Mammalian Species 47(925):78–83
Felis nigripes (Carnivora: Felidae)
Aurélie Renard, Maxime Lavoie, Justin A. Pitt, and Serge Larivière
Départment de Biologie, Université du Québec à Rimouski, 300 Allée des Ursulines, Rimouski, Québec G5L 3A1, Canada; [email protected] (AR); [email protected] (ML); [email protected] (SL)
Department of Biological Sciences, University of Alberta, P.O. Box 221, Pincher Creek, Alberta T0K 1W0, Canada; pitt@ualberta.
ca (JAP)
Present address of SL: Cree Hunters and Trappers Income Security Board, 2700 Blvd Laurier, Champlain #1110, Québec G1V 4K5,
Canada
Abstract: Felis nigripes Burchell, 1824, commonly called the black-footed cat, is the smallest felid in Africa. F. nigripes is a pale
tawny cat with dark markings and prominent horizontal stripes on the limbs. It is distributed in the central part of South Africa and
mainly occurs in open country often near brush cover. F. nigripes is listed on Appendix I of the Convention for the International Trade
of Endangered Species of Fauna and Flora and is listed as “Vulnerable” by the International Union for the Conservation of Nature
and Natural Resources.
Key words: black-footed cat, carnivore, felid, South Africa
© 2015 by American Society of Mammalogists
Synonymy completed 1 July 2010
DOI:10.1093/mspecies/sev008
www.mammalogy.org
Felis Nigripes Burchell, 1824
Black-footed Cat
F. n. nigripes Burchell, 1824:592. See above.
F. n. thomasi Shortridge, 1931:119. See above.
Nomenclatural Notes. Felis is Latin for cat. The specific
name nigripes is from nigri meaning black or dark, and pes
meaning foot (Borror 1960). Other vernacular names include
small-spotted cat, anthill tiger, chat à pieds noirs or chat à pattes
noires (French), gato de pies negro or gato patinegro (Spanish),
Felis nigripes Burchell, 1824:592. Type locality “the town
of Litákun (Letárkoon)…27° 6” 44” [S] …24° 39” 27”
[E];” author implied the country of the Bachapins [= near
Kuruman, northern Cape Province, South Africa].
Microfelis nigripes: Roberts, 1926:250. Name combination.
Felis (Microfelis) nigripes thomasi Shortridge, 1931:119. Type
locality “S South Africa, Eastern Cape Province.”
Microfelis nigripes nigripes: Shortridge, 1934:95. Name
combination.
Microfelis nigripes thomasi: Shortridge, 1934:95. Name
combination.
Context and Content. Order Carnivora, family Felidae,
subfamily Felinae. The genus contains 7 species: F. bieti,
F. catus, F. chaus, F. manul, F. margarita, F. nigripes, and
F. silvestris (Wozencraft 2005); however, a more recent
source (Sunquist and Sunquist 2009) lists only 5 species:
referring to F. catus as the domestic form of F. silvestris, and
placing F. manul in the genus Otocolobus. Two subspecies
have been recognized (Shortridge 1931; Ellerman et al. 1953;
Wozencraft 2005; Sunquist and Sunquist 2009), although
other sources indicate these findings are spurious and that F.
nigripes might be monotypic (Smithers 1971; Olbricht and
Sliwa 1997):
Fig. 1.—Adult Felis nigripes from Victoria West, South Africa. Image
1043 of the American Society of Mammalogists Images Library. Used
with permission of photographer, J. Visser.
78
47(925)—Felis nigripes
mammalian species79
Schwartzfusskatze (German), klein gekolde kat, swart poot
kat, miershooptier (Afrikaans), koirus (Nama), tutchu (Naron
Bushman), sebala, lototsi (Setswana), and ingwe yeziduli
(Xhosa—Rautenbach 1971; Nowell and Jackson 1996).
DIAGNOSIS
Felis nigripes (Fig. 1) is the smallest cat in Africa (Dorst
and Dandelot 1970). It can be differentiated from the sympatric
F. silvestris (wildcat) by the lack of defined spots, the overall grayish appearance, and the larger size (shoulder height >
250 mm) of the latter (Pringle 1977). F. nigripes can be differentiated from other African cats by its small size (< 2.5 kg)
and distinct markings (Guggisberg 1975). The auditory bullae
of this species are proportionally larger (> 33%) than most in
the other species of Felidae (Pocock 1951; Huang et al. 2002;
Peters et al. 2009).
GENERAL CHARACTERS
Felis nigripes is a small, pale tawny cat with dark markings and
prominent horizontal stripes on the limbs. Markings occur in rings
on the throat, blotches on chest and belly, and 2 streaks across the
cheeks, as well as transverse bars across the limbs. Pelage is dark
tawny on the dorsum with pale to white on the venter. Hairs are
25–30 mm in length. Margins of eyes, ears, and mouth are white.
Nose is small, and ears are pointed (Guggisberg 1975; Kingdon
1997). Soles of feet are black. Tail is short, less than one-half the
length of the head and body. Tail is pointy and black-tipped, with
several dark rings, sometimes incomplete at the base (Guggisberg
1975). Mean measurements (mm) with ranges of 5 males and 3
females from Botswana (Smithers 1971; Skinner and Smithers
1990) were, respectively: total length, 579 (540–631), 513 (495–
530); length of tail, 177 (164–198), 153 (126–170); length of hind
foot, 99 (94–104), 92 (89–94); length of ear, 54 (51–57), 47 (45–
50). Mean masses (kg; range) of the same individuals were: 1.6
(1.5–1.7) for the males and 1.1 (1.0–1.4) for the females (Smithers
1971; Skinner and Smithers 1990). Measurements (mm; range) of
3 males from Cape Province (Stuart 1981) were: length of head
and body, 447 (414–490); length of tail, 143 (80–200); length of
hind foot, 102 (95–109); length of ear, 53 (45–58). Mean mass
(kg) for these 3 males was 2.0 (range 1.7–2.4—Stuart 1981) and
for 1 captive female mass was 1.4 (Deem et al. 1998). Mass (kg;
range) for 8 males and 10 females, from South Africa were: 1.9
(1.6–2.5) and 1.3 (1.1–1.7), respectively (Sliwa 2004). Males,
contrary to females, seem to gain weight during the cold season
(May–August—Sliwa 2004). Vocal track length for 1 adult from
South Africa was 13.0 cm (Peters et al. 2009).
Skull (Fig. 2) is small, high, and rounded compared to other
felids but large compared to its own body (Sicuro 2011). The top
of the skull is flat from the eye orbits to the middle of the braincase,
where it slopes abruptly. Rostrum is short and broad. Postorbital
constriction is wide, about 40% of the total length of the skull.
Fig. 2.—Dorsal, ventral, and lateral views of skull and lateral view of
mandible of Felis nigripes (male, Transvaal Museum 25998). Greatest
length of skull is 89.95 mm.
Postorbital bars are incomplete (Skinner and Smithers 1990).
Mean condylobasal length (mm) of skull from 21 males and 11
females was 74.8 and 70.4, respectively (Peters et al. 2009).
DISTRIBUTION
Felis nigripes occurs in Namibia, Botswana, and the central
part of South Africa. It also occurs in southern Zimbabwe (Fig. 3).
Due to its secretive nature, there appear to be discrepancies in
its exact distribution (von Ritcher 1972). It is most abundant in
arid regions of the Kalahari, Karoon, and Western Cape Province
80
mammalian species47(925)—Felis nigripes
Fig. 3.—Geographic distribution of Felis nigripes in Africa. Map modified from Kingdon (1997), Lynch (1975, 1985), Sliwa (2008), Stuart
(1982), and Sunquist and Sunquist (2002, 2009). There are no clear ecological or geographical barriers between the 2 subspecies, F. n. nigripes
and F. n. thomasi.
(Stuart 1982; Kingdon 1997) and is rare north of 20–21° latitude
(Shortridge 1934; Smithers 1971). Recent sightings above 24°
have extended the known distribution of this species northward
(Power 2000). Highest densities are thought to reside in areas
of high gerbil (Muridae, Gerbillinae) densities (Visser 1977).
As with all carnivores, fossil records indicate that cats emerged
from the miacids (Stuart and Wilson 1988); however, no fossils
of F. nigripes are known.
FORM AND FUNCTION
Dental formula is i 3/3, c 1/1, p 3/2, m 1/1, total 30
(Skinner and Smithers 1990). Canines are very sharp, upper
2nd premolars are tiny, as are molars. The pattern of the teeth
follows the characteristic felid arrangement for killing bites,
with long, sharp, blade-like canines, and the sectorial action
of the other teeth in slicing of food into portions small enough
to swallow (Skinner and Smithers 1990). Mean cranial volume
for 6 skulls was 25 ml ± 5.4 (SD), and a mean mass of 2.0 kg
(Sheppey and Bernard 1984). Females have 3 pairs of nipples.
Felis nigripes can subsist without drinking water (Kingdon
1997).
ONTOGENY AND REPRODUCTION
Neonate mass is 60.0–87.9 g, and young Felis nigripes gain
8 g daily (Hemmer 1976). Eyes open at 6–9 days, at 15 days young
are able to walk swiftly, and at 20 days young begin to climb
(Armstrong 1975; Schürer 1978a, 1978b). Young start taking solid
food after 34 days. The study of a wild female revealed the young
stayed with the mother 15–17 weeks and the male played no role
in parental care (Molteno et al. 1998). Young are left unattended
while the female is hunting (Molteno et al. 1998; Sliwa 1999).
Permanent upper canines erupt after 150–158 days, females reach
sexual maturity at 7 months (Olbricht and Sliwa 1997), and a male
showed 1st sign of sexual maturity at 7.5 months (Armstrong
1975). The oldest individual in captivity died at the age of 15 years
and 3 months (Sliwa and Schürer 2001).
Breeding usually occurs in November–December (Kingdon
1997) but births have been recorded until May (Sliwa et al.
2010). Gestations last 63–68 days, and litter size may reach 4,
but usually is 1–2 (Guggisberg 1975; Hemmer 1976; Olbricht
and Sliwa 1997). One hundred and ninety-one litters born in
captivity averaged 1.8 young (Crandall 1964; Leyhausen and
Tonkin 1966; Lucas 1970; Armstrong 1975, 1977; Schürer
1978a, 1978b; Olbricht and Sliwa 1997). In captivity, females
bred continuously with 1 female producing 5 litters in 512 days
(Schürer 1978a). The birth sex ratio for 367 captive individuals was 205:162 (Sliwa and Schürer 2001). Typical estrus lasts
24–48 h with a maximum length of 36 h (Leyhausen and Tonkin
1966; Herrick et al. 2010). The estrus cycle averages 11–12 days
(Herrick et al. 2010). Females might mate while still caring for
young from a previous litter (Molteno et al. 1998). One maternity den in South Africa was located in a hollowed out termite
mound (Sliwa 1998). Head, midpiece, and tail measurements
(μm3) of spermatozoids were, respectively: length 4.310, 9.785,
36.374 and volume 11.268, 2.337, 2.543 (Anderson et al. 2005).
ECOLOGY
Population density for Felis nigripes has been estimated at
0.17 adults/km2 in South Africa (Sliwa 2004). It occurs in open
country near brush cover (Smithers 1971) but the results of a
study on 20 southern Kalahari rangeland sites indicated that F.
nigripes was not impacted negatively by encroaching shrub cover
(Blaum et al. 2007). Home range sizes of males are larger than
those of females. In South Africa, mean home range size (100%
Minimum Convex Polygon) for 5 males and 7 females averaged
20.7 and 10 km2, respectively, and remained stable among years
(Sliwa 2004). Intrasexual overlap was low for males (12.9%),
but substantially higher for females (40%—Sliwa 2004). Adult
male residents overlap with up to 4 females (Sliwa 2004).
Diet consists mainly of gerbils (Gerbillinae) and bush-rats
(Aethomys—Kingdon 1997), but F. nigripes is opportunistic
and will consume anything it can overpower or scavenge (Sliwa
1994b). Birds, spiders, insects, and reptiles may be consumed
occasionally (Kingdon 1997). Stomachs from 5 F. nigripes collected in Botswana contained members of Muridae (number
of occurrence 2/5), arachnids (2/5), birds (1/5), reptiles (1/5),
and coleopteran (1/5). Muridae were gerbils (Tatera), Southern
African pouched mouse (Saccostomus campestris), and Paeba
hairy-footed gerbil (Gerbillurus paeba), the bird was a doublebanded courser, Rhinoptilus africanus, and the reptile a spiny
47(925)—Felis nigripes
mammalian species81
agama, Agama hispida (Smithers 1971). One male collected in
Transvaal had consumed 321 cc of mammals, including 155 cc
of the South African ground squirrel (Xerus inauris) and 50 cc
of gerbil, 3 cc of birds (feathers only), and 1 cc of green grass
(Bothma 1965). Two F. nigripes (1 male, 1 female) were followed for a total of 622 h over late summer, autumn, and winter
in Northern Cape Province, South Africa (Sliwa 1994b). Of 550
prey items consumed, 407 were insects (340 harvester termites,
Hodotermes mossambicus; 22 antlions, Planipennia; 20 locusts
and grasshoppers, Saltatoria; and 25 large moths, Lepidoptera),
37 were birds (14 clapper larks, Mirafra apiata; 5 spike-heeled
larks, Chersomanes albofasciata; 5 kurrichane buttonquail,
Turnix sylvatica; 3 desert cisticola, Cisticola aridula; 2 black
bustard, Eupodotis afer; 1 Richards pipit, Anthus novaeseelandiae; 1 sabota lark, Mirafra sabota; 1 fawn-colored lark; 1
capped wheateater, Oenanthe pileata; and 4 unidentified birds),
and 131 were mammals (34 large-eared African desert mouse,
Malacothrix typica; 23 Paeba hairy-footed gerbil; 17 bushveld
gerbil, Tatera [currently Gerbilliscus] leucogaster; 16 reddishgray musk shrew, Crocidura cyanea; 9 gray African climbing
mouse, Dendromus melanotis; 2 Cape hare, Lepus capensis; 1
Southern African pouched mouse, and 1 springbok, Antidorcas
marsupialis, that was scavenged—Sliwa 1994a, 1994b).
Mammals comprised 72% of consumption by weight, and birds
and insects comprised 26% and 1% of weight, respectively
(Sliwa 1994a, 1994b, 2006).
In one study, prey size averaged 24 g; however, males fed on
larger (27.9 g) prey items than females (20.8 g—Sliwa 2006). In
summer, females captured a higher proportion of small (5–40 g)
and medium (40–100 g) prey although in winter they relied more
on large birds and mammals (> 100 g). The diet of males did
not change between these 2 seasons (Sliwa 2006). In captivity,
F. nigripes has been known to feed on grasses (Leyhausen and
Tonkin 1966); however, this was rarely observed in wild cats
(Bothma 1965; Sliwa 1994a, 1994b).
Felis nigripes can be affected by heartworms (e.g.,
Diroflilaria immitis), protozoans like Babesia, and glomerulonephritis or amyloidosis (Deem et al. 1998; Taugner et al. 2003;
Bosman et al. 2007; Terio et al. 2008). Newborns, young individuals, and possibly adults may be predated by the black-backed
jackal (Canis mesomelas) and large nocturnal raptors like spotted eagle owls (Bubo africanus), Cape eagle owls (B. capensis),
and giant eagle owls (B. lacteus). African marsh owls (Asio
capensis) may benefit from F. nigripes by catching birds flushed
during hunting (Olbricht and Sliwa 1997).
HUSBANDRY
Felis nigripes has been successfully reared and kept in
captivity (Mangold 1989), with a steady increase from 4 individuals in 1974 to 108 in 1990 (Olbricht and Sliwa 1997). This
species has been maintained in North America, Europe, and
Africa (Dmoch 1988). In captivity, 2 major problems have been
identified: (1) high mortality in the very young and adults which
is largely due to high rates of kidney disease and (2) a sex ratio
that is heavily biased toward males (Olbricht and Sliwa 1997).
A limited tolerance to cool and humid air has also been noted
(Olbricht and Schürer 1994). F. nigripes in captivity must have
a diet that includes feathers as they contain an essential amino
acid taurine along with a higher proportion of fiber in relation to amount of protein intake (Olbricht and Schürer 1994).
F. nigripes can live up to 15 years in captivity (Olbricht and
Sliwa 1997). It is possible to use odor for environmental enrichment of captive individuals to encourage an increase of activity and species-typical behavior. Catnip and odor of prey were
more effective than nutmeg (Wells and Egli 2004).
For research purposes, F. nigripes has been anesthetized
with a mixture of ketamine hydrochloride and acetylpromazine
or ketamine and diazepam (Deem 1998), fitted with radio collars
weighing 50 g, and followed at a distance of 10–30 m (Molteno
et al. 1998; Sliwa 2004). Cloned embryos of F. nigripes can be
transferred into domestic cat (Felis silvestris catus) recipients
(Gómez et al. 2006).
BEHAVIOR
The frequency spectrum of the mew call of Felis nigripes
is restricted to the range below 2 kHz (Peters et al. 2009). Scent
marking is common and it appears it may be used to advertise
reproductive status (Molteno et al. 1998). A sharp rise in scenting (5.22–35.90 sprays/h) events occurred 45 days before mating and conception, and when no young were present. Based on
the observation of 1 female, urine-spraying was the predominant
form of marking behavior (Molteno et al. 1998).
Felis nigripes is completely nocturnal, secretive, and solitary (Smithers 1971). Hunting occurs through the night in
all weather conditions from −8 to 28°C (Olbricht and Sliwa
1997). The species hunts on the ground in open, sandy country but close to thickets, termitaries, and clumps of grass, so
as to have access to cover for safety (Smithers 1971). Prey are
captured by stalking (slow stalking winding movements at 0.5–
0.8 km/h), flushing from cover (swift bounds at 1–2 km/h), or
sit-and-wait (waiting by a rodent den for up to 1 h—Olbricht
and Sliwa 1997). F. nigripes can leap up to 1.5 m to catch its
prey (Sliwa 1998; Hunter 1999). Caching behavior has been
observed; however, caches are rarely completely covered (Sliwa
1994a, 1994b, 1998). Dismembering of prey items has been
described in both wild and captive cats, mainly the removal of
the stomach and large intestines from rodents (Leyhausen 1982;
Sliwa 1994b). Observations from 4 individuals in South Africa
determined that F. nigripes makes a vertebrate kill about every
50 min (Sliwa 1994b).
Felis nigripes is not a good climber; however, recent research
has documented climbing behavior on several occasions (Sliwa
1998). F. nigripes is a powerful digger; digging behavior probably arises from the habit of digging holes to use as shelter. During
the day, F. nigripes hides in burrows dug by the Cape porcupine—Hystrix africaeaustralis, the aardvark—Orycteropus afer,
82
mammalian species47(925)—Felis nigripes
or the South African cape hare—Pedetes capensis, and holes in
termitaries (Guggisberg 1975).
Captive behavior of F. nigripes may be influenced by olfactory stimulation (Wells and Egli 2004). Experimental odors initially increased active behaviors (e.g., grooming, exploration),
but responses decreased over a 5-day observation period, suggesting habituation.
GENETICS
The mitochondrial DNA, Y chromosome, cytochrome b, and
12SrRNA sequences of Felis nigripes suggest that it diverged earlier than those of other members of the domestic cat group (Felis),
which diverged about 7–10 million years ago (Masuda et al. 1995;
Johnson et al. 1996; Pecon-Slattery et al. 2004). The karyotype
of F. nigripes is indistinguishable from that of F. silvestris catus.
It has a diploid number (2n) and a fundamental number (FN) of
38 and 72 chromosomes, respectively. It has 16 pairs of metacentric or submetacentric and 1 or 2 pairs of acrocentric chromosomes (Wurster and Benirschke 1968; Wurster-Hill 1973; Pathak
et al. 1998). The Y and X chromosomes are, respectively, a small
subacrocentric and medium-sized submetacentric (Wurster and
Benirschke 1968). A phylogenic reconstruction of the Felidae
using 1,504 (10 karyological, 58 morphological, 1,436 molecular) characters revealed 55 (5 morphological, 50 molecular) autapomorphies for F. nigripes. F. nigripes was placed in the domestic
cat group along with the rusty-spotted cat (Prionailurus rubiginosus), the Pallas’ cat (F. manul), the jungle cat (F. chaus), the
wild cat (F. silvestris), the sand cat (F. margarita), and F. s. catus
(Mattern and McLennan 2000). Hybridizing occurs between this
species and F. s. catus (Tonkin 1972).
CONSERVATION
Felis nigripes is listed on Appendix I of the Convention for
the International Trade of Endangered Species of Wild Fauna and
Flora (2010) and is listed as “Vulnerable” by the International
Union for the Conservation of Nature and Natural Resources
(2010). The remaining population size is estimated at < 10,000
mature individuals with a declining trend due to loss of prey
base, nontarget poisoning, and persecution (Olbricht and Sliwa
1997; Sliwa 2008). Habitat degradation caused by expansion of
human settlement, overgrazing, and agriculture is a major threat
for this species (Nowell and Jackson 1996; Sliwa 2008). Assisted
reproduction might be useful technology to maintain genetically
viable, captive populations of F. nigripes (Swanson 2006).
ACKNOWLEDGMENTS
We are grateful to T. Kearney and M. Burger of the
Transvaal Museum for the skull photos. D. Dyck (University
of Saskatchewan) helped with map and photo editing. The
authors would like to thank A. Sliwa for his valuable comments
and additional references. SL is thankful to NSERC (Natural
Sciences and Engineering Research Council of Canada) for
support through a discovery grant and to the Cree Hunters and
Trappers Income Security Board for logistic support.
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Associate Editor and Editor was Meredith J. Hamilton.