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A conservation assessment of Felis nigripes

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We list Black-footed Cat as Vulnerable C2a(i), as population size is estimated to be fewer than 10,000 mature individuals, where no subpopulation is suspected to be more than 1,000 mature individuals, and there is an inferred continuing decline. We note that this is a precautionary assessment given the low density estimates used in the analysis, the lack of systematic field surveys across the range and the lack of a robust subpopulation definition. This species should be reassessed once such data are available. Key interventions include the establishing of large conservancy areas and sustaining viable Springhare subpopulations in areas where alternative refuge systems are unavailable.
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The Red List of Mammals of South Africa, Lesotho and Swaziland Felis nigripes | 1
Taxonomy
Felis nigripes Burchell 1824
ANIMALIA - CHORDATA - MAMMALIA - CARNIVORA -
FELIDAE - Felis - nigripes
Common names: Black-footed Cat, Small-spotted Cat
(English), Klein Gekolde Kat, Swart-poot Kat,
Miershooptier (Afrikaans), Tsetse (Sesotho), Sebala-
bolokwane, Sebala-molokwane, Sebala, Sebalabala,
Sebala-manokwane, Kêkêtlane (Tswana), Ingwe Yeziduli
(Xhosa)
Taxonomic status: Species
Taxonomic notes: Two subspecies have been described
and these persist in theory (Meester et al. 1986). Felis
n. nigripes, the nominate genotype form described by
Burchell, was first described from near Kuruman in the
Northern Cape. Its range includes present-day
southeastern Namibia, Botswana, Northern Cape, North
West, Gauteng, Limpopo and marginally into
Mpumalanga. Felis n. thomasi Shortridge 1931 is
described from a specimen collected near Grahamstown
Felis nigripes Black-footed Cat
Regional Red List status (2016) Vulnerable C2a(i)*†
National Red List status (2004) Least Concern
Reasons for change Non-genuine:
New information
Global Red List status (2016) Vulnerable C2a(i)
TOPS listing (NEMBA) (2007) Protected
CITES listing (1975) Appendix I
Endemic No
Recommended citation: Wilson B, Sliwa A, Drouilly M. 2016. A conservation assessment of Felis nigripes. In Child MF,
Roxburgh L, Do Linh San E, Raimondo D, Davies-Mostert HT, editors. The Red List of Mammals of South Africa, Swaziland
and Lesotho. South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa.
Black-footed Cat Working Group
in the Eastern Cape. Its range includes the present-day
Eastern Cape, westwards to the southern regions of the
Northern Cape and the Free State.
However, Smithers (1971) doubted the validity of two
subspecies, and his thought is shared by Sliwa (2013)
who suggests that the species may be polymorphic in the
centre of its range but does exhibit some geocline
variations towards the extremes of its range. The northern
race typically tends to be paler with less distinct striping,
whereas the southern race has a more tawny appearance
with bolder patterning. However, since there are no
obvious geographical or ecological barriers between the
ranges which are the leading causes of genetic drift and
speciation (Hoskin et al. 2005), it is likely that the
subspecies status is invalid (Wilson 2016).
Assessment Rationale
Black-footed Cats are endemic to the arid regions of
southern Africa, occurring widely across the western
reaches of the assessment region, and have a relatively
restricted and patchy distribution. The historical paucity of
data has led to inconsistencies and perpetuated
inaccuracies in current literature, which in turn has
affected the accuracy of conservation measures. The
naturally rare, cryptic colouring, small size and secretive
nocturnal nature of this species has contributed
considerably to the lack of information. Black-footed Cats
are known to occur at low densities and it is difficult to
establish population sizes. The stronghold of the species
is suspected to be in the central Karoo region where
highest densities are reached, whereas other regions
(Kalahari, North West Province, northern KwaZulu-Natal
Province, Free State Province, and the Lowveld) are
suspected to have medium or low densities. Using density
estimates of 0.03 km2, 0.02 km2 and 0.01 km2 for high,
medium and low density areas respectively, we converted
kernel densities to isopleths to calculate population size.
Estimated population size ranges from 7,52611,905
individuals, of which 5,2698,334 are considered to be
mature (using a 70% mature population structure), where
no subpopulation is suspected to comprise more than
1,000 mature individuals as the species is patchily
distributed. However, defining subpopulations for this
species needs further work. We also caution that these
estimates will be confounded by uneven sampling across
the range, thus leading to inaccurate density isopleths.
This is a first attempt and should be refined as more data
from across the range become available.
There is a general suspected continuing decline due to
loss of prey base due to bushmeat poaching (especially
Springhare Pedetes capensis), persecution (direct or
incidental), road collisions and predation by domestic
pets. Continuing decline is also inferred from a long-term
study on Benfontein Game Farm, Northern Cape, where
estimated density declined from 0.17 cat / km2 in 1998
1999 to 0.08 cat / km2 in 20052015. Additionally, c. 50%
of radio-collared individuals are lost annually to a
combination of natural mortality and predation from
abundant Black-backed Jackals (Canis mesomelas),
*Watch-list Data Watch-list Threat
Black-footed Cats are very successful hunters
with one vertebrate prey animal caught, on
average, every 50 min (about 1014 rodents or
small birds caught in a night). This represents
about 250300 g of food or 20% of the cat's body
weight, but record intakes can reach 450 g per
night.
Felis nigripes | 2 The Red List of Mammals of South Africa, Lesotho and Swaziland
Figure 1. Distribution records for Black-footed Cat (Felis nigripes) within the assessment region
Caracals (Caracal caracal) or domestic pets. An emerging
threat may be increasing interspecific competition,
including intraguild predation, by overabundant
mesopredators (for example, Black-backed Jackals).
Thus, we list this species as Vulnerable C2a(i), as
population size is estimated to be fewer than 10,000
mature individuals, where no subpopulation is suspected
to be more than 1,000 mature individuals, and there is an
inferred continuing decline. We note that this is a
precautionary assessment given the low density estimates
used in the analysis, the lack of systematic field surveys
across the range and the lack of a robust subpopulation
definition. This species should be reassessed once such
data are available. Key interventions include the
establishing of large conservancy areas and sustaining
viable Springhare subpopulations in areas where
alternative refuge systems are unavailable.
Country Presence Origin
Botswana Extant Native
Lesotho Possibly extant Native
Mozambique Absent -
Namibia Extant Native
South Africa Extant Native
Swaziland Possibly extant Native
Zimbabwe Possibly extant Native
Regional population effects: There is likely to be some
dispersal across regional borders as the range is
continuous across southern Africa. However, the dispersal
capability (c. 20 km; B. Wilson unpubl. data) of this
species is likely to be limited by the species’ small size,
the degree and length of isolation from other
subpopulations caused by human influences and the
presence of competing larger carnivores and large areas
of unsuitable habitat. It is likely that these aspects may be
even greater in the subpopulations outside the region
which will limit these as potential future source
subpopulations and thus limit the rescue effect.
Distribution
Black-footed Cats have the most restricted distribution of
any of the African felid species (Nowell & Jackson 1996).
The species is endemic to the arid grasslands, dwarf
shrub, and savannah of the Karoo and Kalahari in
southern Africa. The majority of the range occurs within
the boundaries of South Africa, thinning out northwards
into Botswana, Namibia and Zimbabwe (Table 1). The
species is found primarily in Namibia and South Africa, but
also Botswana (where there are historical records but only
limited recent ones), marginally in Zimbabwe and likely
marginally in extreme southern Angola (Sliwa 2013).
In South Africa, where the majority of research effort has
taken place, recent records suggest range expansions in
virtually all directions from previous literature (Wilson
2016). It is unlikely that true range extensions have
occurred but is rather an artefact of reporting bias as a
result of recently increased citizen scientist reports
combined with increased research effort (Wilson 2016).
Table 1. Countries of occurrence within southern Africa
The Red List of Mammals of South Africa, Lesotho and Swaziland Felis nigripes | 3
On the eastern side of its range, occurrence records are
confirmed to just west of the Kruger National Park, into
northwestern KwaZulu-Natal, and new data suggest that
the species occurs further south and southwestwards in
the southern Cape and more westwards in the Northern
Cape than previously realised (Wilson 2016). The
occurrence of the species in Swaziland (M. Reilly &
A. Monadjem pers. comm. 2014) and Lesotho
(N.L. Avenant pers. comm. 2014) is considered highly
unlikely and certainly no records exist to date.
Apart from Addo Elephant National Park and Mountain
Zebra National Park in the Eastern Cape Province, and SA
Lombard Nature Reserve in North West Province, there
appears to be no consistent sightings of Black-footed Cats
in formally protected areas, despite considerable research
effort and game drive reporting (Wilson 2016). This is
likely due to unsuitable habitat or the presence of a higher
density of mesopredators which increases competition
and incidences of intraguild predation. They probably
occur in Karoo National Park. However, further field
surveys are needed to determine in which protected areas
subpopulations of the species are resident. Their
presence has now been confirmed through video
evidence in 2016 from Tswalu Kalahari Reserve, Northern
Cape Province (W. Panaino unpubl. data).
The current extent of occurrence in South Africa is roughly
about 930,000 km2 (Wilson 2016). However, we suspect
that area of occupancy is considerably lower, due to
sensitivity to environmental disturbances and threats
(Sliwa 2013).
Population
The Black-footed Cat is naturally rare compared to the
other small cats of southern Africa (Sliwa 2013). Camera
trapping for this species is particularly difficult due its
secretive behaviour and small size. For example, in over
790 records, only one was from a camera trap (B. Wilson
unpubl. data). Black-footed Cats move quickly and do not
habitually use game tracks or roads like other animals.
Since 2006, there has been extensive effort to establish
the historical and current distribution of the species
(Wilson 2016). Prior to 2000, there are only 251
distribution records including fossil specimens. Since
then, more than 545 records have been collected with an
emphasis on locality data. Throughout its entire range
there are only c. 692 verifiable locality records which can
be reliably mapped (Wilson 2016). As such, it is difficult to
determine density, and thus a population estimate, for this
species. High-density estimates come from two study
sites at Benfontein and Nuwejaarsfontein in the Northern
Cape with over 17,000 fixes and 1,600 hours of
observation of radio-collared individuals (Sliwa 2004;
Sliwa et al. 2010): for Benfontein (near Kimberley),
between 1998 and 1999, density (based on radio-collared
individuals) was estimated at 0.17 cat / km2 but only 0.08 cat /
km2 between 2005, whilst at Nuwejaarsfontein, density was
estimated at 0.06 cat / km2 from 20092014 (Sliwa 2004;
Sliwa et al. 2014). However, these two sites may represent
exceptionally high densities due to favourable climate and
human management, and low-quality habitat densities are
probably much lower (Sliwa 2013). Thus, we suspect that
densities of 0.03 cat / km2 represent more realistic
densities across larger scales and represent viable
subpopulations in the long term (B. Wilson and A. Sliwa
unpubl. data).
Using these density estimates, population size was
calculated by converting kernel densities based on
verified records (both historical and recent) to high
(0.03 cat / km2), medium (0.02 cat / km2) and low density
(0.01 cat / km2) isopleths across the entire range of the
species in the assessment region. The population sizes
from the density bands were then summed to produce an
overall population estimate. The best estimate yielded an
estimated 11,905 individuals within the assessment
region, of which 8,334 are mature (assuming a 70%
mature population structure, Table 2). These estimates
were calculated by clipping the range-wide heat map to
South Africa (Figure 2). This was considered a more
biologically realistic method than calculating a separate
heat map for South Africa, as the latter method conserves
linkages and isopleth distributions across borders rather
arbitrarily delineating isopleths for the assessment region.
However, calculating the heat map for the assessment
region alone produces similar population estimates,
ranging from 7,52611,066 individuals (5,2697,747
mature individuals), depending on how the records are
clustered in the isopleths. Additionally, no subpopulation
is suspected to have more than 1,000 mature individuals.
However, the definition of a subpopulation for this species
needs further work depending on dispersal rates and
distances, connectivity and genetic subpopulation
structure, although small home range sizes of the species
minimum estimate for females 7.18.6 km2 (Sliwa 2004;
Kamler et al. 2015) may suggest small subpopulation
sizes. Unpublished data suggest dispersal distance of
20 km (A. Sliwa unpubl. data), and we suspect that
clusters within 50 km of each other and distances of
100 km apart would separate subpopulations. This would
mean a cluster within an area of 1,963 km2 (A = πr2) or
2,500 km2 (A = a2) would yield subpopulation sizes of 334
and 425 individuals (using 0.17 cat / km2) respectively.
These estimates of course suffer from lack of consistent
search effort and thus density zones may be confounded
by field surveys that differ in methodology and single
records may represent substantially higher density areas,
reflecting the lack of field surveys conducted in the region.
As such, population estimates are likely to be
underestimates. This may especially be the case for the
global estimate, where areas outside the assessment
region may be significantly under-sampled, although it is
likely that they exhibit lower densities anyway, as is the
case in Namibia (M. Küsters, pers. comm.). Given that the
Area (km2) / mature population size
High density Medium density Low density Total mature
Assessment region 124,975 / 2,624 179,654 / 2,515 456,293 / 3,194 8,334
Global 124,975 / 2,624 179,654 / 2,515 652,482 / 4,567 9,707
Table 2. Summary of population size estimates for Black-footed Cat (Felis nigripes) both within the assessment region and
globally. Estimates are based on converting existing records into density isopleths and summing the resultant population sizes.
Felis nigripes | 4 The Red List of Mammals of South Africa, Lesotho and Swaziland
assessment region is fairly well-sampled, the population
estimate is likely to be more robust and the analysis
represents a replicable first-step methodology that can be
improved with further field surveys and ecological studies.
It should also be noted that estimates of population sizes
are sensitive to the relative size of the high-density
clusters, thus a more conservative estimate would yield a
lower population size, while a less conservative estimate
would yield a larger population size. For example,
increasing the high-density isopleth to cover 70% of the
records (up from 50%), yields a global population estimate
of 16,853 individuals (11,797 mature). We also note that
the density estimates used for the isopleths are
precautionary and replicated studies in different regions
should be conducted to properly calibrate the density
bands.
There is a higher density in the central part of South Africa,
along a northsouth axis with decreasing frequency of
records either west or east of this region and northwards
in the range (Wilson 2016) (Figure 2).
A decline is suspected over the general range, with only
some central areas indicating stable subpopulations.
However, it is difficult to demonstrate decline in a species
that is so hard to census. Preliminary evidence comes
from the long-term study area Benfontein in the Northern
Cape, where the density in years 19981999 was 0.17
cat / km2 (Sliwa 2004), and over the past 10 years (2005
2015) was only 0.08 cat / km2, which is documented in
capture reports (for example, Sliwa et al. 2014). Local
subpopulations may also be low or even absent in areas
where Black-backed Jackals and Caracals are abundant
due to interspecific competition, including intraguild
predation (but see Kamler et al. 2015), which may be
especially true of juveniles. About 50% of all radio-collared
cats are lost to larger predators (Black-footed Cat Working
Group unpubl. data).
Current population trend: Declining. Inferred from
decline in density from long-term study area.
Continuing decline in mature individuals: Yes, caused
by indirect persecution, predation and disease
transmission.
Number of mature individuals in population: 5,269
8,334 based on extrapolating density across EOO.
Number of mature individuals in largest subpopulation:
Unknown, but it is certainly unlikely that any cluster
numbers more than 1,000 mature individuals.
Number of subpopulations: It is not currently possible to
determine the extent or number of subpopulations.
Severely fragmented: No. Habitat is largely connected
across its range but may be patchy in parts.
Habitats and Ecology
The Black-footed Cat is one of the world’s smallest cats,
with females weighing an average of 1.3 kg and males
larger at 1.93 kg (Sliwa 2013). The conspecific and more
common African Wildcat (Felis silvestris) is considerably
larger (females 3.9 kg; males 5.1 kg) (Sliwa et al. 2010).
Unlike most cat species, these cats are predominantly
ground-dwellers and will not readily take to trees. They
Figure 2. Heat map of distribution records for Black-footed Cat (Felis nigripes); isopleth bands containing the percentage of
distribution records are shown on the left of the figure
The Red List of Mammals of South Africa, Lesotho and Swaziland Felis nigripes | 5
overlapped with those of females by 6067% (Sliwa 2004).
Despite this overlap, all individuals hunted solitarily.
Similarly, annual home range sizes of Black-footed Cats
were 7.1 km2 for an adult female, and 15.6 and 21.3 km2
for two adult males monitored at Benfontein Game Farm,
Nothern Cape Province, from 2006 to 2008 (Kamler et al.
2015). Since range size is dependent on available prey
resources, in more arid regions these home ranges can
be considerably larger. Both sexes spray mark,
particularly during mating season, when spray marks are
deployed in proportion to intensity of use and may play a
role in social spacing (Molteno et al. 1998; Sliwa et al.
2010). Adults travel an average of 8.42 ± 2.09 km per
night a greater distance than the African Wildcat (5.1 ±
3.4 km per night) despite their smaller size (Sliwa et al.
2007).
Ecosystem and cultural services:
A Khoisan legend tells of a Black-footed Cat that
took down a Giraffe (Giraffa camelopardalis) by
piercing its jugular (Sunquist & Sunquist 2002),
which serves only to emphasis the ferocity and
courage contained in such a tiny cat.
This species has the potential to be an iconic
flagship species of southern Africa, particularly in the
Karoo region.
Individuals can consume large amounts of rodents
each year (Figure 4), making this species highly
beneficial to agricultural and livestock farmers where
rodents are considered problematic.
lead a solitary existence except when with kittens or
during brief mating periods. Black-footed Cats are
extremely secretive in nature. They are strictly crepuscular
and nocturnal and are active throughout the night, even
hunting at temperatures of –8˚C (Olbricht & Sliwa 1997).
During the day, the cats make use of dens. The species
prefers hollowed out abandoned termite mounds when
available (especially for the kittens, Figure 3), but will use
dens dug by other animals such as Springhares, Cape
Ground Squirrels (Xerus inauris) and Aardvark
(Orycteropus afer). It is a specialist of open, short grass
areas with an abundance of small rodents and ground-
roosting birds. It inhabits dry, open savannah, grasslands
and Karoo semi-desert with sparse shrub and tree cover
and a mean annual rainfall of between 100 and 500 mm at
altitudes up to 2,000 m asl. It is not found in the driest and
sandiest parts of the Namib and Kalahari Deserts (Sliwa
2013).
Black-footed Cats mate year round but with a distinct
season from late winter (August) over the summer months
up to March. Up to two litters a year may be produced,
with one to four kittens (normally two) born after 6368
day gestation inside a burrow or hollow termitarium
(Olbricht & Sliwa 1997; Skinner & Chimimba 2005). Births
are timed to coincide with rains and food availability.
Kittens are independent after 34 months, but remain
within the range of their mother for extended periods
(Sliwa 2013). Whilst they are reported to live up to 16
years in captive situations, life expectancy in the wild is
about five years (Black-footed Cat Working Group unpubl.
data), although two monitored individuals lived at least
seven years (Sliwa et al. 2014).
As with most small cat species, Black-footed Cats are
obligate carnivores. During a 6-year field study on the
game farm in central South Africa, 1,725 prey items were
observed consumed by 17 free-ranging habituated
individuals (Sliwa 2006). Average prey size was 24.1 g.
Eight males fed on significantly larger prey (27.9 g) than
9 females (20.8 g). Fifty-four prey species were classified
by their average mass into eight different size classes,
three for mammals, three for birds, one for amphibians/
reptiles, and one for invertebrates. Small mammals (5
40 g) constituted the most important prey class (39%) of
total prey biomass followed by larger mammals (> 100 g;
17%) and small birds (< 40 g; 16%). Mammals and birds
pooled comprised 72% and 26% of total prey biomass,
respectively, whereas invertebrates and amphibians/
reptiles combined constituted just 2% of total prey mass
consumed. Heterotherm prey items were unavailable
during winter, when larger birds and mammals (> 100 g)
were mainly consumed. Small rodents like the Large-
eared Mouse (Malacothrix typica), captured 595 times by
both sexes, were particularly important during the
reproductive season for females with kittens. Males
showed less variation between prey size classes
consumed among climatic seasons. This sex-specific
difference in prey size consumption may help to reduce
intra-specific competition (Sliwa 2006). In terms of
interspecific competition, Sliwa et al. (2010) found that
Black-footed Cats captured smaller prey on average than
African Wildcats, although both captured approximately
the same number (1213) of prey animals per night.
Of 17 radio-collared Black-footed Cats (7 males,
10 females) studied at Benfontein Game Farm, Northern
Cape Province, from 1997 to 1998, the home ranges of
adult resident males averaged 16.120.7 km2 while those
of females were 8.610 km2, where male home ranges
Figure 3. A Black-footed Cat protected in a hollowed-out
termitarium
Figure 4. A male Black-footed Cat (Felis nigripes) actively
hunting
Black-footed Cat Working Group
Black-footed Cat Working Group
Felis nigripes | 6 The Red List of Mammals of South Africa, Lesotho and Swaziland
As an enigmatic species, it is a highly prized species
to be seen in the wild by mammal watchers and felid
enthusiasts.
Use and Trade
The Black-footed Cat is accidentally killed in indiscriminate
trapping, hunting (rifle and dogs) and poisoning events
aimed at other damage-causing carnivores. However, as a
protected species these incidents are never reported for
fear of persecution (Wilson 2016). At least one specimen
record reported to B. Wilson was of a Khoisan man in the
Kalahari wearing a skin as part of a loincloth. He indicated
that he obtained the skin from a roadkill from the
neighbouring Botswana area. Similarly, there are
anecdotal reports of skins in households in the central
Karoo (M. Drouilly pers. obs. 2014). Also one flat skin was
obtained from a sheep guard close to the farm
Biesjiesfontein in the Victoria West area in 2012, likely
killed by his hunting dogs (A. Sliwa pers. obs. 2012).
Applications have been made for legal hunting permits in
the Northern Cape and the Eastern Cape provinces, but
these have never been granted. However, several South
African taxidermists have acknowledged having mounted
hunted animals (Wilson 2016), indicating that there is
some interest in the hunting industry for the species.
Recently, there has been an increased interest for this
species in the trophy industry as witnessed by permit
applications and requests made to taxidermists (Wilson
2016).
In the 1970s, there was a demand for the species by
overseas zoos and export permits were granted for cats to
be caught and exported from the Eastern Cape Province.
During further investigations, it was revealed that many of
these cats actually came from the then-Cape Province
(now part of the Northern Cape Province) (Olbricht & Sliwa
1997). More recently, it has become an acknowledged
fact that this species does not thrive in captivity and the
demand for wild-caught individuals has diminished
considerably (Sliwa & Schürer 2006). Zoos now maintain
and exchange animals between institutions. To increase
genetic diversity in these captive populations, in vitro
fertilisation attempts have been made using oocytes of a
captive females and semen collected from a wild-caught
male during annual field research trips conducted by the
Black-footed Cat Working Group (BFCWG) (Herrick et al.
2010), but without generating a viable embryo.
Whilst even predator-proof fencing is not an effective
barrier for Black-footed Cats, the effects of wildlife
ranching are currently unknown and further research is
required to conclusively determine if this land-use practice
differs from other farming practices. Since larger
carnivores are often introduced and managed, the
presence of these is likely to have a negative effect on
Black-footed Cats due to intraguild predation. This is
perhaps the reason why this species does not occur in
formally protected wildlife areas like the Kgalagadi
Transfrontier Park and Kruger National Park but may be
reported in peripheral areas. Effectively, large areas of
intensive wildlife ranching could amplify the island habitat
effect that already prevents movements between
subpopulations, further threatening the species. This
hypothesis should be tested.
Threats
Following 22 years of reach effort by the Black-footed Cat
Working Group, the threats to the species have become
more apparent (Wilson 2016). Additional and previously
unknown threats including, but not limited to, intraguild
predation, diseases, declining Springhare populations and
unsuitable farming practices have now been realised. All
of these suggest that the species is becomingly
increasingly threatened. New distribution data clearly
indicate that the majority of the distribution occurs outside
of formally protected areas (Figure 1). Indeed, the majority
of protected areas are suspected to be too small to
adequately conserve a viable subpopulation. Essentially,
this means the conservation of the species relies on the
cooperation of private landowners and managers,
particularly in large conservancy areas. However, whilst
the geographical distribution of the species may be
greater than previously documented, the actual
occurrence of the species is highly fragmented and
patchy within this area (Wilson 2016), which may have
resulted in the creation of island subpopulations resulting
in limited dispersal opportunities and restricting genetic
exchange between some subpopulations (Wilson 2016).
Habitat loss or changes have previously been considered
the most severe threat to the species. Habitat degradation
that results in the loss of prey base is a serious threat, but
changing agricultural practices may, in some instances,
actually benefit the species by providing artificially high
rodent populations for prey and fewer apex carnivores to
compete with.
Perhaps the most serious long-term threat for Black-
footed Cats is the loss of key resources, such as den sites
and prey, from anthropogenic disturbance or habitat
degradation (for example, from overgrazing). They are
unable to create or maintain their own dens or burrows
and rely on those made by other species. Thus, the
localised removal of a sympatric species, Springhare with
whom they have a crucial inquilistic relationship, can be
detrimental to their continued existence in a region
(B. Wilson unpubl. data; Olbricht & Sliwa 1997; Sliwa
Category Applicable? Rationale Proportion of total
harvest Trend
Subsistence use Yes No reports of bushmeat poaching. However,
skins are probably used on a small scale.
Small Unknown
Commercial use Yes Some illegal trophy hunting. Unknown, but
probably negligible.
Unknown
Harvest from wild population Yes As above Unknown, but
probably negligible.
Unknown
Harvest from ranched population No - - -
Harvest from captive population No - - -
Table 3. Use and trade summary for the Black-footed Cat (Felis nigripes)
The Red List of Mammals of South Africa, Lesotho and Swaziland Felis nigripes | 7
2013). Springhares are often considered a problem or
damage-causing species requiring some control
measures by farmers. Although a rodent, Springhares are
long-lived with a slow reproductive rate and they do not
recover easily from a severe reduction in numbers. In
Botswana, the unregulated subsistence hunting of
Springhares for bushmeat has resulted in the eradication
of the species in some regions (Butynski 1973, 2013), and
with them, in all likelihood the Black-footed Cats (Wilson
2016). Bushmeat hunting may be increasing in both scope
and scale within southern African savannahs (Lindsey et
al. 2013).
The impact of mesopredators may be increasing across
South African rangelands (Avenant & du Plessis 2008). In
general, numbers of Black-footed Cats and other larger
carnivores are negatively related due to intraguild
predation (Sliwa et al. 2009, 2014). Annually, the BFCWG
loses about 50% of all radio-collared cats to larger
predators (BFCWG unpubl. data). The BFCWG believes
that the most vulnerable individuals (kittens and subadults
below 1 kg) are the more likely victims of predation but
since these individuals are difficult to monitor (too small to
fit a radio collar) the actual rate of loss is unknown.
Similarly, during a study over 20062008 on Benfontein
Farm, Northern Cape Province, two Black-footed Cats
were killed by predation; one was killed by Black-backed
Jackals (Kamler et al. 2015; Wilson 2016) and the other by
a Caracal (Wilson 2016). Thus, although Black-footed Cats
can co-exist with Black-backed Jackals by using burrows
during the day (see above for synergistic interaction) and
also taking refuge in them at night during danger (A. Sliwa
pers. obs.), and by partitioning activity and diets (Kamler
et al. 2015), increasing Black-backed Jackal abundance
caused by anthropogenic disturbance and loss of apex
predators is likely to increase Black-footed Cat mortality.
The BFCWG also has evidence of cats killed by traditional
herding dogs and the popularity of Anatolian shepherd
dogs with sheep farmers is potentially a new emerging
threat (Sliwa et al. 2014; Wilson 2016).
Black-footed Cats are also lost through indirect
persecution, such as accidental poisonings (for example
locust spraying, predator control lures/baits) and general
predator persecution throughout most of their range in
South Africa (Nowell & Jackson 1996; Sliwa 2013).
The extent of road mortality on cat subpopulations is not
known. The incident rate recorded in the Endangered
Wildlife Trust’s Road Collision Database is limited to a few
records (W. Collinson unpubl. data). However, 3% out of
790 locality records throughout the species’ entire
distribution area were road collision casualties (Wilson
2016).
Although an inherited trait, and previously thought to be
limited only to inbred captive populations, Black-footed
Cats show a high prevalence for AA-amyloidosis (Olbricht
& Sliwa 1997; Terio et al. 2008). This is a disease
Rank Threat description Evidence in the
scientific literature
Data
quality
Scale of
study Current trend
1 5.1 Hunting & Collecting Terrestrial Animals: bushmeat
poaching and direct persecution of sympatric Pedetes
capensis. Current stress 1.2 Ecosystem Degradation:
causes loss of available den sites.
Butynski 1973, 2013 Indirect Regional Possibly increasing
with rise in bushmeat
poaching in southern
Africa.
2 2.3 Livestock Farming and Ranching: habitat
degradation from overgrazing resulting in loss of prey
base.
Avenant & du
Plessis 2008
Indirect Local Unknown
3 8.2 Problematic Native Species/Diseases: persecution
of large carnivores leads to mesopredator release.
Current stress 1.2 Ecosystem Degradation: removal of
apex predator leads to mesopredator release,
especially Canis mesomelas.
Kamler et al. 2015 Empirical Local Possibly increasing
due to increase in
Black-backed Jackal
numbers.
4 5.1 Hunting & Collecting Terrestrial Animals: incidental
persecution through poisoning, pesticides or traps
intended for other problem animals.
Wilson 2016 Empirical Regional Unknown
5 8.1 Invasive Non-Native/Alien Species/Diseases:
predation by feral dogs and Anatolian shepherd dogs.
Wilson 2016 Empirical Regional Unknown
6 4.1 Roads & Railroads: mortalities from road collisions. Wilson 2016 Empirical Regional Unknown
7 5.1 Hunting & Collecting Terrestrial Animals: illegal
trophy hunting.
Wilson 2016 Empirical Regional Unknown
8 8.2 Problematic Native Species/Diseases: AA-
amyloidosis outbreaks.
Terio et al. 2008 Indirect Local Possibly increasing
with increased contact
with other small
carnivores, domestic
animals and climate
change.
9 11.1 Habitat Shifting & Alternation: climate change may
alter prey densities/distributions, facilitate disease
transmission and increase the frequency of flooding
and droughts.
Wilson 2016 Anecdotal - -
Table 4. Threats to the Black-footed Cat (Felis nigripes) ranked in order of severity with corresponding evidence (based on IUCN
threat categories, with regional context)
Felis nigripes | 8 The Red List of Mammals of South Africa, Lesotho and Swaziland
characterised by fibrillar protein depositions in many
organs as a result of chronic inflammatory processes
usually cumulating in renal failure. About 70% of the
documented deaths of captive cats internationally are as a
result of this disease. However, the presence of amyloid in
free-ranging subpopulations was detected by Terio et al.
(2008) from samples provided by the BFCWG. This
provides additional evidence for a species predilection
and supports the existence of a possible familial type of
amyloidosis in the species. Habitat fragmentation and
subpopulation isolation can only exacerbate this disease
at subpopulation levels, whilst at the same time, the
disease is a major reason why currently global captive
breeding programmes are not self-sustainable (A. Sliwa
pers. comm.). As wild subpopulations become genetically
isolated, reduced genetic variability threatens population
viability by increasing susceptibility to disease and
reducing reproductive fitness. An additional consequence
of habitat fragmentation and population isolation is
increased contact with other carnivores and the
pathogens they carry. Because Black-footed Cats share
their territory, prey base, and infectious disease
susceptibility with many small carnivores, and even
domestic dogs and cats, this provides numerous
opportunities for disease transmission (Lamberski et al.
2009).
Black-footed Cats are also vulnerable to natural disasters
such as flooding of dens and den collapses (Sliwa et al.
2009). The long-term effects of climate change cannot be
overlooked and may lead to changes in range, changes in
timing of breeding events, increases in severe weather
such as flooding and droughts, as well as increased
disease patterns or risks of the spread of pathogens from
parasites.
Fortunately, unlike the African Wildcat, this species does
not hybridise easily with other cat species. The only
confirmed hybrid cases (Black-footed Cats and domestic
cats) were under captive conditions (Leyhausen 1979).
None have been recorded from the wild (A. Sliwa &
B. Wilson pers. obs.).
Current habitat trend: Previously believed to be restricted
mostly to panveld and short grass areas, Black-footed
Cats occupy a wide range of open arid and semi-arid
habitats where they favour any vegetation cover that is low
and not too dense, and they have even been sporadically
recorded on fallow agricultural fields and in extremely
overgrazed areas (Wilson 2016). They are therefore not
restricted by habitats throughout most of their distribution
in southern Africa. However, overgrazing from livestock
farms leads to bush encroachment which reduces the
habitat suitability for their normal prey items. The species
is highly adaptable and reports of individuals utilising
open agricultural areas suggests that the cats may
respond to seasonal fluctuations of rodent populations
associated with sowing and harvesting activities. However,
given the unpredictable nature of these events, and
associated risks due to the proximity to humans and their
domestic carnivores, the benefit to resident cats is
uncertain. Also there is no research as to how these
human-dominated landscapes are used by Black-footed
Cats and whether these specific records are due to the
fact that they are exposed and highly frequented by
human observers (A. Sliwa pers. obs.).
Conservation
Black-footed Cat presence in formally protected areas
remains low to non-existent, particularly in those large
enough to maintain subpopulations; for example,
Kgalagadi Transfrontier Park. It is unclear in how many
protected areas they occur. However, the major Karoo
protected areas Karoo, Mountain Zebra and Addo
Elephant National Parks may each protect a small but
not self-sustaining population. Hunting of this species is
banned in South Africa and Botswana and it is protected
across most of its range excluding Namibia and
Zimbabwe (Nowell & Jackson 1996). Although the Black-
footed Cat has been recorded at least marginally in all of
the South African provinces, the effectiveness of local
protection measures remains in question.
Key interventions for the species include:
1. Judicious management of larger predators and
mesopredators in areas where farming practices have
resulted in unnatural population increases: this is a
holistic approach that includes only controlling true
problem animals, and ensuring that fields are not
overgrazed or overstocked so as to sustain natural
small mammal prey. The effect should be to reduce
unnaturally high mesopredator density. Improved
sheep-farming practices (for example, synchronised
breeding events, kraaling at night, and shepherd
systems, lethal control of only proven problem
mesopredators) should also be trialled.
2. The establishment of large conservancy areas to
create viable Black-footed Cat subpopulations and
facilitate ecological separation between Black-footed
Cats and larger carnivores (Kamler et al. 2015). The
fragmentation of suitable habitats and potential
isolation of subpopulations makes the formation or
maintenance of dispersal corridors important in the
prevention of inbreeding and prevalence of inherited
diseases such as AA-amyloidosis. Despite its small
size, individuals have very large home ranges, and to
conserve subpopulations and create corridors, the
emphasis should be on the establishment of large
conservancy areas with suitable conditions for the
species. This is particularly important in areas where
the prospects of a formally protected area are
unlikely.
3. Similarly, conservancies and private lands with low
apex predator density can be targeted as possible
sites for Black-footed Cat stewardship.
4. Human activities that lead to habitat degradation and
the loss of prey species need to be addressed,
particularly in the Karoo region which is likely to be
the remaining stronghold region for the species. For
example, conservation success may partly depend on
sustaining Springhare numbers, with whom they are
inquilistic, by decreasing persecution and bushmeat
hunting, especially in areas where alternative refuge
systems, such as termite mounds, are unavailable
(Wilson 2016). This should also be achieved by
creating awareness to the presence and needs of the
species among landowners to reduce accidental
persecution, whilst providing information about the
special needs of Black-footed Cats that would enable
them to be actively involved in the protection of the
species.
The Red List of Mammals of South Africa, Lesotho and Swaziland Felis nigripes | 9
5. This may also include generally raising public
awareness of the cryptic species and encouraging
citizen scientists to submit distribution records
through the promotion of citizen science platforms.
This has been done previously with the use of posters
created by the BFCWG requesting the public to report
sightings of the species, the bulk of the data having
been used to establish a more accurate geographical
distribution of the species (Wilson 2016).
6. Other interventions can involve applying stiffer legal
penalties to people involved in deliberate persecution,
illegally keeping or trading, or illegally hunting the
species.
In the long term, the creation of a Biodiversity
Management Plan for the species as an interim or pre-
emptive conservation measure should be investigated
which could be linked to the already existing international
and national ex situ management plans. Currently there is
no consistent and self-sustaining breeding and survival of
cats in South African facilities and some only maintain
their stocks by receiving rescued/confiscated individuals
from the wild. We suggested the establishment of a
national breeding studbook for Black-footed Cats, which
would then allow a better overview on the stocks, which
could then also feed to the international studbook,
maintained by Wuppertal Zoo in Germany since 1991.
Recomm end ations for land m anagers and
practitioners: Currently, there are no conservation plans
for Black-footed Cats and the urgency to create such
plans is undermined by the paucity of data at a national
level. It is anticipated that the numbers will decrease over
the next 20 year period, but during this period, available
data should give a clear picture as to the areas of range
that are most affected, and the best possible interventions
needed.
Currently, at an international level, due to its already
low numbers, it is one of the species listed in the
American Association of Zoos and Aquariums
Species Survival Plan program developed in 1981.
However, the as-yet undetermined subspecies
status (F. n. nigripes and F. n. thomasi) may
undermine and place in jeopardy the international
holdings and ex situ management strategies for the
species as a whole. A number of zoos have captive
individuals and an international stud book is
maintained for the species. Success in captive
conditions is fraught by failure of individuals to thrive
due to their highly specialised needs and the effects
of inbreeding and/or AA-amyloidosis. Thus, ex situ
conservation efforts are not recommended until such
time as expertise exists in South Africa.
Translocation of individuals without the monitoring of
the released cats to determine survival rates is not
advocated or advised as displaced individuals may
suffer a high mortality rate. Systematic monitoring in
areas identified as under-sampled should be
established to improve population estimates and
scan for stewardship sites.
The presence/absence of the species in formally
protected areas must be ascertained.
Research priorities: The Black-footed Cat has been
extensively studied for more than 20 years near the
Kimberley area, along the Free StateNorthern Cape
border. The BFCWG focuses on the ecology, reproductive
biology, geographical range, habitat preferences, health
and conservation of the species. This provided information
about the diet (Sliwa 2006; Sliwa et al. 2010), home range
size and social organisation (Sliwa 2004), ecological
relationships between the species and other sympatric
carnivores (Kamler et al. 2015). More recently, the BFCWG
extended this study to include farms south of De Aar,
Northern Cape Province, in a different habitat type with
different farming practices in place. However, there is little
information for elsewhere within its range and the
following are considered research priorities.
Fine-scale distributional studies across the
distributional range. This will hopefully lead to more
accurate subpopulation estimates. Long-term
monitoring of subpopulation trends are also needed
throughout the species’ geographic range,
particularly in ecologically distinct areas and under
varying farming practices. Kitten mortality and
survival rates also need to be quantified.
Rank Intervention description
Evidence in
the scientific
literature
Data
quality
Scale of
evidence
Demonstrated
impact
Current
conservation
projects
1 2.1 Site/Area Management: form conservancies. - Anecdotal - - -
2 2.1 Site/Area Management: reduce stocking rates and
overgrazing to sustain natural prey base.
- Anecdotal - - -
3 5.3 Private Sector Standards and Codes: identify
private lands with low apex predator density as
stewardship sites.
- Anecdotal - - -
4 3.1.1 Harvest Management: decrease offtake of
Springhare to sustain den site creation for Black-
footed Cats.
- Anecdotal - - -
5 4.3 Awareness & Communications: inform private
landowners of Black-footed Cat occurrence and
habitat requirements.
- Anecdotal - - Black-footed
Cat Working
Group
6 5.4 Compliance & Enforcement: enforce penalties on
those illegally harvesting, persecuting, trading or
breeding Black-footed Cats.
- Anecdotal - - -
Table 5. Conservation interventions for the Black-footed Cat (Felis nigripes) ranked in order of effectiveness with corresponding
evidence (based on IUCN action categories, with regional context)
Felis nigripes | 10 The Red List of Mammals of South Africa, Lesotho and Swaziland
Similarly, studies on changes in density across a
spectrum of habitat quality are needed to refine
conservation plans and subpopulation estimates.
Fine-scale determination on what habitat
characteristics and prey populations are required for
female Black-footed Cats to successfully raise their
kittens is also needed.
The impacts and extent of persecution (both direct
and indirect) and the efficacy of education and
awareness programmes targeted at landowners.
Genetic investigation into the subspecies status is
needed as the results may have conservation
implications.
Investigation into causes and extent of AA-
amyloidosis in wild populations.
The effects of Black-backed Jackals and Caracals,
and possibly other apex carnivores, need to be
quantified.
The degree of inquilism and dependence on
Springhare burrow systems for refuge, and thus
long-term survival of cats following of the removal of
Springhare subpopulations, needs to be determined.
Studies into the dispersal abilities and survival of
subadult Black-footed Cats in different habitats.
Effects of electric fences on local movements and
dispersal patterns must be understood to assess the
effects of long-term geographical isolation on
subpopulation structure. This will enable us to define
and delineate subpopulations more accurately.
Monitoring of the success of re-introduced
individuals into new areas with/without other
individuals present.
The range and impact of transmittable diseases from
sympatric carnivores on Black-footed Cats.
The numbers of Black-footed Cats being removed
for the trophy industry.
Other research projects include:
WAZA Conservation Project 06016 Cologne Zoo
(Germany); McGregor Museum (Kimberley, South
Africa); San Diego Zoo Safari Park (USA); Wuppertal
Zoo (Germany), http://www.waza.org/en/site/
conservation/waza-conservation-projects/overview/
blackfooted-cat-research.
Distribution and Conservation Status of the Black-
Footed Cat Felis nigripes in Namibia http://
www.rufford.org/projects/martina_kusters. Martina
Küsters; bfcat@mail.com or
kusters.m@hotmail.com.
Encouraged citizen actions:
Report sightings (live or dead) to BFCWG
bfc.sightings@gmail.com or via http://black-footed-
cat.wild-cat.org/contact.
Create conservancies, particularly in the Karoo
region.
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Data quality (max) Estimated
Data quality (min) Suspected
Uncertainty resolution Maximum / minimum values
Risk tolerance Precautionary
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Black-footed Cat (Felis nigripes) assessment
Data Sources and Quality
The Red List of Mammals of South Africa, Lesotho and Swaziland Felis nigripes | 11
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Assessors and Reviewers
Beryl Wilson1†, Alex Sliwa2†, Marine Drouilly3
1McGregor Museum, 2Cologne Zoo, 3University of Cape Town
Black-footed Cat Working Group
Contributors
Lizanne Roxburgh1, Jaco Vivier2, R. John Power3,
Matthew F. Child1, Michael Hoffmann4, Kristin Nowell
1Endangered Wildlife Trust, 2The Vale Farm, 3North West Provincial
Government, 4International Union for the Conservation of Nature
IUCN SSC Cat Specialist Group
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be found in Mammal Red List 2016: Introduction and
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Wilson B. 2016. Geographical distribution and status of the black-
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University of Technology, Tshwane, South Africa.
... Based on a literature review of diet and habitat overlap, C. caracal shares some portion of its geographic range in Africa with 67 other mammalian carnivores and was vulnerable from nine of those species when stealing prey (Caro and Stoner 2003) which indicates the relative importance of exploitative competition and interspecific killings for C. caracal in Africa. C. caracal kills and feeds on small carnivores such as African wildcat (Felis lybica), Cape gray mongoose, small-spotted genet (Genetta genetta), bat-eared fox (Otocyon megalotis- Drouilly et al. 2018b), and vulnerable species such as the black-footed cat (Felis nigripes- Wilson et al. 2016). Adult C. caracal can be killed by leopard and spotted hyena (Crocuta crocuta -Skinner 1979), whereas kittens will be predated by these carnivores and jackals if found (Furstenburg 2010;Bothma 2012). ...
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