Philantomba maxwelli

Don E. Wilson & Russell A. Mittermeier, 2011, Bovidae, Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona: Lynx Edicions, pp. 444-779 : 752-753

231.

Maxwell's Duiker

Philantomba maxwelli

French: Céphalophe de Maxwell / German: Maxwell-Ducker / Spanish: Duiker de Maxwell

Taxonomy. Antilope maxwelli C.H. Smith, 1827 , Sierra Leone.

Maxwell's Duiker is sometimes included in P. monticola (particularly in historical reports), although it is now well recognized as a separate species. Reports prior to 2010 listed Maxwell’s Duiker as present from Gambia to Nigeria, but populations from Togo eastwards have been determined to be specifically distinct and are listed under Verheyen’s Duiker (FP. wal teri). Two subspecies recognized.

Subspecies and Distribution.

P. m. maxwelli C.H. Smith, 1827 — Senegal and Gambia to E Ghana (likely limited to the E by the Volta River).

P.m. danei Hinton, 1920 — Yatward and Sherbro Is, Sierra Leone.

Descriptive notes. Head-body 55-75 cm, tail 8-16 cm, shoulder height 32-42 cm; weight 7-5 kg (6:5—11-2 kg) in males and 8-6 kg (6.5-12. 6 kg) in females. Maxwells Duiker is the largest member of the genus Philantomba , although subspecies danei is notably smaller than the mainland form. Females may be up to 15% heavier than males. The body has a distinctively arched back; the forelegs are shorter than the hindlegs, and the head is often held low. The general color is dark brownish-gray, sometimes with a bluish or chestnut-red sheen, with the back darker than the sides. There is significant variation in color between and within populations; this may be age-related, as old animals tend to be paler. The belly and inner thighs are whitish, and this color may extend to the chest, but not the neck. The tail is dark above and white underneath; the lateral hairs are white and tend to grow at right anglesto the tail. The face is generally colored as the body, but the forehead is dark, approaching black in some individuals. The hair on the forehead is quite long, and extends from the crown as a short crest. Distinct pale gray stripes run from the medial corners of the eyes to the crown of the head; these are very narrow above the eyes, widening and becoming more diffuse on the forehead. The contrast between the forehead and superciliary stripes is especially pronounced in young adults, but may be faded or indistinct in old animals. A pair of swollen preorbital glandslies in front of the eyes. The glandular opening is 2-5 cm long and is surrounded by naked skin. The underside of the jaw and upper throat are white. The ears are small and rounded, with a dark brown exterior and white interior. There is a slight coronal crest. Horns are always present in males, but their presence is not consistent in females. In form, the horns are conical and ribbed at the base; the sharp tips usually curve slightly forward. Average horn length in males is 3-5—4-9 cm; maximum recorded length is 6-25 cm. When present in females, the horns are shorter. Most females of subspecies maxwelli are hornless (95% in Liberia; 81% in Ghana); females of subspecies dane: generally have horns. Dental formulais10/3,C0/1,P 3/3,M 3/3 (x2) = 32.

228.

Bush Duiker

Sylvicapra grimmia

French: Céphalophe de Grimm / German: Kronenducker / Spanish: Duiker de sabana

Other common names: Common Duiker, Gray Duiker, Grimm's Duiker

Taxonomy. Capra grimmia Linnaeus, 1758, Cape Colony.

There is considerable geographic variation in this wide-ranging species, and over forty subspecies have been described. Many of these are considered not well founded, butthis species is in need of taxonomic revision. Intermediate variations are often seen where the ranges of two subspecies meet. Eleven subspecies are recognized here; a twelfth subspecies (currently undescribed) may inhabit the higher elevations of Mount Kilimanjaro, Tanzania.

Subspecies and Distribution.

S.c.grmmiaLinnaeus,1758—SSouthAfrica(CapeProvince).

S.c.altivallisHeller,1912—CKenya(AberdareMtsandMtKenya).

S.gcaffraFitzinger,1869—SMozambique,EZimbabwe,Swaziland,Lesotho,NESouthAfrica.

S.g.campbelliaeGray,1843—EGuinea,SMali,NIvoryCoast,SBurkinaFaso,NGhana,Togo,Benin,SNiger,Nigeria,NCameroon,SChad,CentralAfricanRepublic,SWSudan,WUganda,Rwanda,Burundi.

S.c.hindeiWroughton,1910—SEKenyaandNTanzania.

S.c.lobeliarumLonnberg,1919—EUganda,WKenya(restrictedtoMtElgon).

S.c.madoquaRuppell,1835—WEthiopia(highlands),WEritrea.

S.gnyansaeNeumann,1905—SESudan,WEthiopia,EUganda,andWKenya.

S.c.orbicularisPeters,1852—SSomalia,EKenya,Tanzania,EZambia,Malawi,NMozambique.

S.c.splendidulaGray,1871—SEGabon,SRepublicoftheCongo,SDRCongo,NAngola,EBotswana,WZambia,WZimbabwe.

S. c. stemhardti Zukowsky, 1924 — S Angola, Namibia, W Botswana, NW South Africa.

Descriptive notes. Head-body 70-105 cm (males) and 90-115 cm (females), tail 10-20 cm, shoulder height 39-68 cm; weight 9.7-22.4 kg (males) and 10.3-26.3 kg (females). Weights vary regionally; animals from the northern parts of the species’ distribution (including West Africa) tend to be smaller than those from the south. Females are typically 2-4 kg heavier than males. In form, the Bush Duiker generally resembles other savanna-dwelling small antelopes more than other duikers: the back is relatively straight (not arched), the legs are long and slender, the neck is longer, and the head is often held above the line of the back. Pelage characteristics vary widely between locations. Coat length is longest in montane forms from East Africa (notably in the subspecies altivallis and lobeliarum), but is often short and glossy in lowland forms. Overall color is grayish-yellow or sandy brown, but may be very red or pale. Each hair is banded, creating a speckled or grizzled appearance. The underparts and inside of legs are white or pale gray. The legs are generally colored as the body, with a black patch above the hooves and a variable dark stripe on the front of the forelegs. A similar stripe is present on the hindlegs in some specimens. The tail has a variable extent of black on the upper surface (ranging from totally black to just the tip); the underside of the tail is bushy and white. The head is generally more rufous than the body, especially the forehead. The rhinarium is black, and the underside of the jaw is whitish. A black or dark brown blaze runs up the bridge of the nose from the muzzle and is highly variable in length. In some individuals it fades away below the eyes (typical of the subspecies orbicularis, splendidula, and steinhardti), whereas in others it continues clearly to the top of the head (usually seen in subspecies altivallis, campbelliae, and hinder). A whitish ring may be present around the eyes (especially in subspecies orbicularis). The slit-like openings of the preorbital glands are large and conspicuous. The ears are large and pointed; the outer surfaces are gray, the inner surfaces fringed with white hair. The size of the ears appears closely linked to climate and habitat: they are largest in hot, arid regions. The coronaltuft is generally long, narrow, and wispy; its color varies from rufous to black. Horns are generally present in males only; they are slender, straight, and pointed, rising almost vertically from the forehead. Average horn length is 9-9 cm (typical range 7.5-12.5 cm); horns up to 18 cm long have been recorded. Short, stunted horns are occasionally seen in females; nearly 13% of subspecies steinhardti females in eastern Botswana are horned. Dental formulais 10/3, C0/1, P3/3,M3/3(x2)=32.

Habitat. A wide range of mixed woodland and savanna habitat types are used. Bush Duikers may be found from sea level to over 4300 m on Mount Kenya (where the species may be found at the snow line). Cover (shrubbery, thickets, or woodland) is needed for shelter; short open grasslands and woodlands without underbrush are avoided. Rainforests and deserts are also avoided, although forest edges may be used for shelter. Bush Duikers are found in a diverse array of woodlands, including those dominated by Brachystegia—fulbernardia (miombo), Colophospermum mopane and Isoberlinia (mopane), Terminalia, or Acacia. Population densities in Zimbabwe average 1-9-9 ind/km?, and are highest in Baikiaea woodland. Typical population density across the species’ range is 2-3 ind/km®. Bush Duikers frequent seasonally burned savannas; the flush of new growth 1-2 months after the burn may cause duiker population densities to double. Baboons (Papio sp.) will occasionally prey on young duikers; other predators include Leopards (Panthera pardus), Servals (Leptailurus serval), African Wild Dogs (Lycaon pictus), martial eagles (Polemaetus bellicosus), and African rock pythons (Python sebae). If pressed, Bush Duikers will readily take to water; they are strong swimmers and may swim for more than 1 km. They are able to jump over 1-8 m, but usually run beneath obstacles when avoiding predators.

Food and Feeding. A browser. Small, ground-level herbs constitute the main part of the diet, accounting for 656% of feeding time. Leaves, stems, and sometimes the roots of such herbs are all eaten. Fallen, often partially dry leaves and fruits are also consumed. Grass is eaten only very rarely, and then only as very young shoots. Stomach content analysis in eastern Zambia revealed a preference for the leaves of Bauhinia petersiana, Brachystegia spp., Combretum molle, Diospyros kirkii, Julbernardia sp., Securidaca longipedunculata, Strychnos sp., Syzygium sp., Tricalysis sp., and Xeromphis obovata. Fruits from Brachystegia spp., Dichrostachys cinerea, Gardenia subacaulis, Julbernardia sp., Pseudolachnostylis maprouneifolia, Ximenia caffra, and Ziziphus abyssinica are preferentially eaten, as are the flowers from Brachystegia spp., Dolichos spp. or Vigna sp., Julbernardia sp., and Protea angolensis. Observations of foraging Bush Duikers in Zimbabwe revealed a very high preference for leaves of Albizia amara, Commelina welwitschii and C. bengalensis, Grewia flavescens, Pterocarpus angolensis, and Securinega virosa, and the fruits of Acacia robusta, Flacourtia indica, Grewiaflavescens, and Ziziphus mucronata. Resin from Acacia trees may be chewed orlicked from tree trunks. Animal matteris occasionally eaten, including termites, eggs, reptiles, and birds; there are several records of Bush Duikers actively hunting and killing ground-dwelling birds. This species is generally independent of free water, but tends to drink daily if it is available. In the Kalahari Desert, this species feeds on moisture-rich melons (Citrullus lanatus), berries, and tubers that are dug up using the hooves. Hanging fruits may be reached by standing upright on the hindlegs.

Breeding. Reproduction occurs throughout the year across most of the species’ distribution. There is a peak in births during the rainy season (November-March) in southwestern Zimbabwe. Estruslasts 1-5-3 days, but is difficult to detect. Courtship behavior involves frequent chases of the female by the male. Males frequently vocalize with a low-pitched bleat during courtship; females may bleat while running away during preliminary chases. Pursuing males will frequently lick or bite at the hindquarters of the female. As estrus peaks, the female slows her pace and will urinate, allowing the male to perform flehming to determine receptivity. “Laufschlag” (a ritualized foreleg kick) precedes copulation, which is very short (2-5 seconds). Gestation is approximately 200 days (a range of 186-216 days has been accurately recorded in captivity). Births typically occur in early morning, and occasionally in the late afternoon or evening. Litter size is one; weight at birth is approximately 1-7 kg. Infants are darker in color and have woollier coats than adults; they are precocial, but are cached for several days. The initial hiding spotis typically less than 100 m from the birth site. The mother will return to the infant 4-6 times during the day to nurse (such behavior at night has not been studied). Infants begin testing solid food at two weeks. Weaning occurs at three months, although older animals up to 4-5 months may attemptto suckle; the association between mother and infant does not last past six months. There is a postpartum estrus. The average interbirth interval in captive Bush Duikers is 234-259 days. Females experience theirfirst estrus between eight and twelve months of age; males become sexually interested at approximately 16 months. Maximum longevity in captivity is 14-15 years.

Activity patterns. Diurnal, trending towards crespuscular. Activity commences at 05:00 h orslightly thereafter and continues until around 08:00 h, when resting places are returned to. Active foraging commences again in the late afternoon, typically at 16:30-17:30 h, and may continue well into the evening. Night-time resting places in dense cover or against boulders are typically retired to by 23:30 h. Foraging accounts for 7-8 hours of the day, split between morning and evening. Midday activity is most frequent on cool or cloudy days, or during light rain. Activity levels during the day tend to be higher in more open habitats where visibility, and therefore the ability to detect predators, is greater.

Movements, Home range and Social organization. Primarily solitary. Associations between males and females only occur during estrus, and may last for several days. Observed groups of three are usually an adult male accompanying an estrous female and her most recent offspring. Males and females occupy individual home ranges. Ranges of females may overlap with those of other females as well as males, but males appear to exclude other males from their home ranges. Preorbital gland secretions are used by males to mark their territories; the boundaries are defended through combat. Dung middens are not used, although areas with relatively high fecal density often occur around fruiting trees (a function of increased time spent in the area). Home ranges typically have areas of dense vegetation at their core, containing several resting places that are frequently used. There is little variation in home range size between the sexes. In South Africa, territories are 0-21-0-27 km?; in Zimbabwe, the territories of males average 0-14 km? and female home ranges 0-13 km? When two duikers meet, such as when both are foraging around fruiting trees, they may mutually groom each other and remain together for up to several hours before parting. Bush Duikers may associate with baboons (Papiosp.), foraging on leaves and fruit dropped from the treetops by the primates.

On following pages: 229. Sahel Duiker (Sylvicapra pallidion); 230. Crowned Duiker (Sylvicapra coronata); 231. Maxwell's Duiker (Philantomba maxwell); 232. Verheyen's Duiker (Philantomba walter).

Status and Conservation. Classified as Least Concern on The IUCN Red List. The Bush Duikeris one of the most common and widespread African antelopes. The total population has been estimated at 1,660,000 animals, although it has been suggested that the actual number could be twice this figure. The population trend appears to be stable, in part due to this species’ ability to colonize a wide variety of habitat types and its relative resiliency to hunting.

Bibliography. Grubb & Groves (2001), Happold (1987), IUCN/SSC Antelope Specialist Group (2008ba), Kingdon (1982), Weigl (2005), Wilson (2001).

229.

Sahel Duiker

Sylvicapra pallidior

French: Céphalophe du Sahel / German: Sahel-Kronenducker / Spanish: Duiker del Sahel

Other common names: Chad Crowned Duiker

Taxonomy. Sylvicapra grimmia pallidior Schwarz, 1914, Mani, Lower Shari River.

Formerly included as a subspecies of S. grimmia, but is considered to be a separate species here. Scientific literature often combines reports of S. pallidior with the neighboring S. grimmia campbelliae to the immediate south. Monotypic.

Distribution. S Chad, N Central African Republic, C Sudan, and W Ethiopia. Further research may indicate a range extension west of Lake Chad.

Descriptive notes. No specific measurements available. Size is similar to small subspecies of the Bush Duiker (S. grimmia). Western individuals tend to be larger than those from eastern parts of the species’ distribution. The overall coloration is a pale sandy brown or buff, significantly paler than the Bush Duiker. The mid-dorsal region is darker than the sides. Banding of hairs is reduced, creating a weakly speckled appearance. The undersides and inner surfaces of the legs are white, and this coloration always extends down to the hooves. The forelegs have a diffuse dark stripe on their front surface for most of their length. The distal half of the tail is dark; the underside is white. The face is the same buffy color as the body, with a dark facial blaze extending from the muzzle to the top of the head. The blaze is more diffuse and the ears are longer than in neighboring the Bush Duiker subspecies campbelliae. There is a crest of long hair at the top of the head. Horns,similar in form to Bush Duikers, are only present in males. Dental formulais10/3,C0/1,P 3/3, M 3/3 (x2) = 32.

Habitat. Sahelo-Sudanian savannas with interspersed woodland. Characteristic woodland tree species (recorded in northern Central African Republic) are predominantly Acacia sp., with Vitellaria paradoxa, Isoberlinia doka, Combretum hypopilinum, Anogeissus leiocarpus, Afzelia africana, Terminalia sp., and Burkea africana. Population densities of 0-02-0-1 ind/km? have been estimated in this habitat.

Food and Feeding. There is no specific information available for this species, but presumably a browserlike the Bush Duiker.

Breeding. There is no specific information available for this species, but likely similar to the Bush Duiker. A captive individual identified as S. pallidior lived at least 6-5 years.

Activity patterns. There is no specific information available for this species, but probably diurnal/crepuscular like the Bush Duiker.

Movements, Home range and Social organization. There is no specific information available for this species, but probably solitary like the Bush Duiker.

Status and Conservation. Classified as Least Concern on The IUCN Red List (under S. grimmia). Population estimates are unreliable due to confusion with S. grimmia campbelliae, which may occur in close proximity to the Sahel Duiker.

Bibliography. Bouché et al. (2009), Fay et al. (1990), Grubb & Groves (2001), IUCN/SSC Antelope Specialist Group (2008ba), Spinage (1988), Weigl (2005).

230.

Crowned Duiker

Sylvicapra coronata

French: Céphalophe couronné / German: Senegal-Kronenducker / Spanish: Duiker coronado

Taxonomy. Cephalophora coronata Gray, 1842, Western Africa.

Formerly considered a subspecies of S. grimmia, S. coronata is herein considered to be a separate species. The Bush Duiker subspecies S. grimmia campbelliae and the Sahel Duiker (S. pallidior) are sometimes attributed to S. coronata in literature reports, although this species, strictly speaking, is restricted to coastal West Africa. Monotypic.

Distribution. Senegal, Gambia, Guinea-Bissau, and Guinea.

Descriptive notes. No specific measurements available, but bodysize is generally smaller than in the Bush Duiker (S. grimmia). The build is trim, with long, slender legs. The pelage is bright orange-yellow. Only scattered hairs have black tips, and thus speckling or grizzling of the coat is minimal. The mid-dorsal region tends to be darker and have a reddish sheen. The underparts are yellowish-white, and the upper halves of the limbs are whitish on their inner surfaces. All four legs are colored as the body, with dark brown patches just above the hooves. A faint brown vertical stripe marks the front of the forelegs, beginning at the hoof-patch but not reaching above the carpus. The tail has a dark brown or black tip. The ears are short compared to the Bush Duiker. A deep red facial blaze runs from the muzzle to the eyes, but it does not extend to the top of the forehead. There is a long coronal tuft. Vertically oriented spike-like horns are present in males only. Dental formula is 10/3, C0/1,P 3/3, M 3/3 (x2) = 32.

Habitat. Savanna woodland and forest—savanna mosaics, typically dominated by Isoberlinia trees. Areas with secondary grass growth are preferred, and the color of the coat provides excellent camouflage in dry grass. Sufficient coveris a necessary habitat component. Crowned Duikers are generally present in low densities across their distribution; an estimate of 0-2 ind/km? was obtained from Niokolo Koba National Park in Senegal.

Food and Feeding. There is no specific information available for this species, but likely a browser.

Breeding. There is no specific information available for this species, but records from captivity indicate a life span of at least six years.

Activity patterns. There is no specific information available for this species, but presumably diurnal or crespuscular like the Bush Duiker.

Movements, Home range and Social organization. There is no specific information available for this species, butlikely similar to the Bush Duiker.

Status and Conservation. Classified as Least Concern on The IUCN Red List (under S. grimmia). Widespread across its limited range, Crowned Duikers occupy most of their historical distribution, although the population density is much lower than it once was. The species has been assessed as uncommon in Gambia and Guinea-Bissau, and common in Guinea. The Senegal population is thought to number at least 2000 animals. This species is still locally common, especially in protected areas like Niokolo Koba National Park. Principal threats include hunting and habitat loss, but the population appears to be stable.

Bibliography. East (1999), Galat et al. (1992) Galat-Luong et al. (2006), Grubb & Groves (2001), IUCN/SSC Antelope Specialist Group (2008ba), Lydekker (1914), Weigl (2005), Wilson (2001).

231.

Maxwell's Duiker

Philantomba maxwelli

French: Céphalophe de Maxwell / German: Maxwell-Ducker / Spanish: Duiker de Maxwell

Taxonomy. Antilope maxwelli C.H. Smith, 1827, Sierra Leone.

Maxwell’s Duiker is sometimes included in P. monticola (particularly in historical reports), although it is now well recognized as a separate species. Reports prior to 2010 listed Maxwell's Duiker as present from Gambia to Nigeria, but populations from Togo eastwards have been determined to be specifically distinct and are listed under Verheyen’s Duiker (FP. wal teri). Two subspecies recognized.

Subspecies and Distribution.

P.m.maxwelliC.H.Smith,1827—SenegalandGambiatoEGhana(likelylimitedtotheEbytheVoltaRiver).

P.m. danei Hinton, 1920 — Yatward and Sherbro Is, Sierra Leone.

Descriptive notes. Head-body 55-75 cm, tail 8-16 cm, shoulder height 32-42 cm; weight 7-5 kg (6:5—11-2 kg) in males and 8-6 kg (6.5-12.6 kg) in females. Maxwell's Duiker is the largest member of the genus Philantomba, although subspecies dane: is notably smaller than the mainland form. Females may be up to 15% heavier than males. The body has a distinctively arched back;the forelegs are shorter than the hindlegs, and the head is often held low. The general color is dark brownish-gray, sometimes with a bluish or chestnut-red sheen, with the back darker than the sides. There is significant variation in color between and within populations; this may be age-related, as old animals tend to be paler. The belly and inner thighs are whitish, and this color may extend to the chest, but not the neck. The tail is dark above and white underneath; the lateral hairs are white and tend to grow at right angles to the tail. The face is generally colored as the body, but the forehead is dark, approaching black in some individuals. The hair on the forehead is quite long, and extends from the crown as a short crest. Distinct pale gray stripes run from the medial corners of the eyes to the crown of the head; these are very narrow above the eyes, widening and becoming more diffuse on the forehead. The contrast between the forehead and superciliary stripes is especially pronounced in young adults, but may be faded or indistinct in old animals. A pair of swollen preorbital glandslies in front of the eyes. The glandular opening is 2-5 cm long and is surrounded by naked skin. The underside of the jaw and upper throat are white. The ears are small and rounded, with a dark brown exterior and white interior. Thereis a slight coronal crest. Horns are always present in males, but their presenceis not consistent in females. In form, the horns are conical and ribbed at the base; the sharp tips usually curve slightly forward. Average horn length in males is 3.5-4.9 cm; maximum recorded length is 6-25 cm. When present in females, the horns are shorter. Most females of subspecies maxwelli are hornless (95% in Liberia; 81% in Ghana); females of subspecies dane: generally have horns. Dental formula is 10/3, C0/1,P 3/3, M 3/3 (x2) = 32.

Habitat. Wide-ranging in both primary and secondary tropical forests in West Africa. Maxwell’s Duiker is highly adaptable and may be found in relic forest patches, gallery forests, and farmbush, where farm plots are interspersed with brushy thickets. Open fields and bamboo thickets tend to be avoided. The introduced Akyeampong weed (Chromolaena odorata ) provides a significant source of cover in disturbed areas and allows Maxwell's Duikerto live in close proximity to human settlements. This species rarely ventures more than 20-30 m from cover. In Ivory Coast, population densities in primary forest have been estimated to be 63 ind/km?; in secondary forest densities tend to be higher, approximately 79 ind/km?. In Ghana and Liberia, population densities are much lower, estimated at 10 ind/ km? even in protected areas.

Food and Feeding. Primarily frugivorous, with fleshy fruits and leguminous seeds comprising, on average, 76-9% ofthe diet (based on eight stomachs collected near Tai National Park, Ivory Coast; this averagerises to 87-9% if one outlier is excluded). Leaves and flowers are also regularly consumed; these are eaten in a significantly greater proportion during the rainy seasons when fruit is scarce. Over 78 fruiting plant species have been identified in the diet of Maxwell's Duiker in Ivory Coast. Studies performed in central Ivory Coast (near Toumodi) indicate that preferred fruiting species during the dry season include Alchornea cordifolia, Blighia sapida, Ficus capensis, Griffonia simplicifolia, Mangifera indica, Nauclea latifolia , and Phoenix reclinata; during the rainy season, fruits of Canthium vulgare, Spondias mombin, Nauclea latifolia , and Blighia sapida comprise the greatest proportion of fruits eaten. In other regions of Ivory Coast, fruits of Dialium aubrevillei and Musanga cecropioides and seeds from Turracanthus africanus are heavily utilized. Maxwell’s Duiker is unable to ingest large fruits; the maximum size that can be swallowed whole is approximately 3 cm. Larger fruits must be soft enough for the flesh to be chewed off. Browse species in the diet include Thonningia sanguinea, Landolphia owariensis, Homalium aylmeri, Canthium vulgare, and the cultivated Manihot esculenta . Blossoms are frequently consumed; fungi and bark are eaten in small quantities. Because many of these food sources are most plentiful in the forest canopy rather than at ground level, Maxwell’s Duikers will forage beneath groups of primates, picking up dropped and dislodged items. Several species of ants, particularly Paltothyreus tarsatus and Oecophylla longinoda, are opportunistically consumed.

Breeding. Breeding occurs throughout the year in Ghana, although in Ivory Coast most births occur during the dry seasons, from January to March and from August to September. During courtship, the male chases the female, licking or biting at her tail. As the female becomes increasingly receptive, her pace slows and the male begins to perform ritualized leg-kicking (“laufschlag”); he determines female receptivity by testing her urine with the flehmen response. Copulation is brief. Gestation is approximately 205 days (from four reliable captive records). Litter size is one, and neonates weigh 620-954 g. Births typically occur in the early morning or late afternoon. Infants andjuveniles are much darker in color than adults, with distinct white markings above the hooves. Although they can stand soon after birth, infants spend the majority of time cached in an isolated and hidden spot, usually at the base of a tree (especially between buttress roots). The mother visits regularly to nurse the infant. The same bedding spot may be used for over a month, and the infant remains immobile there unless approached within 3 m. After two weeks, the infant becomes increasingly mobile and begins nibbling on dead leaves; at four weeks, the juvenile forages daily but consistently returns to the hiding spot to spend the night. Weaning occurs around two months and the caching bed is abandoned by three months. Females may reach sexual maturity as early as eight months, although 10-12 months is more typical. In captivity, this species may live up to 18 years.

Activity patterns. Diurnal. Activity levels are highest in the early morning (06:30-08:00 h) and late afternoon (16:30-18:00 h). A general trend to lower activity levels is seen during midday (10:00-14:00 h), although 60% of observations are still of active individuals. In captivity, Maxwell's Duikeris active for 64% of daylight hours (06:30 18:00 h) and only 21% of the hours of darkness (18:30-06:00 h). Females tend to be less active than males. Night-time is spent bedded down in dense cover;if disturbed, individuals will move away and then settle down quickly. The same sleeping spotis rarely used for two consecutive nights.

Movements, Home range and Social organization. Maxwell's Duiker is a highly sedentary species, with individuals inhabiting stable home ranges that are typically 0-05 km?, but may be as small as 0-03 km?®. There is no significant difference in home range size between the sexes. Home ranges in primary forest tend to be larger than in secondary forest. Networks of paths are used within the home range. Adult males make occasional forays outside of their home range boundaries, usually returning via a circular route within 3-5 hours. Such travel is very different from the typical slow foraging within the home range and may involve reproductive or territorial activity. The basic social unit is a male-female pair; maximum recorded group size is five (one male, two adult females, and two juveniles). A bonded pair effectively shares a home range, with an overlap of approximately 80% between the two individuals. Adjacent pairs have distinct home ranges with less than 1% overlap across seasons. Home range boundaries tend to be stable across years. Borders are marked with latrine areas; males may also horn vegetation at the boundaries. Males are territorial and frequently (5-6 times/ 10 minute interval) mark boundary objects with their preorbital glands (females also mark, but at half that rate: 2-8 times/ 10 minutes). Males do nottolerate the presence of other males at the periphery of the home range. Agonistic interactions between males are brief but very intense and lack a ritualized pre-combat stage. Fights generally last less than two minutes, and involve head-to-head collisions that may result in one competitor being flipped into the air with the force of impact. Intense chases usually follow. Paired individuals often reciprocally groom and mark each other using preorbital gland secretions. In the wild, pairs account for 39% of observations; however, records of solitary animals (seen in 53% of observations) may be misleading: the second partner may not be visible to the observer. Pedal scent glands may play a role in following or locating other duikers. Night-time observations are primarily of solitary individuals resting. Despite maintaining a close association with a single partner, males will mate with other females. While walking or foraging, the tail is constantly flicked up and down,flashing the white underside; it has been suggested that this is a visual cue for paired individuals, although visibility in some habitats is too low for visual signals. When startled, Maxwell’s Duiker freezes in position, even if in mid-stride; the tail also ceasesits regular motion. Only two vocalizations are known: an alarm whistle and a loud bleat.

Status and Conservation. Classified as Least Concern on The IUCN Red List. Maxwell's Duiker is common throughout most ofits range; the total population is estimated to be several million individuals. The adaptability ofthis species to disturbed habitats has enabled it to continue to thrive despite large tracts of original habitat being modified or destroyed. Hunting for bushmeatis the principal threat to survival.

Bibliography. Aeschlimann (1963), East (1999), Hofmann & Roth (2003), IUCN/SSC Antelope Specialist Group (2008ar), Newing (1994), Ralls (1969, 1973, 1975), Weigl (2005), Wilson (2001).