Black-footed cat

The scientific name Felis nigripes was used by the British explorer William John Burchell in 1824 when he described the species based on skins of small, spotted cats that he encountered near Litákun (now known as Dithakong), in South Africa. Felis (Microfelis) nigripes thomasi was proposed as a subspecies by the South African mammalogist Guy C. Shortridge in 1931, who described black-footed cat skins collected in Griqualand West that were darker than those of the nominate subspecies. When the British zoologist Reginald Innes Pocock reviewed cat skins in the collection of the Natural History Museum, London, he corroborated that the black-footed cat is a Felis species. The validity of a subspecies was doubted as no geographical barriers matching the observed differences exist between populations. In 2017, the IUCN Cat Specialist Group revised felid taxonomy and noted that the black-footed cat is most probably a monotypic species. Phylogeny and evolution Phylogenetic analysis of the nuclear DNA from all Felidae species revealed that their evolutionary radiation began in Asia in the Miocene around . Analysis of mitochondrial DNA of all Felidae species indicates that they radiated at around . The black-footed cat is part of an evolutionary lineage that is estimated to have genetically diverged from the common ancestor of all Felis species around , based on analysis of their nuclear DNA. Analysis of their mitochondrial DNA indicates a genetic divergence of Felis species at around . Both models agree on the jungle cat (F. chaus) having been the first Felis species that diverged, followed by the black-footed cat. Fossil remains of the black-footed cat have not been found. It possibly migrated during the Pleistocene into Africa. This migration was possibly facilitated by extended periods of low sea levels between Asia and Africa. The following cladogram shows the phylogenetic relationships of the black-footed cat as derived through analysis of nuclear DNA: ==Characteristics==

The black-footed cat has tawny fur that is entirely covered with black spots. Its head is darker than the rest of the body but paler above the eyes. Its whiskers are white, and its ears bear grizzled dark brown hairs. On the neck and back, some spots are elongated into stripes. The spots form transverse stripes on the shoulders. The forelegs and the hind legs bear irregular stripes. Its tail is confusedly spotted. The underparts of the feet are black or dark brown. The throat rings form black semi-circles that vary in colour from dusky brown to pale rufous and are narrowly edged with rufous. Some individuals have a pure white belly with a tawny tinge where it blends into the tawny colour of the flanks. The ears, eyes and mouth are lined with pale off-white. Two black streaks run from the corners of the eyes across the cheeks. Individuals vary in background colour from sandy and pale ochre to dark ochre. In the northern part of its range, it is lighter than in the southern part, where its spots and bands are more clearly defined. The three rings on the throat are reddish brown to black, with the third ring broken in some individuals. Females measure in head and body length with a long tail. Males are between with a long tail. Its tapering tail is about half the length of the head and body. Females weigh and males . ==Distribution and habitat==

The black-footed cat is endemic to Southern Africa; its distribution is much more restricted than other small cats in this region. Its range extends from South Africa northward into southern Botswana, where it was recorded in the late 1960s. It has also been recorded in Namibia, extreme southern Angola and southern Zimbabwe. It is unlikely to occur in Lesotho and Eswatini. It inhabits open, arid savannas and semi-arid shrubland in the Karoo and the southwestern Kalahari with short grasses, low bush cover, and scattered clumps of low bush and higher grasses. The mean annual precipitation in this region ranges from . In the Drakensberg area, it was recorded at an elevation of . ==Behaviour and ecology==

The black-footed cat is nocturnal and usually solitary, except when females care for dependent kittens. Unlike most other cats, it is a poor climber, as its stocky body and short tail are thought not to be conducive for climbing trees. However, one black-footed cat was observed and photographed resting in the lower branches of a camelthorn tree (Vachellia erioloba). A female roams in an average home range of in a year, and a resident male in an area of . The range of an adult male overlaps the ranges of one to four females. It uses scent marking throughout its range. Other forms of scent marking include rubbing objects, raking with claws, and depositing faeces in visible locations. Its calls are louder than those of other cats of its size, presumably to allow calls to be heard over relatively large distances. When close to each other, however, it uses quieter purrs or gurgles; when threatened, it hisses and growls. It is difficult to survey because of its highly secretive nature; moreover, it tends to move fast without using roads or tracks like other cats. In South Africa, a density of was estimated in Benfontein near Kimberley during 1998 to 1999, that fell to during 2005 to 2014. Farther south, in the Nuwejaarsfontein area, the estimated number of individuals during 2009 to 2014 was . These were probably exceptionally high densities, as both areas feature good weather and management conditions, while the number of individuals in less favourable habitats could be closer to . In a "sit and wait" hunt, it waits for the prey motionlessly in front of a rodent den, sometimes with closed eyes. Its ears keep moving, and it opens the eyes as soon as it hears a sound. It is able to satisfy its daily water requirements through its prey, but drinks water when available. A male pounced on a lamb resting in the grass, but abandoned the hunt after the lamb got up on its feet. It later scavenged the carcass of a recently deceased lamb weighing nearly . It consumed around meat in each of several bouts of eating, starting from the thighs, making its way from the lower back through the flanks to the neck; later it opened up the chest and fed on the inner organs. Insects like harvester termites, grasshoppers and moths constituted about 2% of the prey mass consumed. Reproduction and life cycle In captivity, male black-footed cats become sexually mature at the age of nine months, and females at the age of seven months. The female gives birth to up to two litters per year during the Southern Hemisphere summer between October and March. The litter size is usually one or two kittens, in rare cases also four kittens. They become independent after three to four months and tend to stay within their mother's home range. Captive black-footed cats can live for up to 15 years and three months. Wild black-footed cats are susceptible to transmission of infectious diseases from domestic dogs and cats. == Threats ==

Known threats include methods of indiscriminate predator control, such as bait poisoning and steel-jaw traps, habitat destruction from overgrazing, declining South African springhare populations, intraguild predation, diseases, and unsuitable farming practices. Several black-footed cats have been killed by herding dogs. The majority of protected areas may be too small to adequately conserve viable sub-populations. == Conservation ==

The black-footed cat is listed on CITES Appendix I and is protected throughout most of its range including Botswana and South Africa, where hunting is illegal. In November 2012, this project was extended to Biesiesfontein Farm located in the Victoria West area. Between 1992 and 2018, 65 black-footed cats were radio-collared and followed for extended periods to improve the understanding about their social organisation, sizes and use of their home ranges, hunting behaviour and composition of their diet. Camera traps are used to monitor the behaviour of radio-collared black-footed cats and their interaction with aardwolves (Proteles cristatus). In captivity The Wuppertal Zoo acquired black-footed cats in 1957, and succeeded in breeding them in 1963. In 1993, the European Endangered Species Programme was formed to coordinate which animals are best suited for pairing to maintain genetic diversity and to avoid inbreeding. The International Studbook for the Black-footed Cat was kept in the Wuppertal Zoo in Germany. , detailed records existed for a total of 726 captive cats since 1964; worldwide, 74 individuals were kept in 23 institutions in Germany, United Arab Emirates, US, UK, and South Africa. Several zoos reported breeding successes, including Cleveland Metroparks Zoo, Fresno Chaffee Zoo, Brookfield Zoo, and Philadelphia Zoo. The Audubon Nature Institute's Center for Research of Endangered Species is working on advanced genetics involving cats. In February 2011, a female kept there gave birth to two male kittens – the first black-footed cats to be born as a result of fertilization using frozen and thawed sperm and frozen and thawed embryos. In 2003, the sperm was collected from a male and then frozen. It was used to fertilize an egg in 2005, creating embryos that were thawed and transferred to a surrogate female in December 2010. The female carried the embryos to term and gave birth to two kittens. The same center reported that on 6 February 2012, a female black-footed cat kitten, Crystal, was born to a domestic cat surrogate after interspecies embryo transfer. ==References==