Not to be cited or copied without written permission from the authors

Draft Not to be cited or copied without written permission from the authors Colobus rufomitratus Struhsaker Final edit JK & TB Sept 007 Procolobus rufomitratus Eastern Red Colobus, with subspecies having the following names: Tana River Red Colobus, Ashy or Uganda Red Colobus, Tshuapa or Thollon s Red Colobus, Oustalet s Red Colobus, Kisangani Red Colobus, Lulindi River Red Colobus, Kivu Red Colobus, Lomami River Red Colobus, Semliki Red Colobus. Fr. Colobe bai de l est Ger. Ost Afrikanische Stummelaffe Procolobus rufomitratus (Peters, 1879). Monatsb. K. Preuss. Akad. Wiss. Berlin, 1879: 829. Muniuni, Tana River, Kenya. Taxonomy

Polytypic species. Taxonomy extremely complex and much debated (Rahm 1970, Dandelot 1971, Napier 1985, Colyn 1991, Kingdon 1997, Gautier-Hion et al. 1999, Groves 1989, 2000, 2001, Groves & Thorington 1970, Grubb and Powell 1999, Grubb et al. 2003). As treated here includes at least eight subspecies: Tana River Red Colobus P. r. rufomitratus; Ashy (or Uganda) Red Colbous P. r. tephrosceles (synonym: gudoviusi); Tshuapa Red Colobus P. r. tholloni; Oustalet s Red Colobus P. r. oustaleti (synonyms: brunneus, powelli, nigrimanus, schubotzi); Kisangani Red Colobus P. r. langi; Lulindi River Red Colobus P. r. lulindicus; Kivu Red Colobus P. r. foai (synonym: graueri); and Lomami River Red Colobus P. r. parmienteri. A ninth, Semliki Red Colobus P. r. ellioti (synonym: semlikiensis), may represent a very large hybrid swarm between the latter five taxa, Verheyen (1962). Most early authorities considered all to be subspecies of a single species of Red Colobus Colobus badius (e.g., Schwarz 1928, Rahm 1970, Haltenorth & Diller 1980). Dorst & Dandelot (1970) confined badius to a single western species while a second species, pennanti embraced all the rest of the Red Colobus forms. Groves (1989) regarded the nominate population, P. rufomitratus, from the Tana river delta, to be a species in its own right and most of the forms listed above as subspecies of a single Central species, P. pennanti, ranging from the Bight of Benin to Zanzibar island. Groves (2001) again kept Tana rufomitratus as a separate species, one of nine in the red colobus group; tholloni, foai and tephrosceles were also given specific status. Groves (2001) placed lulindicus, ellioti, langi, oustaleti, parmentierorum and semlikiensis as subspeces or synonyms of Piliocolobus (Procolobus) foai. Kingdon 1997 had followed a similar course but provisionally allied tephrosceles with oustaleti and other populations from north and east of the Congo (Zaire) river provisionally

clustering them as subspecies of a Central African Red Colobus P. oustaleti. Gautier- Hion et al. (1999) divided P. r. oustaleti into six subspecies, and placed tholloni, oustaleti, langi, lulindicus, foai, parmienteri, powelli, nigrimanus, and semlikiensis as subspecies of Pennant s Red Colobus Procolobus pennantii. See authors listed above and Napier (1985), Colyn (1991), Grubb et al. (2003) for reviews and additional details. Chromosome number: currently unknown. Description A medium-sized, arboreal monkey with variable amounts of reddish-brown or orangebrown on top of the head, back, tail, and lateral surface of the arms and legs, and paler ventrum. Males generally same or similar color as females. Adult males of some subspecies up to ca. 25% heavier than adult females in some subspecies (e.g. 9-12.5 kg vs 7 9 kg for P. r. tephrosceles; Kingdon 1971, Struhsaker 1975), but this size difference is not present in all subspecies (e.g., P. r. rufomitratus; Struhsaker, pers. obs.). In all subspecies, males have larger canines, and nuchal and sagittal crests (Verheyen 1962, Grubb, pers. obs.). Colour and patterning highly variable among subspecies and often so within a subspecies, such as tephrosceles, oustaleti, and particularly in the ellioti/semlikiensis hybrid swarm. See Geographic Variation for information on extent of phenotypic variation. Back and tail often dark-grey, lacking any red or brown depending on subspecies. Tail typically tufted, especially in adult males. P. r. tholloni is the most brightly colored subspecies with bright orange or reddish over most of the dorsum. Hair usually of medium length. Face dark grey or blackish in most subspecies, but tholloni

distinct from the others by having light skin around the eyes, nose, and mouth, but not as contrasting as in the Udzungwa Red Colobus Procolobus gordonorum or Zanzibar Red Colobus Procolobus kirkii. P. r. tholloni is also distinguished by its more prognathous skull. See Verheyen 1962, Napier (1985), Colyn 1991, Kingdon (1971, 1997), Gautier- Hion et al. (1999), Groves (2001), and Grubb et al. (2003) for detailed descriptions, illustrations, and discussions of the complex of subspecies here recognized within P. rufomitratus. Newborns differ in colour from adults, are all dark gray to blackish on the dorsum and top of head, and light grey below. All newborns so far described for the subspecies within P. rufomitratus (tephrosceles, oustaleti, tholloni) have distinct pink lips and noses that are joined by a pink stripe. These pink parts of the face contrast sharply with the otherwise dark face. This facial pattern is lost in P. r. tephrosceles at about 3-4 months of age, but not lost in P. r. oustaleti until about 2-3 years. Red and brown colors do not appear in the pelage until about 2 months of age in P. r. tephrosceles, but may appear earlier in P. r. tholloni. Adult coloration in P. r. tephrosceles occurs at about 3.5 months (Struhsaker 1975 and unpub. data). There is tremendous variation in color and other phenotypic characters between and within the subspecies of this group (see above). For example, P. r. tephrosceles in the Mbisi Forest of S Tanzania have longer hair than those in Kibale, Uganda, and a much higher incidence of stump tails. Fifteen percent of the Mbisi adults and subadults in a sample of 35 had stump tails, whereas none were seen amongst juveniles (Rodgers et al. 1984). These characters are attributed to the high altitude (2,200m) and southerly latitude (7 0 40 S.) that result in much colder temperatures at Mbisi. Sexual dimorphism,

particularly in body size, is most pronounced in P. r. tephrosceles and least in P. r. rufomitratus. Geographic Variation P.r. rufomitratus. Predominantly grayish-brown dorsum. Darker on shoulders and tail, but paler on rump and lateral surface of limbs. Crown and nape dull orange contrasting with black brow and dark greyish black face. Cow-lick (whorl) of stiff hairs behind each ear). Dorsum dull grayish-brown to charcoal gray. Ventrum grayish-white. P. r. tephrosceles. Resembles rufomitratus; back and tail dark grey to black; lighter grey sides and ventrum; arms and legs grey with varying amounts of brown on lateral surface especially on the upper or proximal part of the arm; brown usually absent from hindlegs; facial whiskers grey. Dark face; rusty-red cap with whorled hair above/behind ear, (sometimes edged with a black tuft) very conspicuous in some individuals, less so in others. Black on brow extends to ears, sometimes up onto crown. Considerable variation in color and some males develop conspicuous cape on shoulders. P. r. tholloni. Circumfacial hair short, black, except on lower cheeks where hair is whitish. Muzzle square and prominent. Crown deep brown or chestnut, upperparts and outer surfaces of limbs orange-russet. Ventrum whitish with yellowish tone. Upper surfaces of hands and feet dark brown. Tail red for about 70% of its length, terminal tuft black.

P. r. oustaleti. Medium sized. Highly variable colouration, but predominantly reddishbrown. Crown reddish-brown; Face dark.. Dorsum varies from dark smoky-brown to olive-brown to coppery- red to dark mahogany. Ventrum paler than dorsum. Hands and feet often black. Tail light brown to black. P. r. langi.. Crown, shoulders and front legs deep orange, reddish or chestnut. Dorsum, hindlegs and tail blackish. Ventrum grey. P. r. lulindicus. Crown reddish. Crest poorly developed. Brow band black. Ears with prominent tufts at base. Proximal dorsum blackish. Distal dorsum and legs reddish. Ventrum grayish. Hands, feet and tail blackish. Pelage short. P. r. foai. Pelage long, with ragged appearance. Ears with prominent tufts at base. Cheeks often streaked with dark brown freckles. Brows black beneath red tuft on forehead; crown reddish. Proximal dorsum blackish. Distal dorsum reddish. Ventrum grayish. Limbs, hands and feet unicolour, light reddish, paler than distal dorsum. Tail reddish. P. r. parmientieri.. Face black with pinkish lips, chin, and base of nose. Ears with prominent tufts at base. Frontal band black. Crown brownish-red without crest. Midback, shoulders, and upper arms black, rest of dorsum and lateral side of arms and legs brick red. Ventrum and throat whitish. Feet and hands black.

P. r. ellioti. Highly variable subspecies and probably represents a large hybrid swarm involving langi and semlikiensis (Colyn 1993). Apart from an orange cap there is no consistent coat colour or pattern (Lorenz von Libernau, L. 1917, Colyn 1993, Struhsaker pers. observ.).. Predominantly reddish-brown. Crown orange-red. Proximal dorsum and shoulders generally reddish. Saddle grey or reddish tones, hind legs and tail dark brown, grey or black. Arms rusty orange or grey. Forearms orange-red or charcoal grey. Similar Species None within the geographic range. Distribution Endemic to Central and East Africa. Rainforest, Afromontane and Afroalpine, and Coastal Forest Mosaic Biotic Zones. From the Sangha R. in W Central African Republic, east to the lower Tana River in E Kenya, north to S Central African Republic and S Sudan, and south to N Zambia and SW Tanzania (Kingdon 1974, Rodgers 1981, Colyn 1991, 1993, and Gautier-Hion et al. 1999, Groves 2001). P.r. rufomitratus. Lower Tana River forests, Kenya. P. r. tephrosceles. Eastern border of Western Rift in Uganda and Tanzania from Kibale National Park, Uganda to Mbisi Forest Reserve, Tanzania (ca. 0 o 41 N 7 o 40 S). P. r. tholloni. South of the Congo R. east to Lomami R, and south to the Kasai-Sankuru R., DRC.

P. r. oustaleti. North of Congo (Zaire) river from Sangha R. in west across the Oubangui R. east to L. Albert. Northern limit is the gallery forests north of the Uele R. and Congo/Nile watershed in S. Sudan. Southeast limit is the Aruwimi-Ituri R., DRC. P. r. langi..lowland forest east of Congo river from about Bumba to Boyoma Falls. North of Maiko R. to valley of Aruwimi R., east to piedmont of Albertine Rift highlands P. r. lulindicus. Riverine (lowland) forest subspecies.western limit is the Lualaba R. Between Lowa-Oso R. in north and Elila R. in the south, DRC. P. r. foai. Montane forest species. Between Lowa R. on west side of L. Kivue and ca. 6 S on west shore of L. Tanganyika, DRC. P. r. parmientieri.. Between Lomami R. and Lualaba R, south to Ruiki R. and Lutanga R. Face black with pinkish lips, chin, and base of nose. P. r. ellioti. From Congo-Lualaba R. east to Semliki Valley, and west shore of L. Edward and L. GeorgeL. Kivu. Northern limit the Aruwimi-Ituri R. Southern limit the Maiko R. Habitat In a variety of forest types, including gallery forest, forest-miombo mosaic, oldgrowth lowland, mid-altitude, and montane moist forest, degraded secondary forests, and raphia palm swamps; ranging in altitude from near sea level to 2,200 m (Struhsaker 1975, Marsh 1978, 1981, Rodgers, et al. 1984, Maisels et al. 1994, Chapman & Chapman 2000a). Most of the research on P. rufomitratus has focused on P. r. tephrosceles in Kibale Forest, W. Uganda, and on P. r. rufomitratus in floodplain/ground water forests along the lower Tana River, E Kenya. In Kibale they are most abundant in moist, oldgrowth, evergreen forest with emergent trees reaching 50 m in height. Prominent trees in

this habitat include, Parinari excelsa, Aningeria altissima, Celtis durandii, Celtis africana, Markhamia platycalyx, Uvariopsis congensis, Funtumia latifolia, Lovoa swynnertonii, Mimusops bagshawei, Albizia grandibracteata, Newtonia buchananii, and Piptadeniastrum africanum (Struhsaker 1975, Chapman & Chapman 2000a). Common trees in the Tana River forests include, Sorindeia madagascariensis, Ficus sycomorus, Diospyros mespiloformis, Albizia gummifera, and Albizia glaberrima. Except for the drier lower Tana River site. P. rufomitratus usually occurs where annual rainfall is >1,000 mm Abundance Extremely common at some sites and rare at others. Population densities of P. r. tephrosceles in Kibale range from about 25 to 300 individuals/km 2 (Struhsaker 1975, 1997, Skorupa 1988, Chapman & Chapman 1999, Chapman & Lambert 2000, Teelen 2005). Densities of P. r. rufomitratus along the Tana River range from 33-253 individuals/km 2 (Marsh 1978, Decker 1994, Mbora 2003). Moderate to heavy levels of selective logging result in very reduced population densities in Kibale (Struhsaker 1975,1997, Skorupa 1986) that persist for at least several decades (Chapman et al. 2000, Struhsaker 2005). Unusually high rates of predation by chimpanzees Pan troglodytes in Kibale, and at Gombe, Tanzania, have also led to major reductions in P. r. tephrosceles populations (Wrangham & Bergmann-Riss 1990, Stanford 1998, Mitani & Watts 1999, Mitani et al.2000, Struhsaker 2005, Teelen 2005). Struhsaker (2005) estimates that there are at least 17,000 P. r. tephrosceles in Kibale alone, plus a few thousand more in the small and isolated forests of W Tanzania.

Forest destruction and fragmentation along the lower Tana River has led to very significant population declines of P. r. rufomitratus, which may now number fewer than 1,500 individuals (Decker 1994, Butynski & Mwangi 1994,1995, Wieczkowski & Mbora 2000, Mbora 2003, Struhsaker 2005). There are no population estimates for other subspecies of P. rufomitratus, but P. r. oustaleti and P. r. tholloni are likely the most numerous because of their vast geographic ranges and large amount of habitat remaining. Adaptations Diurnal and arboreal. See the Subgenus and Genus Procolobus profiles, as well as the Subfamily Colobinae profile for anatomical and physiological adaptations. Foraging and Food Folivorous. Daily travel distance for P. r. tephrosceles in Kibale is highly variable within and between groups, ranging from ca. 180-1,185 m. During a 15-month sample period, the daily travel distance of one group of 22 individuals ranged from 222-1,185 m. Average daily travel distance among five groups in Kibale was between 500 m and 600 m. (Struhsaker 1975, Struhsaker & Leland 1987). Daily distances are similar amongst the smaller groups living in the drier and more seasonal riparian forests of the Tana River, Kenya (Decker 1994).] There is disagreement as to whether or not group size affects travel distance (Struhsaker & Leland 1987, Gillespie & Chapman 2001). Mean annual home range size varies from ca. 35 ha ( (Kibale).- to 100 ha (Gombe) for P. r. tephrosceles (Clutton-Brock 1975, Struhsaker 1975, Struhsaker & Leland 1987), whereas they are only 4-19 ha (for P. r. rufomitratus in the riverine forest patches along the Tana River (Decker 1994). Territoriality is absent amongst the

populations studied and groups often have extensive, if not complete, overlap in home ranges (Struhsaker 2000b). Exceptions exist in heavily logged areas of Kibale where there is little overlap in home ranges and intergroup encounters are rare (Skorupa 1988, Struhsaker 2000b). Feeding occurs throughout the day, but is frequently alternated with periods of rest and travel (Struhsaker 1975, Marsh 1981a). Time spent feeding during the daylight hours is ca. 23-30% for P. r. rufomitratus (Decker 1994) and ca. 45% for P. r. tephrosceles (Struhsaker 1975). Young leaves dominate (~ 30-50%) the diets of all three subspecies for which there are data, i.e., P. r. tephrosceles at Kibale (Struhsaker 1975, 1978, Clutton-Brock 1975, Chapman & Chapman 2000a), P. r. rufomitratus at Tana River (Marsh 1981, Decker 1994), and P. r. tholloni at Salonga N.P., DRC (Maisels et al. 1994). Mature leaves are eaten to a much lesser extent and even then it is usually the petioles and not the lamina that are consumed (Struhsaker 1975, 1978). In the Salonga National Park, DRC, P. r. tholloni eat large quantities of seeds (31% of diet). Seeds sometimes dominate the diet of P. r.tephrosceles in Kibale for periods of several weeks (Struhsaker, pers. obs.). Hundreds of tree and liana species are fed upon. Some of the common food species are: 1) P. r. tephrosceles: C.africana, C. durandii, N. buchananii, M. platycalyx, Aningeria altissima, Milletia dura, L. swynnertonii, P. excelsa, and A. grandibracteata; 2) P. r. rufomitratus: Ficus sycomorus, Sorindeia obtusifoliolata, Acacia robusta, Albizia gummifera, and Pachystela brevipes; and 3) P. r. tholloni: Guibourtia demeusei, Dialium sp., Symphonia globulifera, Cynometra pedicellata, Gilbertiodendron dewevrei, and Daniella pynaertii.

P. r. tephrosceles of Kibale occasionally eat soil from the castings of subterranean termites (Struhsaker, pers. obs.). Social and Reproductive Behaviour Social. Groups of P. r. tephrosceles average about 45-50 individuals (range 8-80,) in Kibale, and 55-59 individuals (range 30-82,) in Gombe (Struhsaker 1975, 2000a,b), but are much smaller in the high-altitude and disturbed Mbisi Forest with a mean of ca. 25 (range <14 30,) (Rodgers et al. 1984). One group of P. r. tholloni numbered at least 60 individuals (Maisels et al. 1994). Groups of rufomitratus in the forest fragments along the Tana River are much smaller, averaging 18 individuals) in the 1970s (Marsh 1979), 11 in the late 1980s and early 1990s (Decker 1994), and 10 (4-31) in 1999-2000 (Mbora 2003). Fission-fusion occurred in a P. r. tephrosceles group in a heavily logged area (Skorupa 1988, Struhsaker 1997, 2000a), and for another group in an unlogged part of Kibale (Chapman & Chapman 2000a). Groups of P. r. tephrosceles in Kibale and Gombe usually contain several adult males (x = 7-8, range 1-13,), but the smaller groups in Mbisi appear to have fewer adult males (Struhsaker 2000a,b). Similarly, the large groups of P. r. oustaleti in the Ituri Forest, DRC, have several adult males (Struhsaker, pers. obs.). P. r. rufomitratus at Tana River are exceptional in that most groups have only one adult male and never more than two (Marsh 1979, Decker 1994). Groups of P. rufomitratus in Kibale and Gombe contain many adult females (x = 18, range 2-32,), and the adult sex ratio (females/male) averages 1.7 to 3.3 (range 1.3-10, (Struhsaker 2000a,b). Adult sex ratios are much higher in the one-male groups of P. r.

rufomitratus at Tana River, averaging 7.3 females/male in the 1970s (Marsh 1979) and 5.1 females/male in the late 1980s and early 1990s (Decker 1994). In general, about 35-50% of group members are immature. Solitary adult and subadult males are uncommon. Temporary all-male parties have been seen only in P.r. rufomitratus (Marsh 1979). Amongst the P. r. tephrosceles in Kibale and P. r. rufomitratus along the Tana River, all females leave their natal groups to join other groups. Parous females transfer between groups and this can occur several times in a lifetime. Even females with 1-2 year-old juveniles occasionally transfer between groups. Female transfers appear to be immediate without any aggression from resident group members (Marsh 1979, Struhsaker & Leland 1979, 1985, Struhsaker & Pope 1991). In contrast, although about half of the P. r. tephrosceles males in Kibale leave their natal groups, they are rarely able to join other groups Most of them disappear and presumably die. Males remaining in their natal groups form a coalition among themselves. There is a dominance hierarchy within these male coalitions, which is expressed through displays, and priority of access to food and space. The highest-ranking male undertakes most, but not all, of the copulations. Females sometimes mate with more than one male. Red colobus are multiple mounters, i.e. a complete copulation with ejaculation is preceded by a series of incomplete mounts by the same male on the same female. No vocalizations are given during copulation, but female P. r. tephrosceles often shake their heads and forequarters back and forth near the end of a complete copulation, perhaps indicating orgasm. Among P. r. tephrosceles, copulating pairs are often harassed by other adult males and juveniles.

Male dominance ranks and mating success change over time. In Kibale, no male has been dominant and the main copulator for more than ca. 3-4 years, but some males were reproductively active for nearly 12 years (Struhsaker 1975, Struhsaker & Pope 1991, Struhsaker 2000b). Coalitions of males in neighboring groups fight one another and the outcome of these encounters may be important in determining patterns of female transfer. Females do not usually participate in these fights nor those within groups (Struhsaker 1975). In contrast, P. r. rufomitratus along the Tana River are exceptional in that adult males are much less tolerant of one another and groups usually contain only one or, occasionally, two adult males. On the Tana, males are able to immigrate and take over groups from incumbent males. All-male groups are uncommon and unstable in the Tana population (Marsh 1979). Allogrooming is common and adult females are the primary groomers. In contrast, allomothering is extremely rare (Struhsaker 1975). Only once was a P. r. tephrosceles female seen carrying another female s infant. She carried this infant along with her own infant for several days until one of the infants disappeared (Struhsaker, pers. obs.). Play amongst immatures is common during their first 2 years. The vocal repertoire size of P. r. tephrosceles is complex and may contain more than 25 different calls, many of which are components of a graded system (Marler 1970, Struhsaker 1975). The repertoires of P. r. oustaleti, P. r. ellioti, and P.r. rufomitratus are less well studied, but they appear identical to P. r. tephrosceles (Struhsaker, pers. obs.). It, appears that P. r. rufomitratus on the Tana vocalize much less than the other subspecies of P. rufomitratus (Struhsaker 1975). This may be a consequence of their smaller, one-male, social groups.

Polyspecific associations are common for P. rufomitratus and have been particularly well documented for the P. r. tephrosceles of Kibale. During line transects 72% of all encounters with red colobus groups in unlogged forest were with other monkey species (Struhsaker 2000a). Although they associated with 5 other monkey species in Kibale, they most typically associated with redtails (Cercopithecus ascanius). It appears that these asssociations are formed primarily in response to predation pressure from crowned eagles Stephanoaetus coronatus and secondarily to habitat quality, the cost to benefit ratio of foraging together, convergence at common foods, and by chance alone. (Struhsaker 1981, 2000b, Chapman & Chapman 2000b). Reproduction and Population Structure Adult females have perineal swellings, which are very large (ca. 7-10 x 7-10 cm) in P. r. oustaleti (Struhsaker, pers. obs.) and much smaller (ca. 1/3 this size) in P. r. tephrosceles and P. r. rufomitratus (Struhsaker 1975). These swellings appear to occur during estrus and ovulation, but also occur at other times, such as during pregnancy (Struhsaker & Leland 1985, Struhsaker, pers. obs.). Estrous P. t. tephrosceles may give off olfactory cues because males often nuzzle a female s perineum before mounting her (Struhsaker 1975). Gestation length is ca. 6-7 months (Struhsaker & Leland 1985). Copulations and births occur in all months, but at Kibale most P. r. tephrosceles births occurred during three relatively wet months (April, May, and November) and one dry month (June) (Struhsaker 1997). Likewise, no seasonality in births was observed during an 18-month study of P. r. rufomitratus (Marsh 1978). Single births are the rule and no twins have

been recorded. Weight at birth unknown, but an infant P. r. tephrosceles estimated to be <4 months old weighed 0.5 kg (Struhsaker 1975). The average interbirth interval for 16 recognizable P. r. tephrosceles in Kibale over 18-years was 24.4 months (n=73 births to 16 females) and 27.5 months when cases of neonatal mortality were excluded (Struhsaker & Pope 1991). A similar interbirth interval appears to occur among the rufomitratus of the Tana based on ratios of infants per adult female (Marsh 1979, Decker 1994, Struhsaker et al. 2004). Weaning is a gradual process and is usually complete in P. r. tephrosceles by about 12-18 months. Reproductive and physical maturity are reached in about 5-5.5 years in both male and female P. r. tephrosceles (Struhsaker & Leland 1985, Struhsaker & Pope 1991). Infanticide by one male was observed during a long-term study of a multi-male group of P. r. tephrosceles (Struhsaker & Leland 1985) and suspected in P. r. rufomitratus when a male replaced the resident male (Marsh 1979). In both cases the evidence supported the hypothesis that infanticide is a reproductive strategy. Mowry (1995) observed possible infanticidal behaviour by a male P. r. rufomitratus, but the infant survived the attack. Population structure is essentially the same as that given for group composition because solitaries are uncommon. Maximum longevity is unknown, but one adult female P. r. tephrosceles was observed for nearly 15 years. Because she was fully adult at the initiation of observations, she was probably at least 20 years old when she died (Struhsaker & Pope 1991). Mean annual birth rate in P. r. tephrosceles is 0.49 (Struhsaker & Pope 1991). Mean survivorship to adulthood is estimated as 39%.

Assuming a 1:1 sex ratio at birth, survivorship to adulthood was 47% for females and 34% for males (Struhsaker & Pope 1991). Predators, Parasites and Diseases Major predators on P. r. tephrosceles are crowned eagles (Struhsaker & Leakey 1990, Mitani et al. 2001) and chimpanzees (Wrangham & Bergmann-Riss 1990, Stanford 1998, Mitani & Watts 1999, Mitani et al.2000, Struhsaker 2005, Teelen 2005). Predation pressure appears to influence population size and age-sex composition. A herpes-like disease afflicted some P. r. tephrosceles in Kibale and apparently killed five out of 10 adult males in one group over a 2.5-year period (Struhsaker 2000b). Predation by humans is particularly important in DRC, e.g. P. r. tholloni (Thompson 2000). Conservation IUCN Category (2007): Critically Endangered as for P. r. rufomitratus. Other subspecies Least Concern. CITES: Appendix I for P. r. rufomitratus. Other subspecies Appendix II. The main threat to P. r. rufomitratus is habitat loss due primarily to agricultural clearing and extraction of forest products by local communities (Decker 1994, Butynski & Mwangi 1994, 1995, Wieczkowski & Mbora 2000, Mbora 2003) and secondarily to alteration of river flow volume and cycles by five hydroelectric power dams upriver (Hughes 1984, Butynski 1995). Less than 35% of the P. r. rufomitratus population is legally protected within the Tana River Primate National Reserve, but even there the habitat is insecure because of inadequate law enforcement (Butynski & Mwangi 1994, 1995, Wieczkowski & Mbora 2000). It is recommended that: 1) the legally

protected area be increased in size; 2) law enforcement be made effective; 3) increased involvement of local community through conservation education; and 4) ecological monitoring increased and made an integral component of conservation management. Other subspecies of P. r. rufomitratus are not on the current Red List. P. r. tephrosceles should be assessed for the Red List because of its restricted range and declining habitat. The largest population of this taxon is in the Kibale National Park where the habitat is well protected. In Kibale and Gombe, however, the greatest threat to P. r. tephrosceles is predation by chimpanzees where it is estimated that they kill between 6.5% and 40% of these monkeys annually Stanford 1998, Watts and Mitani 2002, Teelen 2005. As a result, P. r. tephrosceles populations have declined in both of these Parks in the last 20-25 years (Struhsaker 2005). The Kibale population should be monitored throughout the Park on an annual or semi-annual basis. Status surveys should be done of smaller populations of P. r. tephrosceles in areas such as Mahali Mountains National Park, Mbisi Forest Reserve, and the Biharamulo region of Tanzania. Nothing is known of the population status of the other subspecies of P. rufomitratus. The very extensive range of some, such as P. r. oustaleti and P. r. tholloni, suggest that their conservation status might not be as dire as those taxa living farther east. Thompson (2000), however, describes the very extensive and intensive hunting pressure on P. r. tholloni, which has likely increased with continuous expansion of the bush-meat trade and its extension into rapidly growing urban centres, with the proliferation of weapons and ammunition, all associated with the long-standing conflicts and insecurity in DRC. The same problem likely applies to all other subspecies of P. rufomitratus living in DRC. Obvious conservation actions include: more conservation areas; greater law

enforcement; conservation education; and surveys to better understand the status and conservation problems facing the P. rufomitratus of DRC. Key References Decker, B.S. 1994 Maisels, F. et al. 1994 Marsh, C.W. 1978 Struhsaker, T.T. 1975 Struhsaker, T. T. & Pope, T. R. 1991 Authors Thomas T. Struhsaker Peter Grubb Full References Butynski, T.M. 1995. Report says dam could threaten Kenya s endangered primates. African Primates 1: 14-17. Butynski, T.M. & Mwangi, G. 1994. Conservation status and distribution of the Tana River red colobus and crested mangabey. Unpublished report to the Kenya Wildlife Service and Zoo Atlanta, Nairobi. Butynski, T.M. & Mwangi, G. 1995. Census of Kenya s endangered red colobus and crested mangabey. African Primates 1: 8-10.

Chapman, C.A. & Chapman, L.J. 1999. Implications of small scale variation in ecological conditions for the diet and density of red colobus monkeys. Primates 40: 215-231. Chapman, C. A. & Chapman, L.J. 2000a. Constraints on group size in red colobus and red-tailed guenons: examining the generality of the ecological constraints model. Internatonal Journal of Primatology 21: 565-585. Chapman, C.A. & Chapman, L.J.. 2000b. Interdemic variation in mixed-species association patterns: common diurnal primates of Kibale National Park, Uganda. Behavioral Ecology and Sociobiology. 47: 129-139. Chapman, C. A., Balcomb, S. R., Gillespie, T., Skorupa, J. & Struhsaker, T.T. (2000b). Long-term effects of logging on African primate communities: A 28 year comparison from Kibale National Park Uganda. Conservation Biology 14: 207-217. Chapman, C.A. & Lambert, J.E.. 2000. Habitat alteration and the conservation of African primates: case study of Kibale National Park, Uganda. American Journal of. Primatology 50: 169-185. Clutton-Brock, T.H. 1975. Ranging behaviour of red colobus (Colobus badius tephrosceles) in Gombe National Park. Animal Behaviour 23: 706-722, Colyn, M.M. 1991. L importance zoogeographique du Bassin du Fleuve Zaire pour la speciation: le cas des Primates simiens. Annales sciences Zoologiques Musee Royale de L Afrique Centrale, Tervueren, Belgique 264: 1-250. Colyn, M.M. 1993. Coat colour polymorphism of red colobus monkeys (Colobus badius, Primates, Colobinae) in eastern Zaire: taxonomic and biogeographic implications. Journal African Zoology/ Revue de Zoologie Africaine 107 :301-320.

Decker, B.S. 1994. Effects of habitat disturbance on the behavioral ecology and demographics of the Tana River red colobus (Colobus badius rufomitratus). International Journal of Primatology 15: 703-37. Delson, E., Terranova, C.J., Jungers, W.L., Sargis, E.J., Jablonski, N.G., anddechow, P.C. 2000. Body mass in Cercopithecidae (Primates, Mammalia): estimation and scaling in extinct and extant taxa. Anthropological Papers of the AmericanMuseum of Natural History No. 83. 159 pp. Gautier-Hion, A., Colyn, M. & Gautier, J.-P.. 1999. Histoire Naturelle des Primates D Afrique Centrale. ECOFAC, Libreville. 162 pp. Gevaerts, H. 1992. Birth seasons of Cercopithecus, Cercocebus and Colobus in Zaire. Folia Primatologica 59: 105-113. Gillespie, T.R. & Chapman, C.A.. 2001. Determinants of group size in the red colobus monkey (Procolobus badius): an evaluation of the generality of the ecological constraints model. Behavioral Ecology and Sociobiology 50: 329-338. Grubb, P., Butynski, T.M., Oates, J.F., Bearder, S.K., Disotell, T.R., Groves, C.P. & Struhsaker, T.T.. 2003. Assessment of the diversity of African primates.i International Journal of Primatology. 24: 1301-1357. Hughes, F.M.R. 1985. The Tana River floodplain forest, Kenya: ecology and the impact of development. Ph.D. thesis, University of Cambridge. Kingdon, J. 1974. East African Mammals. vol. 1. Academic Press, London. 446 pp. Lorenz von Libernau, L. 1917 Beiträge zur Kenntnis der Affen und Halbaffen von Zentral-Afrika. Annalen naturh. Mus. Wien 31: 169-241

Maisels, F., Gautier-Hion, A. & Gautier, J.-P.. 1994. Diets of two sympatric colobines in Zaire: more evidence on seed-eating in forests on poor soils. International Journal of Primatology. 15: 681-701. Marler, P. 1970. Vocalizations of East African monkeys. I. Red colobus. Folia Primatologica. 13: 81-91. Marsh, C.W. 1978. Ecology and social organization of the Tana River red colobus (Colobus badius rufomitratus). Ph.D. dissertation, University of Bristol, Bristol. Marsh, C.W. 1979. Comparative aspects of social organization in the Tana River red colobus, Colobus badius rufomitratus. Zeitschrift fur Tierpsychologi., 51: 337-362. Marsh, C.W. 1981. Diet choice among red colobus (Colobus badius rufomitratus) on the Tana River, Kenya. Folia Primatologica. 35: 147-178. Mbora, D.N.M. 2003. Habitat quality and fragmentation and the distribution and abundance of the Tana River red colobus monkey, Procolobus rufomitratus, in Eastern Kenya. Ph.D. thesis, Miami University, Oxford, Ohio. Mitani, J. C., Struhsaker, T. T. & Lwanga, J.S. (2000). Primate community dynamics in old growth forest over 23.5 years at Ngogo, Kibale National Park, Uganda: implications for conservation and census methods. Internatioanl Journal of Primatol-ogy. 21: 269-286. Mitani, J.C. &. Watts, D.P. 1999. Demographic influences on the hunting behavior of chimpanzees. American Journal of Physical Anthropology 109: 439-454. Mowry, C.B. 1995. Possible infanticidal behaviour by a Tana River red colobus Procolobus badius rufomitratus. African Primates 1: 48-51.

Napier, P.H. 1985. Catalogue of Primates in the British Museum (Natural History) and Elsewhere in the British Isles. Part III: Family Cercopithecidae, Subfamily Colobinae, British Museum (Natural History), London. O Leary, R. 2003. An annotated catalog of the African primate genera Colobus and Procolobus (Cercopithecidae: Colobinae) in the collections of the American Museum of Natural History. Novitates No. 3399. 26 pp. Rahm, U.H. 1970. Ecology, zoogeography, and systematics of some African forest monkeys. In: Old World Monkeys. (eds. J.R. Napier and P.H. Napier). Academic Press, New York and London. pp.589-626. Rodgers, W.A., Struhsaker, T.T. & West, C.C.. 1984. Observations on the red colobus (Colobus badius tephrosceles) of Mbisi Forest, south-west Tanzania. African J-ounal of. Ecology. 22: 187-94. Rodgers, W. A., 1981. The distribution and conservation status of colobus monkeys in Tanzania. Primates 22: 33-45. Skorupa, J.P. 1986. Responses of rainforest primates to selective logging in Kibale Forest, Uganda: a summary report. In: Primates: the road to self-sustaining populations (ed. K. Benirschke). Springer-Verlag, New York. Pp. 57-70. Skorupa, J. P. 1988. The effects of selective timber harvesting on rain-forest primates in Kibale Forest, Uganda. Ph.D. thesis, University of California, Davis. Stanford, C. B. 1998. Chimpanzee and Red Colobus: The Ecology of Predator and Prey. Harvard University Press, Cambridge. 296 pp. Struhsaker, T.T. 1975. The Red Colobus Monkey. University of Chicago Press, Chicago. 311 pp.

Struhsaker, T.T. 1978. Interrelations of red colobus monkeys and rain-forest trees in the Kibale Forest, Uganda. In: The Ecology of Arboreal Folivores (ed. G. G. Montgomery). Smithsonian Institution Press, Washington, D.C. Pp. 397-422. Struhsaker, T.T. 1981. Polyspecific associations among tropical rainforest primates. Zeitschrift fur Tierpsychologi. 57: 268-304. Struhsaker, T. T. 1997. Ecology of an African Rain Forest: Logging in Kibale and the Conflict between Conservation and Exploitation. University Press of Florida, Gainesville. 434 pp. Struhsaker, T. T. 2000a. The effects of predation and habitat quality on the socioecology of African monkeys: lessons from the islands of Bioko and Zanzibar. In: Old World Monkeys (eds. P. F. Whitehead & C. J. Jolly). Cambridge University Press, Cambridge. Pp. 393-430. Struhsaker, T. T. 2000b. Variation in adult sex ratios of red colobus monkey social groups: implications for interspecific comparisons. In: Primate Males. Causes and Consequences of Variation in Group Composition (ed. P. M. Kappeler). Cambridge University Press, Cambridge. Pp. 108-119. Struhsaker, T.T. 2005. Conservation of red colobus and their habitats. International Journal of Primatologyl. 26: 525-538. Struhsaker, T. T. & Leland, L. 1979. Socioecology of five sympatric monkey species in the Kibale Forest, Uganda. In: Advances in the Study of Behavior (eds. J. Rosenblatt, R. A. Hinde, C. Beer & M. C. Busnel). Academic Press, New York. 9: 158-228. Struhsaker, T.T. & Leland, L.. 1985. Infanticide in a patrilineal society of red colobus monkeys. Zeitschrift fur Tierpsychologie 69: 89-132.

Struhsaker, T. T. & Leland, L.. 1987. Colobines: infanticide by adult males. In: Primate Societies (eds. B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham & T. T. Struhsaker). The University of Chicago Press, Chicago. Pp.83-97. Struhsaker, T. T. & Pope, T. R. 1991. Mating systems and reproductive success of two African forest monkeys (Colobus badius and Cercopithecus ascanius). Behaviour 117: 182-205. Teelen, S. 2005. The impact of hunting by chimpanzees (Pan troglodytes) on demography and behavior of red colobus monkeys (Procolobus rufomitratus) at Ngogo, Kibale National Park, Uganda. Ph.D. thesis, Yale University, New Haven. Thompson, J.A.M. 2000. Conservation of Thollon s red colobus Piliocolobus tholloni, Democratic Republic of Congo. African Primates 4: 27-32. Verheyen, W.N. 1962 Contribution a la craniologie compare des primates. Musee Royale Afrique Centrale-Tervuren, Belgique Ser 8 Sci. Zool. 105 Watts, D. & Mitani, J.C.. 2002. Hunting behavior of chimpanzees at Ngogo, Kibale Natinal Park, Uganda. International Journal of Primatology, vol. 23, no. 1: 1-28. Wieczkowski, J. & Mbora, D.N.M.. 2000. Increasing threats to the conservation of endemic endangered primates and forests of the lower Tana River, Kenya. African Primates 4: 32-40. Wrangham, R.W. & Bergmann-Riss, E.L. 1990. Rates of predation on mammals by Gombe chimpanzees, 1972-1975. Primates 38: 157-170. Body Measurements Procolobus rufomitratus tholloni

HB (female): 590 (580-600) mm, n=2 T (female): 690 mm, n=1 HF (female): 165 (150-170) mm, n=2 E (female): 35 mm, n=1 WT estimated to be in lower range for Red Colobus at 7 10 kg Average WT estimate for Red colobus at 8-12 kg. Procolobus rufomitratus oustaleti (cf. powelli) HB (male): 540 (455-590) mm, n=17 HB (female): 524 (480-565) mm. n=12 T (male): 726 (650-800) mm. n=17 T (female): 713 (645-790) mm, n=12 HF (male): 186 (170-198) mm, n=17 HF (female): 182 (170-202) mm, n=12 E (male): 39 (35-42) mm, n=17 E (female): 37 (35-40) mm, n=12 WT estimated to be in mid to low range for Red Colobus at 8 11 kg. Procolobus rufomitratus oustaleti (cf. brunneus) HB (male) 555 (550-560) mm, n=2 HB (female): 490 (450-520) mm, n=5 T (male) 700 (630-770) mm, n=2 T (female): 580 (430-650) mm, n=5

HF (male) 178 (160-195) mm, n=2 HF (female): 163 (154-172) mm, n=5 E (male): 30 (25-35) mm, n=2 E (female): 32 (29-35) mm, n=5 Procolobus rufomitratus oustaleti (cf. brunneus) HB (male): 582 (525-610) mm, n = 10 HB (female): 559 (510-585) mm, n=5 T (male): 714 (633-785) mm, n=10 T (female): 709 (650-750) mm, n=5 HF (male): 191 (180-200) mm, n=10 HF (female): 189 (183-203) mm, n=5 E (male): 40 (38-42) mm, n=10 E (female): 39 (35-42) mm, n=5 WT (female): 5.4 (SD=0.5) kg, n=21 Gevaerts (1992) for 21 non-pregnant adult female Procolobus rufomitratus from bushmeat markets in Bondo, Kisangani and Kindu, DRC. These are likely to be mostly P. r. oustaleti, but probably also include some P. r. langi, P. r. lulindicus, P. r. parmentieri, and perhaps other subspecies. Procolobus rufomitratus tephrosceles HB (male): 615 (584-648) mm, n=5 HB (female): 584 mm, n=1

T (male): 691 (660-724) mm, n=5 T (female): 686: mm, n=1 HF (male): 182 (171-191) mm, n=5 HF (female): 171 mm, n=1 E (male): 35 (32-41) mm, n=5 E (female): 29 mm, n=1 WT (male): 10.5 kg (n=1) WT ( female): 5.8 kg (n=1) Kibale Forest, Uganda (Struhsaker1975) Procolobus rufomitratus ellioti (cf. semlikiensis) HB (male): 512 (480-540) mm, n=5 HB (female): 520 (500-540) mm, n=3 T (male): 640 (540-750) mm, n=5 T (female): 643 (600-680) mm, n=3 HF (male): 168 (160-189) mm, n=5 HF (female): 170 (165-175) mm, n=3 E (male): 31 (29-33) mm, n=5 E (female): 30 (29-30) mm, n=3. END NOTE Sources for some measurements?