, Louisiana, Oklahoma, and Texas showed that red wolves existed only in small groups due to mixing with coyotes. This was likely because red wolves could not adapt to changes in their environment caused by human activities and the arrival of coyotes from the west, or because coyotes caused red wolves to disappear through hybridization.

Since 1987, red wolves have been released into northeastern North Carolina, where they live across 1.7 million acres. This area includes five counties (Dare, Hyde, Tyrrell, Washington, and Beaufort) and features three national wildlife refuges, a U.S. Air Force bombing range, and private land. The red wolf recovery program is special because more than half of the land used for reintroduction is on private property. About 680,000 acres (2,800 square kilometers) are federal or state lands, and 1,002,000 acres (4,050 square kilometers) are private lands.

In 1991, red wolves were also released into the Great Smoky Mountains National Park in eastern Tennessee. However, they struggled to survive due to disease (parvovirus), parasites, competition with coyotes, and fighting among wolves. Low prey numbers also forced them to leave the park to find food in lower areas. In 1998, the U.S. Fish and Wildlife Service moved the remaining wolves from the park to Alligator River National Wildlife Refuge in eastern North Carolina. Other wolves have been released on coastal islands in Florida, Mississippi, and South Carolina as part of breeding programs. St. Vincent Island in Florida is the only active island site for raising red wolves today.

Captive breeding and reintroduction

After the Endangered Species Act of 1973 was passed, the U.S. Fish and Wildlife Service began official efforts to save the red wolf from extinction. A captive-breeding program was started at the Point Defiance Zoological Gardens in Tacoma, Washington. Between 1973 and 1980, 400 animals were captured from southwestern Louisiana and southeastern Texas by the USFWS.

Scientists used measurements, vocalization analyses, and skull X-rays to identify red wolves and distinguish them from coyotes and hybrids. Of the 400 animals captured, only 43 were believed to be pure red wolves and sent to the breeding facility. The first litters were born in captivity in May 1977. Some of the pups were hybrids, and they and their parents were removed from the program. Of the original 43 animals, only 17 were considered pure red wolves. Three of these could not breed, leaving 14 to be the breeding stock. These 14 were so closely related that they had the genetic makeup of only eight individuals.

In 1996, the International Union for Conservation of Nature listed the red wolf as critically endangered. Over 30 facilities participate in the red wolf Species Survival Plan, which oversees breeding and reintroducing more than 150 wolves.

In 2007, the USFWS estimated that 300 red wolves remained worldwide, with 207 in captivity. By late 2020, only about 7 wild red wolves had radio collars, and about a dozen were uncollared. No wild pups had been born since 2018. This decline was linked to wolves being shot or poisoned by landowners and reduced conservation efforts by the USFWS.

A 2019 study by the Center for Biological Diversity found that over 20,000 square miles of public land across five areas had suitable habitat for red wolves to be reintroduced. These areas were chosen based on prey availability, distance from coyotes and human development, and connectivity between sites. The sites include the Apalachicola and Osceola National Forests, the Okefenokee National Wildlife Refuge, and nearby protected lands; national parks and forests in the Appalachian Mountains such as Monongahela, George Washington & Jefferson, Cherokee, Pisgah, Nantahala, Chattahoochee, and Talladega National Forests, Shenandoah National Park, and 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 mostly resembling coyotes were found on Galveston Island, Texas. These animals had red wolf genes from a past population. Scientists proposed selectively breeding these coyotes with the captive red wolf population. A 2019 study found that about 55% of sampled canids in southwestern Louisiana had red wolf ancestry, with one individual showing 78–100% red wolf genes. This suggests that some red wolf genes may still exist in the wild but are not present in the captive population.

From 2015 to 2019, no red wolves were released into the wild. In March 2020, the FWS released a new breeding pair of red wolves, including a young male from St. Vincent Island, Florida, into the Alligator River National Wildlife Refuge. The pair did not produce a litter. In March 2021, two male red wolves from Florida were paired with two female wild red wolves from eastern North Carolina and released. One male was killed by a car shortly after release. On April 30 and May 1, 2021, four adult red wolves were released, and four pups were fostered by a wild female. In total, nearly 30 wild red wolves lived in the area, including those not wearing radio collars.

A 2020 study reported that camera traps in northeast Texas captured images of a large canid with wolf-like features. Later, hair samples and tracks confirmed the presence of red wolves.

By fall 2021, six red wolves had been killed, including four adults released in spring 2021. Three were killed in vehicle collisions, two died from unknown causes, and one was shot by a landowner. These losses reduced the wild population to about 20 individuals. In winter 2021–2022, the USFWS released nine captive adult red wolves into the wild. A family of five was placed in Pocosin Lakes National Wildlife Refuge, while two breeding pairs were released into Alligator River National Wildlife Refuge. This increased the wild population to less than 30 individuals.

On April 22, 2022, a breeding pair of adult red wolves produced six pups, four females and two males. These pups were the first born in the wild since 2018. As of 2023, between 15 and 17 wild red wolves live in Alligator River National Wildlife Refuge.

In April and May 2023, two captive male red wolves were paired with two wild females in acclimation pens and later released. At the same time, a wild breeding pair gave birth to a second litter of five pups, two males and three females. A male pup from a captive litter was fostered into the pack, increasing the Milltail pack to 13 individuals. This brought the wild population to 23–25 individuals.

In May 2023, two families of red wolves were placed in acclimation pens for release into Pocosin Lakes National Wildlife Refuge. One family included a breeding pair and three pups, while the other included a breeding pair, a yearling female, and four young pups born in the acclimation pen. In early June 2023, these families were released into the wild, increasing the population to about 35 individuals. In addition to the wild population, approximately 270 red wolves live in zoos and captive breeding programs across the U.S.

Interbreeding with coyotes has been identified as a major threat to red wolf recovery. Adaptive management efforts have reduced coyote genes in the wild to less than 4% by 2015. Other threats, such as habitat loss, disease, and human-caused deaths, remain concerns. Efforts to address these threats are ongoing.

By 1999, the mixing of coyote genes with red wolves was recognized as the greatest threat to wild red wolf recovery. An adaptive management plan, which included sterilizing coyotes, was successful in reducing coyote genes in the wild by 2015. Since the 2014 programmatic review, the USFWS has not implemented

Relationship to humans

Before Europeans arrived in the Americas, the red wolf held an important place in Cherokee spiritual beliefs. The Cherokee call the red wolf "wa'ya" (ᏩᏯ) and believe it is the companion of Kana'ti, a hunter and the father of the Aniwaya, also known as the Wolf Clan. Traditionally, the Cherokee usually do not kill red wolves because they believe killing one may cause revenge from the wolf's pack members.

Gallery

• Displaying different colors
• During the winter season
• Contrasting colors
• A red wolf participating in a breeding program. There are fewer than 100 red wolves living in the wild.
• Wearing a radio collar
• A red wolf that was captured in Oklahoma in 1944

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 .

In 2021, the American Society of Mammalogists considered the red wolf as its own species ( Canis rufus ).

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:

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.

C. mosbachensis was a wolf that once lived across Eurasia before going extinct. It was smaller than most North American wolf populations and smaller than C. rufus , and has been described as being similar in size to the small Indian wolf , Canis lupus pallipes . He further proposes that C. mosbachensis invaded North America where it became isolated by the later glaciation and there gave rise to C. rufus . In Eurasia, C. mosbachensis evolved into C. lupus , which later invaded North America.

The paleontologist and expert on the genus Canis ' natural history, Xiaoming Wang , looked at red wolf fossil material but could not state if it was, or was not, a separate species. He said that Nowak had put together more morphometric data on red wolves than anybody else, but Nowak's statistical analysis of the data revealed a red wolf that is difficult to deal with. Wang proposes that studies of ancient DNA taken from fossils might help settle the debate. In 2009, Tedford, Wang and Taylor reclassified the purported red wolf fossils as Canis armbrusteri and Canis edwardii .

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 no