Nandinia binotata, Gray, 1843

Don E. Wilson & Russell A. Mittermeier, 2009, Nandiniidae, Handbook of the Mammals of the World – Volume 1 Carnivores, Barcelona: Lynx Edicions, pp. 50-53 : 52-53

African Palm Civet

Nandinia binotata

French: Nandinie / German: Pardelroller / Spanish: Nandinia

Taxonomy. Viverra binotata Gray, 1830 ,

Ashanti region, Ghana.

Four subspecies recognized, but taxonomic boundaries at the subspecific level poorly defined.

Subspecies and Distribution.

N. b. binotata Gray, 1830 — from Gambia to DR Congo, including Bioko I.

N. b. arborea Heller, 1913 — Kenya, S Sudan, N Tanzania, and Uganda.

N. b. gerrardi Thomas, 1893 — Malawi, Mozambique, S&E Tanzania (presumably also Zanzibar I), NE Zambia, and E Zimbabwe.

N. b. intensa Cabrera & Ruxton, 1926 — Angola, S DR Congo, and NW Zambia.

Descriptive notes. Head-body 37-62.5 cm, tail 34-76.2 cm; weight 1.2- 3 kg. Two pairs of mammae. Adult males may be slightly larger than females. Head is broad; fairly pointed muzzle; nose pad brown to dark. Ears are short, broad at the base and rounded; there is a single brown-black spot behind ear pinnae. Iris is brownish-orange. Pelage is short, woolly, and dense. Coat color is relatively uniform across the body, and varies from grayish to dark brown; ventral pelage slightly paler. There is a wide range of individual variation in both color and spot markings. Albinism seems extremely rare (one albino individual collected in DR Congo). Back exhibits a characteristic pair of whitish yellow spots on shoulders, and irregularly distributed dark spots that may partly coalesce in the nape (nucchal) region. One to three clearly defined dark lines on neck. Limbs are short, thickset, and poorly spotted. Five claws are partially retractile. Feet plantigrade; pads extremely well-developed, reaching wrist in forefoot and ankle in hindfoot. Tail is longer than length of head and body (110-120%), densely furred, and has nine to 15 irregularly marked dark rings; rings are generally incomplete on the underside. Both sexes have a glandular abdominal pouch situated anterior to the vulva or penis, about 55 mm long, 20 mm deep, and 20-30 mm wide when open. It shows a simpler structure than the perineal glands usually found in viverrids, and produces a yellowish liquid with a musky odor comparable to the “musk” ofcivets and genets. Skull differs from other extant carnivorans by the persistence throughoutlife of the cartilaginous condition of the posterior chamber of the auditory bulla. Rostrum is moderately elongated; zygomatic arches are wide and sagittal crest is generally thin and elevated. Dental formula: I 3/3, C 1/1, P 4/4, M 2/2 = 40. Variation in the presence/absence of M?* exists. Chromosome number: 2n = 38; FN = 66. Geographic variations in coat pattern have been described. In gerrardi, dark stripes on neck are absent, body is sparsely and finely spotted, the pair of bright scapular spots is weakly distinct, and dark tail rings are narrow and sharply defined. In arborea, nucchalstripes are present but narrow, lower sides of body are not spotted, and dark tail rings are thin. In intensa, coat color is redder and brighter, spots are intense black and the pair of bright scapular spots is more striking to the eye.

Habitat. Occurs from lowland to montane tropical rainforest, up to 2500 m (e.g. Cameroon, Tanzania). Also present in riparian and deciduousforests, savannah woodlands, and disturbed forests after logging or cultivation.

Food and Feeding. Although predominantly frugivorous, they are able to adjust energetic resources with opportunistic foraging on vertebrates (essentially murids, more rarely pottos), birds’ eggs, and arthropods. In Gabon, average stomach content was about 80% fruit and 20% diverse prey items. May frequently raid crops and poultry; can be found eating carrion. Fruits are generally consumed in 5-10 minutes, followed generally by a rest taken on a branch 10-20 m away; such frugivorous meals are taken approximately every two hours. Transit is fast and fruit remains are eliminated 2-3 hours after consumption, which might contribute to seed dispersal across the forest (in Kenya, supposed to spread twelve species of plants). There have been reports of 12-15 individuals feeding at the same time on the fruit of the parasol tree (Musanga cecropioides). During encounters between genders, females seem to be given priority access to food. Forefeet used extensively when attacking. Prey is usually pressed and held on the substratum, and then killed byseveral rapid bites all over the body. They are reported to drink palm wine directly from vessels in palm trees in Sierra Leone and the Niger Delta. Captive individuals showed attraction for alcohol that may result from consumption of fermented fallen fruits and tree exudates in the wild.

Activity patterns. Nocturnal and arboreal. During the day, usually rests on large horizontal branches and lianas, in elevated tree holes, or in crevices formed by vegetation 12-15 m above ground. They use the ground to cross deforested areas or to forage. Plantigrade locomotion and large, deeply carunculated, transversely ridged pads of hindfeet allow agile tree climbing. Able to climb a smooth post and descend head first by “vertical looping”, i.e. gripping the trunk, spread-eagled, with claws and hairless pads of forefeet, then bringing both hindfeet forward until they touch the forefeet. Tail maybe used as a balancing rod when walking thin branches; although not prehensile, can be wrapped around branches for support. Able to walk on steel wire andjump across gaps up to 1 m wide and as high as 1.8 m. An individual was observed soaring from high branches, with tail extended and legs stretched out, and making a perfect four-point landing.

Movements, Home range and Social organization. Movements of individuals remain poorly understood. In Gabon, males have home ranges between 34 and 153 ha (mean 85 ha), larger than those of females (29-70 ha, mean 45 ha). Home range size in males is dependent on number of available females rather than nutritional resources. Although adult male home ranges often overlap broadly, they consist of distinct sectors. Large males tend to visit all sectors regularly while smaller males stay in their own sectors to avoid dominant males. Home ranges of dominant males may cover home ranges of several females. Adult females have clearly delimited home ranges with narrow overlap zones along borders, where scent marks are deposited. Immature daughters usually remain with their mothers; young males apparently abandon the maternal home range as soon as they are weaned. Dominant males seldom meet, but fights may be extremely violent and fatal to the loser. Sexual competition appears to accentuate weight differ ences in males, with dominant ones often becoming heavy. Non-dominant males tend to be thinner, with smallertesticles and perineal glands as symptoms of socialstress. In Gabon, estimate of minimum mean density was about 5 ind/km®, with higher means in inundated forests and river borders (reaching 8 ind/km?). In Bwindi Impenetrable National Park, Uganda, mean density was higher above 2000 m elevation than below 1800 m (3-32 ind/km? and 2-17 ind/km?, respectively). Usually, individuals sleep alone, but occasionally a male and female can be found resting close to one another. Scent marking plays a significant role in communication. The perineal gland is rubbed on branches by flexing the hindlimbs; the odor can persist for several months. Other scent glands, such as the one located between the third and fourth toe of each foot, may also be used to scent-mark trails. Individuals produce a typical loud, plaintive cry similar to a lengthy “hoo”, which can be heard over a distance of almost 1 km. In Gabon, calls between genders were at their maximum when a female was in estrus, in June.

Breeding. During courtship, a captive pair at close range emitted sounds akin to coughing. In Gabon, births mostly occurred from September to January (wet season and start of short dry season), occasionally in other months, but never during March—June. Gestation is about 64 days; there are two litters per year, usually of two young each. Birth weight averages about 56 g. Neonates have eyes and ears closed; they accompany mother to visit fruit trees as soon as they are able. Theyattain adult size and weight at 6-9 months. Captivity records report maximum longevity of 16 to 18 years.

Status and Conservation. CITES: not listed. Classified as Least Concern on The [UCN Red List. Probably the most common small carnivore in African forests, although field censuses are rare. In Gabon, biomass estimates neared 11 kg /km? which is greater than that of sympatric carnivorans such as genets and civets. Local threats include destruction of individuals caught preying on poultry. They are also hunted for the bushmeat and traditional medicine markets: they are the most common carnivore sold in markets in Equatorial Guinea and Guinea. The skin is used to make ceremonial clothes, wrist-bracelets, and hats. Their association with closed-canopy habitat suggests vulnerability to intense logging, which is occurring at a high rate in western Africa, and to deforestation in eastern and southern Africa. They are preyed upon by larger carnivorans, birds of prey, and pythons. In Cameroon, it is part of the natural reservoir stock for trypanosomiasis. Due to their arboreal mode oflife, African Palm Civets are subject to falls and bone fractures.

Bibliography. Allen, G.M. (1939), Allen, J.A. (1924), Andama (2000), Ansell (1960b), Anstey (1991), Basilio (1962), Bates (1905), Bourliere et al. (1974), Carpaneto & Germi (1989a), Chapuis (1966), Charles-Dominique (1977, 1978), Coetzee (1977), Colyn, Dudu & Mankoto Ma Mbaelele (1987), Colyn, Dufour etal. (2004), Crawford-Cabral (1989), Eisentraut (1973), Engel (2000), Ewer (1973), Flynn (1996), Flynn & Nedbal (1998), Flynn et al. (2005), Gaubert (In press a), Gaubert & Veron (2003), Gaubert, Veron & Tranier (2002), Gaubert, Wozencraft et al. (2005), Gregory & Hellman (1939), Hamerton (1941, 1945), Happold (1987), Henschel (2001), Herder et al. (2002), Hunt (1974, 1987, 1989), IUCN (2008), Jeffrey (1977), Juste et al. (1995), Kingdon (1997), Leyhausen (1965), Malbrant & Maclatchy (1949), McKenna & Bell (1997), Meester et al. (1986), Mertens (1925), Morales et al. (2005), Naughton-Treves (1998), Ngandjui (1998), Ososky (1998), Perkin (2004, 2005), Peters (1984b), Pimley (1999), Pocock (1915f, 1929), Ray & Sunquist (2001), Rettig & Divers (1978), Rosevear (1974), Schouteden (1945), Simpson (1945), Skinner & Chimimba (2005), Smithers & Lobao (1976), Taylor (1970a, 1971, 1974, 1976), Thompson (1858), Thorneycroft (1958), Todd (1967), Van Rompaey (1997), Van Rompaey & Colyn (In press d), Van Rompaey & Powell (1999), Vosseler (1928), Wemmer (1977), Werneck (1948), Wozencraft (1993, 2005), Waurster & Benirschke (1967), Yoder et al. (2003).

Figures

African Palm Civet

Subspecies and Distribution. N. b. binotata Gray, 1830 — from Gambia to DR Congo , including Bioko I. N. b. arborea Heller, 1913 — Kenya , S Sudan , N Tanzania , and Uganda . N. b. gerrardi Thomas, 1893 — Malawi , Mozambique , S&E Tanzania (presumably also Zanzibar I), NE Zambia , and E Zimbabwe . N. b. intensa Cabrera & Ruxton, 1926 — Angola , S DR Congo , and NW Zambia .