The Department of Mammalogy of the American Museum of Natural History houses over 300 specimens of African colobus monkeys (genera Colobus and Procolobus). The collection consists of red, olive, and black-and-white colobus (Procolobus badius, P. verus, Colobus angolensis, C. guereza, C. polykomos, C. satanas, and C. vellerosus). Taxonomic notes, pelage descriptions, and range are noted for each species and subspecies. For each specimen, the following data are given: catalog number, date collected or recorded, sex, age, nature of specimen, locality, name of collector, and measurements.
INTRODUCTION
The American Museum of Natural History currently houses over 300 specimens of African colobus monkeys. Herbert Lang and James P. Chapin collected the majority of the red colobus and over half of the black-and-white colobus during the American Museum Congo Expedition (1909–1915). All of these specimens were collected from the northeast of the Democratic Republic of the Congo, near Faradje. A map of the collecting sites of the expedition was published by Allen (1925). The remainder of the colobus collection represents a spotty sampling from selected geographic areas.
The collection consists mostly of specimens preserved as skins and skulls. There are also skeletons, casts, wet specimens, and mounted specimens present. The collection contains the type series of Colobus langi (Allen, 1925), consisting of the holotype (Lawrence, 1993) and two paratypes. This material is listed herein under the heading for Procolobus badius ellioti, currently regarded by Napier (1985) as a subjective senior synonym.
The classification used in this catalog for the black-and-white colobus follows Oates et al. (1994) in separating these species at the generic level from the red and the olive colobus, and in recognizing five species. At the subspecific level, the classification used by Napier (1985) is followed, except where noted. All five species of black-and-white colobus are represented in the museum collection. These are Colobus angolensis, C. guereza, C. polykomos, C. satanas, and C. vellerosus.
The taxonomy of the red colobus is unstable and has yet to be fully resolved. The classification used in this catalog is conservative and follows Oates et al. (1994) at the generic level. The red colobus is recognized as a subgenus, Piliocolobus of the genus Procolobus. At the specific level, the catalog follows Napier (1985) in recognizing only two species, P. badius and P. kirkii. Procolobus kirkii is considered by many authors to be a valid species, and although P. badius may in fact be comprised of several species, it is recognized here as a single species. This arrangement conflicts with Groves (1993), who recognizes four species, and others (Dandelot, 1971; Delson et al., 1982).
Seven recognized subspecies or populations of red colobus are represented in the collection: Procolobus (Piliocolobus) badius badius, P. b. bouvieri, P. b. ellioti, P. b. oustaleti, P. b. rufomitratus, P. b. tephrosceles, and P. b. tholloni. A second species of red colobus, P. kirkii, is not represented in the AMNH collection.
Only one species of olive colobus is recognized, Procolobus (Procolobus) verus, of which there are two specimens in the collection.
The primary purpose of this catalog is to record and present the data on all the colobus specimens in the AMNH. Recent reviews of colobus taxonomy have raised several subspecies to specific status (e.g., Groves, 2001). However, since no consensus has been reached and since this catalog is not meant as a review of colobus taxonomy, I have followed a conservative classification throughout. Similarly, pelage descriptions refer only to specimens in the collection, not to the species or subspecies as a whole. It is hoped that the information contained here will be a useful resource for colobus studies and assist in the resolution of the taxonomy of these primates.
Taxonomic Notes
In 1781, Pennant published the first description of two colobus species—the “full-bottom monkey” (the black-and-white colobus) and the “bay monkey” (the red colobus). The descriptions were based on one specimen of each species in the Leverian Museum, London, brought from Sierra Leone. The type specimens were subsequently lost during the dispersal of the Leverian collection. Later authors (e.g., Rochebrune, 1886–1887) suggested that Pennant's description and figures were unreliable. These factors, among others, contributed to the ensuing confusion regarding colobus nomenclature and classification.
The taxonomy of the African colobus has undergone a number of changes in the more than two hundred years since Pennant's publication. The debates center on the number of genera that should be used to represent the African colobus and the number of species that should be used to reflect the diversity of the red colobus and the black-and-white colobus.
The black-and-white, red, and olive colobus are separated at the generic and subgeneric levels. Initially, all three were considered to be members of a single genus (Martin, 1841) but subsequent research has revealed significant differences between each group. Pocock (1936) examined the external characters of the red colobus and compared the results to work he had carried out previously on black-and-white colobus (Pocock, 1926). His data led him to suggest that the two should be distinguished generically. Hill (1952) conducted studies of the external and visceral anatomy of the olive colobus and concluded that his findings suggested a reason to separate verus (the olive colobus) from badius (the red colobus). Many similarities between the red and olive colobus were being uncovered at this time and there was a need to represent these shared features in the taxonomy. Hill and Booth (1957) suggested uniting Procolobus (the olive colobus) and Piliocolobus (the red colobus) within the genus Procolobus based on their distinctive shared features. These features include sexual swelling in the female, the presence of a perineal organ in the male, separate ischial callosities in the female, and small larynx (Pocock, 1936; Hill, 1952; Hill and Booth, 1957; Kuhn, 1967, 1972). All of these features were observed to be common to the red and the olive colobus, and to distinguish both from the black-and-white colobus. In addition, Clutton-Brock (1974) and Struhsaker and Oates (1975) noted significant differences in the behavior and ecology of the red and the black-and-white colobus in the Kibale Forest, Uganda, which also suggested that the two groups might not be members of the same genus.
Currently, Oates et al. (1994) and others (Brandon-Jones, 1984; Strasser and Delson, 1987) agree that the red and the olive colobus monkeys are more closely related to each other than either is to the black-and-white colobus. These authors recognize two genera—Colobus, for the black-and-white colobus and Procolobus to represent the red and the olive colobus. Procolobus is further subdivided into two subgenera, Piliocolobus (red) and Procolobus (olive). This follows an arrangement suggested by Hill and Booth (1957). It is also the arrangement followed in this catalog.
The taxonomy of the black-and-white colobus has been worked out with some resolution. At least four and, more recently, five distinct species are recognized (Oates et al., 1994; Groves, 2001). The elevation of vellerosus to specific status was suggested by Dandelot (1971) and investigated further by Oates and Trocco (1983), whose findings led them to conclude that vellerosus is a distinct species.
The taxonomy of the red colobus at specific level has yet to be resolved with satisfaction. The same 14 forms are recognized by many authors (Rahm, 1970; Dandelot, 1971; Napier, 1985). However, there is no agreement yet on which of these should be assigned specific or subspecific status. It is generally agreed that the Zanzibar red colobus, P. kirkii, is sufficiently distinct morphologically and ecologically from the other red colobus to be distinguished at the specific level (Verheyen, 1962; Napier, 1985).
A comparison of classifications for the red colobus can be found in Oates et al. (1994), while Groves (2001) contains a more recent classification. The number of species used to describe the group varies considerably. Rahm (1970) recognized only one species, Dandelot (1971) eight species, and Napier (1985) two species. More recently, Colyn (1991) suggested that the subspecies ellioti may actually consist of four distinct subspecies (langi, lulindicus, foai, and semlikiensis) together with subspecific hybrids. Groves (2001) recognized nine species.
There has been little resolution to this subject, however, and Oates et al. (1994) observed that an appropriate classification may not be reached until further comprehensive research is carried out.
There is only one recognized species of olive colobus, Procolobus (Procolobus) verus.
Format of the Catalog
The African colobus collection in the AMNH is arranged according to genus and subgenus; the two genera are Colobus, the black-and-white colobus, and Procolobus, including the subgenera Procolobus (the olive colobus) and Piliocolobus (the red colobus). This catalog followed the format used in An annotated catalogue of Malagasy primates… (Buettner-Janusch and Tattersall, 1985). For each level of classification, information was given under the following headings:
Taxonomic Notes: As discussed above, due to the unstable nature of red colobus systematics in particular, there are a number of different classifications proposed at both the specific and subspecific level. Under this heading, alternative classifications of the groups and the opinions of different authors are presented.
Pelage Notes: Different formats of description were used for the red and the black-and-white colobus in order to best describe features of the pelage of each group. For the red colobus, a description of coat coloration was given for each subspecies if relevant, followed by the distinguishing characteristics of the pelage of each subspecies. Wherever possible, the pelage description followed this order: head, cheeks, nape, shoulders, back, sides, underparts, forelimbs, hind limbs, tail. In the case of the black-and-white colobus, general pelage descriptions were outlined for each species. The key formatted by Napier (1985) was used to distinguish subspecies in C. angolensis and C. guereza. Napier's key used the most appropriate features of the pelage to describe the variation in each subspecies. As mentioned previously, the pelage descriptions referred only to the specimens in the AMNH collection.
Species Lists
Eight columns were used to describe each taxon in this collection, under the following headings:
Catalog Number: The number assigned by the museum to each specimen.
Date: This was the date of collection of wild-caught specimens or, in the case of zoo specimens, the date on which they were recorded in the permanent catalog of the AMNH.
Sex and Age: Information on sex was taken from the field tag and confirmed from the actual specimen whenever possible. Three age classes were recorded: adult, juvenile, and infant. In four instances, the collector's tag noted a specimen as an embryo. Following Napier (1985), an adult was defined as having complete permanent dentition. Adults included individuals whose canines were not fully grown. An infant was defined as having entirely deciduous dentition, without M1 erupted. A juvenile included all stages of development between adult and infant, as well as individuals with nonerupted M3. For some of the specimens without skulls, the age was inferred from the skin size. In these cases, the age was enclosed in parentheses.
Description: Skull, skin, and skeleton were noted when present. Skull included cranium and mandible; specimens with anything less were noted as partial skull. Skin denoted whole skin, either flat or mounted.
Locality: Data were taken from field tags, and the localities were verified in a modern atlas. Field notes and gazetteers of the original collectors were also used as references for locality information. When available, the altitude of the locality was also included. Allen (1925) included a map of localities from the Lang and Chapin expedition which was useful for many of the specimens in this collection. If information was not available, or if the specimen was taken from a zoo, “No Data” was entered in this column.
In the case of one species and three subspecies (i.e., Colobus polykomos, C. angolensis cottoni, C. guereza guereza, and C. g. kikuyuensis) specimens without exact locality data were included in the list. This was done when the pelage characters corresponded accurately with the descriptions given by Napier (1985) and Dandelot (1971) and with the other specimens of that species or subspecies in the collection.
The Democratic Republic of the Congo was abbreviated throughout as DRC. This country was known as the Belgian Congo at the time most of the specimens were collected. It was later called Zaire. The Republic of Congo was noted as Congo (Brazzaville) to distinguish the two countries. This country was formerly known as French Congo.
Remarks: In this column the name of the collector was listed, along with the original field number of the specimen. All zoo specimens were from the NYZS (New York Zoological Society) and were noted accordingly.
Measurements: The measurement column lists data taken from the collector's field tag and includes total length, tail, hind foot, and ear length. The measurements were given in millimeters for most specimens and were included in parentheses in inches if originally measured as such by the collector.
Specimens taken from zoos or without sufficient data to enable identification at the subspecific level are included at the end of the appendix, nos. 22–25.
Gazetteer
Geographic coordinates for the majority of the collecting localities were listed in the gazetteer. A map of the central African localities (fig. 1) included the distributions of the specimens collected from that area.
GENUS COLOBUS ILLIGER, 1811
Taxonomic Notes: The genus Colobus represents only the black-and-white colobus monkeys. For some time, four species were recognized: Colobus angolensis, C. guereza, C. polykomos and C. satanas However, Oates and Trocco (1983) suggested elevating C. polykomos vellerosus to specific status based on their study of vocalizations, cranial dimensions, coat pattern, and geographical distribution. Following their findings, five species of black-and-white colobus are recognized in this catalog.
Pelage Notes: The five species of the genus Colobus were easily identified by their black-and-white or all black (C. satanas) coats. The black-and-white colobus are large monkeys and have long tails which in C. guereza are tufted. The infant pelage contrasts markedly with the adult pelage, being completely white at birth in all species except C. satanas whose coat is brown at birth (Oates et al., 1994). Each species has distinctive pelage characters; for example, C. angolensis has shoulder hairs or epaulettes, which readily distinguish it from other Colobus species. The various forms of these features were also used to identify subspecies.
Range of Specimens: The specimens of the genus Colobus in the museum were collected from Liberia, Ivory Coast, Cameroon, and Gabon in the west and across the continent from Congo (Brazzaville) to Ethiopia and south to Tanzania. The entire range of black-and-white colobus is more widespread, from Guinea Bissau in the west, including northeastern Angola and possibly Zambia to the south and Sudan and the Central African Republic to the north (Dandelot, 1971; Oates and Trocco, 1983; Napier, 1985).
Colobus angolensis Sclater, 1860
Taxonomic Notes: Rahm (1970) and Napier (1985) recognized eight subspecies. Dandelot (1971) recognized only six, synonymizing adolfifriederici with ruwenzorii and palliatus with sharpei. Groves (2001) recognized the same six subspecies as Dandelot (1971) and included an unnamed subspecies from the northern part of the Mahale mountains, east of Lake Tanganyika. This catalog followed Napier (1985) in maintaining the separate status of adolfifriederici and ruwenzorii and of palliatus and sharpei.
Pelage Notes: The long white shoulder hairs, which are continuous with the white hairs of the cheeks and throat, distinguish Colobus angolensis from other Colobus species. Some subspecies have a white brow band. There is no white mantle and the tail is untufted. The proximal portion of the tail is black and the distal portion is white or grizzled. The proportion of the grizzled part of the tail varies in length according to subspecies. Napier (1985) used features such as the brow band, epaulettes, pubic patch, and tail coloration to identify subspecies; that format is followed here.
Range of Specimens: The Colobus angolensis specimens in the AMNH were collected from areas across northern DRC, Uganda, and southwestern Tanzania. Colobus angolensis is distributed across mainland Africa from southern Congo (Brazzaville), northeastern Angola, DRC (excluding the northwest and southeast regions), Rwanda, Uganda, and Tanzania and is also found at the Kenyan coast (Dandelot, 1971; Napier, 1985; Groves, 2001).
Colobus angolensis adolfifriederici Matschie, 1914b
Taxonomic Notes: Dandelot (1971) considered adolfifriederici to be synonymous with ruwenzorii. He proposed that the only distinction between the two forms is in pelage density; adolfifriederici, living at higher altitudes than ruwenzorii, has a longer pelage. Hull (1979) also united adolfifriederici with ruwenzorii based on craniometric studies. Rahm (1970), however, did not agree, contesting that there are significant differences between the two in shoulder hairs and in tail coloration. Napier (1985) also noted distinctions in the pelage between the two forms, including less profuse epaulettes in adolfifriederici (mentioned by Rahm, 1970) and a broader white pubic patch in ruwenzorii. Groves (2001) united the two subspecies. Given the uncertainty, adolfifriederici and ruwenzorii are treated in the catalog as separate subspecies in line with following a conservative classification.
Pelage Notes: There is only one skin specimen of adolfifriederici in the collection. Narrow brow band, epaulettes less profuse than those of C. a. angolensis. Pubic patch small and grizzled. Tail almost entirely black, distal 15% grizzled.
Range of Specimens: Southern Uganda. The distribution of the subspecies is eastern DRC, Rwanda, and southwestern Uganda (Napier, 1985). Rahm (1970) extended the distribution to Minziro forest in Tanzania, which is just over the border with Uganda.
Specimens: Appendix 1, no. 1.
Colobus angolensis angolensis Sclater, 1860
Pelage Notes: Brow band absent, epaulettes long. Approximately 25% of the distal portion of the tail is grizzled.
Range of Specimens: Western and southern DRC. The subspecies is found in northeastern Angola and DRC (Rahm, 1970; Dandelot, 1971) from the left bank of the Congo and Lomani Rivers to the southern border of Katanga (Dandelot, 1971).
Specimens: Appendix 1, no. 2.
Colobus angolensis cottoni Lydekker, 1905
Pelage Notes: Brow band absent, epaulettes short, not very profuse, pubic region entirely black. Tail black at base, grizzled for the remaining 80%.
Range of Specimens: All of the specimens for which there are locality data available are from northeastern DRC, which is the range of the subspecies noted by Rahm (1970) and Dandelot (1971). The last two specimens in the list do not have locality data. However, Allen (1925) included them as C. a. cottoni, and the pelage markings of the two specimens are comparable to the rest of the collection.
Specimens: Appendix 1, no. 3.
Colobus angolensis ruwenzorii Thomas, 1901
Pelage Notes: There are two skin specimens of ruwenzorii in the museum collection, only one of which is complete. The following description is based on that skin. Long, dense pelage. Narrow brow band, long epaulettes, broad grizzled pubic patch. Tail almost all black, distal 5% grizzled.
Range of Specimens: Ruwenzori Mountains. This is the range of the subspecies according to Rahm (1970) and Napier (1985). Dandelot (1971) and Groves (2001) extended the range of ruwenzorii, as they have united it with adolfifriederici.
Specimens: Appendix 1, no. 4.
Colobus angolensis sharpei Thomas, 1902
Taxonomic Notes: Dandelot (1971) united sharpei with palliatus for the same reason he united adolfifriederici with ruwenzorii; he argued that increased pelage density in sharpei is the only distinction between the two. Groves (2001) also untied the two. Napier (1985), however, noted various other differences in the pelage of the two forms, and Hull (1979) suggested keeping sharpei and palliatus separate pending further study. The two are considered separate subspecies in this catalog.
Pelage Notes: Long pelage, distinct, broad white brow band. Long epaulettes, white, broad pubic patch. Distal 25% of tail grizzled.
Range of Specimens: Southern Tanzania. Rahm (1970) noted the distribution as north of Lake Malawi, and Rungwe, Poroto, Livingstone, and Uzungwa Mountains, Tanzania.
Specimens: Appendix 1, no. 5.
Colobus guereza Rüppell, 1835
Taxonomic Notes: Nine subspecies were recognized by Schwarz (1929) and Rahm (1970). Dandelot (1971) recognized only six, synonymizing percivali with guereza and uellensis and dodingae with occidentalis. Hull's study (1979) of craniometric characters led him to recognize eight subspecies, uniting uellensis with occidentalis. Groves (2001) and Napier (1985) concurred with this arrangement, and that classification is followed here.
Pelage Notes: The distinguishing features of the pelage of C. guereza are the long white mantle and the long flowing tail, which ends in a white tuft. This species is also noted for its wide white ring around the callosities. The features used by Dandelot (1971) and Napier (1985) to distinguish between subspecies are the length of the mantle, color of tail base, and description of the white tail tuft.
Range of Specimens: The Colobus guereza specimens in the AMNH were collected from localities in Congo (Brazzaville), northeastern DRC, Uganda, Kenya, and Ethiopia. Napier (1985) and Dandelot (1971) recorded the range of the species from eastern Nigeria through Cameroon, northern Gabon, Congo (Brazzaville), eastwards through northern DRC, Central African Republic, southern Sudan and Ethiopia, and southwards to Kenya, Uganda, and northern Tanzania.
Colobus guereza gallarum Neumann, 1902
Pelage Notes: Long mantle, covering tail base which is black, grizzled underneath. White tuft is almost half of the total tail length.
Range of Specimens: Eastern Ethiopia. Ethiopia, northeast of Rift Valley, Galla country (Dandelot, 1971).
Note: The colobus monkeys collected by T. D. Carter at Mount Ansha include what appear to be specimens of both C. g. gallarum and C. g. guereza. Napier (1985) and Dandelot (1971) distinguished the two subspecies by the color of their tail base. Although the tail base of the specimens of C. g. gallarum is somewhat grizzled underneath, it is distinguishable from the specimens of C. g. guereza. Yalden et al. (1977) expressed dissatisfaction with the basis of the separation, stating that the variation in tail color may be clinal in nature. Furthermore, it is not clear if the distribution or range of the two subspecies overlaps. Rahm (1970) included a map of the distribution of all the subspecies of Colobus. In this map, the distribution of C. g. gallarum does not extend as far south or west as Dandelot (1971) suggested. Yalden et al. (1977) stated that there appears to be no geographic boundaries between the two forms to separate them or to prevent dispersal.
Furthermore, the location of Mt. Ansha could not be determined exactly. It is north of Mt. Cacca (Kaka), and the province Carter (collector) called Arussi is now called Arsi (Yalden, personal commun.) While it is acknowledged that the same locality is noted for two subspecies, they were separated in the collection according to pelage differences.
Specimens: Appendix 1, no. 6.
Colobus guereza guereza Rüppell, 1835
Taxonomic Notes: Dandelot (1971) united percivali with guereza, while other authors (Rahm, 1970; Hull, 1979; Napier, 1985; Groves, 2001) maintained the two as separate subspecies.
Pelage Notes: Long mantle, covering base of tail, which is grizzled. White tuft comprises half of the tail.
Range of Specimens: Western Ethiopia. Groves (2001) reported the range from forested areas of the Ethiopian highlands, extending into lowland forests along the Omo River and in the Blue Nile gorge.
Specimens: Appendix 1, no. 7.
Colobus guereza kikuyuensis Lönnberg, 1912
Pelage Notes: Very long mantle, covering up to a quarter of the tail. Base of tail black, white tuft voluminous, forming about two-thirds of the length of the tail.
Range of Specimens: Southwestern Kenya. The subspecies occurs at Ngong Escarpment, Mount Kenya, and the Aberdare Range (Dandelot, 1971; Groves, 2001).
Specimens: Appendix 1, no. 8.
Colobus guereza matschiei Neumann, 1899
Pelage Notes: Long mantle, covering base of tail. Tail base black, tail tuft full, comprising about one-third of total length.
Range of Specimens: Eastern Uganda and western Kenya. Dandelot (1971) noted the range of the subspecies as Kenya, west of Rift Valley, Mau Escarpment, and Mount Elgon.
Specimens: Appendix 1, no. 9.
Colobus guereza occidentalis (Rochebrune, 1886–1887)
Taxonomic Notes: Rahm (1970) maintained occidentalis and uellensis as separate subspecies, while Dandelot (1971), Napier (1985), and Groves (2001) treated uellensis as a synonym of occidentalis. Hull's craniometric studies (1979) confirmed the view that the two are synonymous.
Pelage Notes: Short mantle, not extending beyond base of tail, which is black. There is variation among the specimens in the length of the tail tuft—from one-fifth to one-third of the total length.
Range of Specimens: There are three main collection localities for the specimens of occidentalis here: southern and western Uganda, northeastern DRC, and western Congo (Brazzaville). The subspecies is found across Africa from eastern Nigeria, southern Cameroon, Gabon, and northern DRC to Uganda and southwestern Sudan (Napier, 1985; Groves, 2001).
Specimens: Appendix 1, no. 10.
Colobus polykomos (Zimmermann, 1780)
Taxonomic Notes: Until recently, most authors (Rahm, 1970; Dandelot, 1971; Napier, 1985) recognized three subspecies (polykomos, dollmani, and vellerosus). Dandelot (1971), however, noted that vellerosus might actually be a distinct species. More recently, Oates and Trocco (1983) suggested raising C. polykomos vellerosus to specific status based on vocalization and other analyses. Oates et al. (1994) followed Oates and Trocco (1983) in recognizing vellerosus as a separate species. Booth (1958) and Rahm (1970) suggested that the intermediate form, dollmani, might in fact be a hybrid swarm, a theory investigated by Groves et al. (1993). They conclude that dollmani is in fact a hybrid and that polykomos and vellerosus should be recognized as distinct species. In this catalog, I follow Groves et al. (1993) and Groves (2001) in recognizing vellerosus and polykomos as separate species. There are no specimens labeled C. p. dollmani in the AMNH collection.
Pelage Notes: Gray brow band, gray long epaulettes. Tail white from base to tip and untufted. Callosities bordered by narrow ring of white hair.
Range of Specimens: Liberia and southern Ivory Coast. The range of the species is from Guinea to Sassandra River, Ivory Coast (Napier, 1985; Groves, 2001). Rahm (1970) included Gambia in the range. However, Oates and Trocco (1983) suggested that it is unlikely that C. polykomos is present in Gambia. Schwarz (1929) stated that the type of C. leucomeros was almost certainly not from “River Gambia”. Oates and Trooco (1983) were unaware of other specimens collected in Gambia and therefore excluded Gambia from the range of C. polykomos.
Specimens: Appendix 1, no. 11.
Colobus satanas Waterhouse, 1838
Taxonomic Notes: Dandelot (1971) and Eisentraut (1973) agreed that the subspecies metternichi was in fact based on partial albinos. Eisentraut (1973) also suggested that the mainland population may be subspecifically distinct from the island population on Bioko.
Range of Specimens: The specimens in this collection were taken from southwestern Cameroon and northwestern Gabon. The subspecies occurs in Bioko, southwestern Cameroon, Equatorial Guinea, Gabon, and northwestern Congo (Brazzaville) (Rahm, 1970; Carpaneto 1995).
Specimens: Appendix 1, no. 12.
Colobus vellerosus (I. Geoffroy, 1834)
Taxonomic Notes: As already noted, this species was previously considered a subspecies of C. polykomos, due, according to Dandelot (1971), “to a craze for simplification”.
Pelage Notes: The six skins in the AMNH collection are not complete and consist only of the trunk of the body up to the forearms and down to the thighs. The grizzled patches on the thighs, the distinguishing feature of this species, are visible on the skins.
Range of Specimens: There were no locality data for the specimens of vellerosus in this collection. The range of the subspecies is from Bandama River, Ivory Coast, to western Nigeria (Dandelot, 1971; Rahm, 1970; Groves, 2001).
Specimens: Appendix 1, no. 13.
GENUS PROCOLOBUS, SUBGENUS PILIOCOLOBUS ROCHEBRUNE, 1886–1887
Taxonomic Notes: The subgenus Piliocolobus of the genus Procolobus represents the red colobus monkey.
Pelage Notes: The red colobus have been described as large, slender monkeys with long, untufted tails (Napier, 1985). There is no sexual dimorphism in the coat coloration. The pelages of infants and adults differ somewhat in hue but not as markedly as in the black-and-white colobus. The amount of red coloration in the pelages of these monkeys varies considerably among subspecies. Some of the subspecies (e.g., P. b. tholloni) are strikingly red in color. Others exhibit little red color; rather, their pelts are drab brown, as seen in P. b. oustaleti. Napier (1985) has published useful descriptions and pelage notes on this group.
Range of Specimens: The range of the red colobus monkeys in the AMNH collection is somewhat limited, as only 7 of the approximate 14 subspecies are represented. The range extends from Liberia and Ivory Coast in the west, across DRC and Uganda, with a few specimens from Tana River, Kenya. Dandelot (1971) stated that the red colobus monkeys are to be found across the African continent from Senegal in the west to the island of Zanzibar in the east.
Procolobus (Piliocolobus) badius (Kerr, 1792)
Taxonomic Notes: This species may actually be comprised of as many as eight species (Dandelot, 1971).
Pelage Notes: All of the P. badius subspecies in this collection have a dark brown or black band across the brow extending to the ears. They are similar in body size and can be easily distinguished by their respective pelage colors which have variable amounts of black, brown, and red.
Range of Specimens: The seven subspecies of P. badius in this collection are representative of populations in north-central Liberia, western Ivory Coast, western DRC, northeastern DRC (the majority of the specimens was from this region), Uganda, and Tana River, Kenya. As noted for the subgenus Piliocolobus, the actual range of these monkeys extends across the African continent.
Procolobus (Piliocolobus) badius badius (Kerr, 1792)
Pelage Notes: Two rich colors are seen in this subspecies—black and maroon. Neither color lessens in intensity throughout the coat, as is often seen in the varying shades of brown observed in the pelages of other subspecies. Black head, maroon cheeks. Nape black with maroon shoulders. Back and rump black. Sides maroon. Upper arms black, lower arms and hands maroon. Thighs black, legs and feet maroon. Tail black from base to tip.
Range of Specimens: Liberia to Ivory Coast. The range of the subspecies is from Sierra Leone, eastern Guinea, to western Ivory Coast. Eastern limit of their range is thought to be the Bandama River (Dandelot, 1971; Groves, 2001).
Specimens: Appendix 1, no. 14.
Procolobus (Piliocolobus) badius bouvieri (Rochebrune, 1886–1887)
Taxonomic Notes: This subspecies was considered by Dandelot (1971), Delson et al. (1982), and Groves (2001) to be a subspecies of pennantii.
Pelage Notes: The skin specimen in the collection is not in good condition. The head, feet, and hands are missing. The shoulders are cream-colored, the midline of the back is dark red to brown. The rest of the pelage is bright red to orange.
Range of Specimens: The collectors of this specimen noted the location as Ouesso, Congo; the skin is more than likely a trade skin, so the exact locality cannot be determined. The distribution of the subspecies is northern Congo (Brazzaville). Rahm (1970) noted the distribution as including the Rivers Sangha and Likouala. Dandelot (1971) stated that the distribution may also include the confluence of the Congo and Oubangui Rivers.
Specimens: Appendix 1, no. 15.
Procolobus (Piliocolobus) badius ellioti (Dollman, 1909)
Taxonomic Notes: Three specimens collected by Lang and Chapin were classed as a new species, Colobus langi by Allen (1925). This name was later considered a synonym of ellioti. However, as mentioned previously, it is now thought that the species ellioti may actually be comprised of a number of distinct species and hybrids (Colyn, 1991). In fact, a comparison of Allen's langi specimens to the one considered by him to be C. ellioti (52286) may support this proposition. All five specimens in the collection are from the northeast of DRC. The collection localities on the Lang and Chapin map appear to be separated by rivers, although it is difficult to conclude from the map if real geographic boundaries exist. For the purposes of this catalog, all of the specimens are tentatively included in P. b. ellioti. Two distinct phenotypes can be seen in the collection. The two morphological characters used by Allen (1925) to distinguish the C. ellioti and C. langi specimens were body size (C. langi being smaller) and the abrupt transition in color observed on the back of C. langi. That transition can be seen in specimens 52290, 52291, 52292, and 119607. However, the latter is larger in body size than the first three specimens. The abrupt color transition is not evident in specimen 52286. It remains to be seen how the taxonomy of this group will be resolved.
Dandelot (1971) considered ellioti to be a potential species, while Delson et al. (1982) considered it a subspecies of pennantii. Groves (2001) included ellioti as a subspecies of foai.
Pelage Notes: Group 1: Specimen 52286. Auburn head and cheeks. Nape speckled lightly with black. Shoulders light auburn. Upper back deep auburn-brown, grading to brown at sides of mid-back and rump. No abrupt transition in back coloration. Sides brown, underparts cream. Forelimbs auburn, hands reddish-brown. Hind limbs light brown from thigh to foot, feet darker brown. Tail brown. Group 2: Specimen 119607. Auburn head and cheeks. Nape, shoulders, top of back rich auburn. Abrupt transition to dark brown at mid-back, grading into lighter brown at rump. Forelimbs and hands auburn. Hind limbs brown from thighs to feet, feet darker brown. Tail dark brown to tip. Specimens 52290, 52291, and 52292 are similar in coloration but are considerably smaller in size than specimen 119607. The major difference in coloration between these two groups is the transition between auburn and brown on the back. It is an obvious and distinguishing feature of the latter group.
Range of Specimens: All specimens of P. b. ellioti in this collection were collected from northeastern DRC. Procolobus b. ellioti is found in eastern DRC, the Ituri forest, and at least as far south as Lake Edward (Dandelot, 1971).
Specimens: Appendix 1, no. 16.
Procolobus (Piliocolobus) badius oustaleti (Trouessart, 1906)
Taxonomic Notes: Lang and Chapin collected all but one of these specimens. In his notes on this collection, Allen (1925) divided the specimens into two subspecies—Colobus powelli powelli and Colobus powelli brunneus. The main reason behind this separation was the variation in color. The specimens from the forest galleries of the savannah country were considered much paler and duller in tone (C. p. brunneus) than the more richly colored specimens from the rainforests (C. p. powelli). The author did however remark on the “extremely wide range of color variation” within the entire group. Almost every description of pelage in terms of sex, age, and locality has to be qualified by numerous exceptions. Both names (powelli and brunneus) were later considered synonyms of oustaleti. This subspecies itself is also considered to be a subspecies of rufomitratus (Dandelot, 1971), pennantii (Delson et al., 1982), or foai (Groves, 2001).
Pelage Notes: Considerable variation in pelage color is seen in the skins of this group. The amount of red in the pelage varies from its complete absence to auburn-infused brown. The following description is general. Head usually dark brown, lighter brown cheeks. Nape of neck dark brown, shoulders brown. Back coloration grades from dark brown to lighter at rump. Sides medium brown to reddish brown. Underparts light colored. Forelimbs brown, tinged with red, cream underneath. Hands and feet dark brown or black. Hind limbs medium brown to reddish brown. Tail dark brown. The overall impression of coat color in this group is one of nondescript, dull brown tones. The near absence of red coloration distinguishes the group from other subspecies. There is little evidence of color transition or of the rich hues seen in the other subspecies in this collection. There are three specimens of infant skins in the P. b. oustaleti collection. Two of the specimens (52274 and 52328) have no red coloration and are duller in tone than the adult skins. The third infant pelage (52318) has some red coloration appearing on the head.
Range of Specimens: Most of the specimens collected are from northeast DRC, in the vicinity of Faradje. The range of this subspecies extends from eastern Congo (Brazzaville) across DRC and as far east as Lake Albert and the Ituri River (Dandelot, 1971).
Specimens: Appendix 1, no. 17.
Procolobus (Piliocolobus) badius rufomitratus (Peters, 1879)
Taxonomic Notes: Many authors have considered rufomitratus a valid species, including Dandelot (1971), Delson et al. (1982), and Groves (1993, 2001).
Pelage Notes: Head capped with russet, light brown cheeks. Nape dark brown, shoulders light brown to creamy white. Back grades from dark to medium to light brown at rump. Sides light brown grading to creamy-white underparts. Forelimbs and hind limbs very light brown, almost creamy. Hands and feet light brown. Tail medium brown, darker at tip. The distinguishing characteristic here is the prominent red cap. The rest of the coat is brown, similar in tone to the brown of P. b. oustaleti.
Range of Specimens: Specimens were collected from the mouth of the Tana River, Kenya. Range of the subspecies is restricted to Tana River gallery forests from Garsen north nearly to Wenje (Groves, 2001).
Specimens: Appendix 1, no. 18.
Procolobus (Piliocolobus) badius tephrosceles (Elliot, 1907)
Taxonomic Notes: Dandelot (1971) considered tephrosceles to be a subspecies of rufomitratus, while Delson et al. (1982) included it as a subspecies of pennantii. Groves (2001) considered it a good species.
Range of Specimens: All the specimens are from the Kibale Forest, western Uganda. The range of the subspecies extends from western Uganda, Rwanda, Burundi, and northwestern Tanzania (Dandelot, 1971; Groves, 2001).
Specimens: Appendix 1, no. 19.
Procolobus (Piliocolobus) badius tholloni (Milne-Edwards, 1886)
Taxonomic Notes: Dandelot (1971) considered tholloni to be a valid species, while it was considered to be a subspecies of pennantii by Delson et al. (1982). Groves (2001) regarded it as a good species.
Pelage Notes: Rich auburn head, cheeks a lighter auburn. Nape red, speckled with black. Shoulders orange-red. Top half of back black to dark brown. Rest of back and sides rich auburn. All limbs rich auburn to wrists and ankles. Wrists, ankles, hands, and feet all black. Underparts creamy, infused with auburn. Tail deep auburn grading to red-brown at tip.
This subspecies is distinctly red in color. The red tail and the abrupt transition of red to black at the wrists and ankles are distinguishing features in the specimens in the AMNH collection. No other group in the collection has as much red in its pelage.
Range of Specimens: Western DRC, just south of the River Congo, on the border with Congo. Dandelot (1971) noted that this subspecies is found south of the Congo River; the southern limit may be the Rivers Kasai and Sankuru. Groves (2001) recorded the eastern limit as the River Lomami.
Specimens: Appendix 1, no. 20.
GENUS PROCOLOBUS, SUBGENUS PROCOLOBUS ROCHEBRUNE, 1886–1887
Procolobus (Procolobus) verus (van Beneden, 1838)
Taxonomic Notes: The subgenus Procolobus contains only one species, Procolobus verus, the olive colobus.
Pelage Notes: Coat has short hair and is a dull reddish olive color. Limbs and tail are olive. There is a longitudinal crest on the crown.
Range of Specimens: Liberia. The range extends from Sierra Leone to eastern Nigeria (Dandelot, 1971; Oates et al., 1994).
Specimens: Appendix 1, no. 21.
GAZETTEER
The exact location and geographic coordinates of as many localities as possible were determined. Information on localities was taken from a number of sources: The Times Atlas of the World (9th edition), collectors' field notes, maps and gazetteers in the Department of Mammalogy archives, and publications of other authors (e.g., Allen, 1925; Colyn, 1991).
Specimens were collected from three broad geographic regions in Africa. They are listed under those regions below. When available, the geographic coordinates are noted for the collection locality. A map of the localities in the Congo Basin is represented in figure 1.
West African Forest Block
Gabon
Kango, 00°15′N, 10°11′E (Times Atlas, 1992).
Ghana
Manso Wassaw. It is likely that the collection locality is between two towns, Manso and Wassaw. Manso is at 05°08′N, 01°50′W (Times Atlas, 1992). There is a town called Wassaw Dunwa at 05°46′N, 02°32′W (US Board on Geographic Names No. 102, 1967).
Ivory Coast
Cavally River, Georgetown, 04°36′N, 07°33′W (US Board on Geographic Names No. 89, 1965).
Cavally River, Niroyé. A map for the locations of Georgetown and Niroyé can be found in Tate (1942). Niroyé is found in the region of the Lower Cavally River, south of Taté, 04°48′N, 7°32′W (Times Atlas, 1992).
Duékoué, 06°40′N, 07°15′W (Times Atlas, 1992).
Liberia
Lofa, 11 km north, 3 km east of Ziggida, Gabayea River. Dickerman's field notes contain a map. Ziggida is at 08°02′N, 09°29′W (US Board on Geographic Names No. 106, 1968).
Newtown, 04°54′N, 08°26′W (US Board on Geographic Names No. 106, 1968).
Sawtown, 11 mi east of the Farmington River (letter from Tate to H.E. Anthony, correspondence file, AMNH). The Farmington River is located at 06°08′N, 10°22′W (US Board on Geographic Names No. 106, 1968).
Congo Basin
Cameroon
Efulan, 02°42′N, 10°30′E (Chapin, 1954), 53 miles east of Kribi at 02°56′N, 09°56′E (Times Atlas, 1992).
15 km west of Eséka, 03°40′N, 10°48′E (Times Atlas, 1992).
Mfume = Mfuni, 05°40′N, 09°16′E (US Board on Geographic Names No. 60, 1962).
Congo (Brazzaville)
Kelle Abole. Again, it is likely that the collection locality is between two towns, Kelle and Abole. A town called Kellé is located at 00°05′S, 14°33′E (Times Atlas, 1992). There is a town called Abolo at 00°08′N, 14°16′E (US Board on Geographic Names No. 61, 1962)
Makoua, 00°01′S, 15°40′E (Times Atlas, 1992).
Ouesso, 01°38′N, 16°03′E (Times Atlas, 1992).
Democratic Republic of the Congo
Abawe, 02°30′N, 26°50′E (Allen, 1925).
Akenge, 02°54′N, 26°49′E (Chapin, 1954).
Angumu, 00°10′S, 27°42′E (Colyn, 1991).
Avakubi, 01°18′N, 27°35′E (Colyn, 1991).
Bafuka, 04°20′N, 27°50′E (Allen, 1925).
Bafwabaka, 02°10′N, 27°39′E (Colyn, 1991).
Beni, 3,900 ft., 00°28′N, 29°28′E (Chapin, 1954).
Bolobo, 02°10′S, 16°17′E (Times Atlas, 1992).
Faradje, 03°44′N, 29°43′E (Colyn, 1991).
Gamangui, 02°10′N, 27°17′E (Colyn, 1991).
Kasai District, Kananga (formerly Luluabourg), 05°53′S, 22°26′E (Times Atlas, 1992).
Kivu, west base of Ruwenzori. This information probably refers to the district of Kivu. The Ruwenzori Mountains run from 00°05′N, 29°42′E to 00°53′N, 30°12′E (Chapin, 1954).
Lukolela, 01°10′S, 17°11′E (Times Atlas, 1992).
Lukolela, Plaine de Tervueren (the latter not found).
Medje, 02°25′N, 27°18′E (Colyn, 1991).
Mokoko (location from Chapin's field notes), “near Mokoko, 114 km from Irumu [01°31′N, 29°49′E (Chapin, 1954)] on road to Beni”, 00°28′N, 29°28′E (Chapin, 1954).
Niapu, 02°25′N, 26°28′E (Colyn, 1991).
Poko, 03°08′N, 26°58′E (Colyn, 1991).
Risimu, 00°31′N, 25°35′E (Chapin, 1954).
Rutshuru, 01°11′S, 29°27′E (Colyn, 1991).
Ukaturaka, 02°00′N, 20°30′E (Allen, 1925).
Vankerckhovenville, 03°17′N, 29°26′E (Chapin, 1954).
Yakuluku, 04°21′N, 28°49′E (Chapin, 1954).
Uganda
Kakuka area, NW Ruwenzori Mts., 9000 ft. This is probably Kakuka Mt., 00°35′N, 29°59′E (Colyn, 1991).
Karamoja Dist., Mt. Kadam, 01°45′N, 34°43′E (Times Atlas, 1992).
Karamoja Dist., Namalu, foot of Mt. Kadam. Namalu is at 01°49′N, 34°38′E (Delany, 1975).
Kibale Forest, Kanyawara, 5,000 ft, 00°34′N, 30°21′E (from collector's field tags).
Kibale Forest, near Dubona camp, 00°14′N, 30°22′E (from collector's field tags).
Malabigambo Forest, 00°57′S, 31°35′E (Colyn, 1991), Sango Bay area, 4,000 ft.
Mubende area, 4,000 ft. Mubende is at 00°35′N, 31°24′E (Times Atlas, 1992).
West Nile Dist., Aringa Co., junction of Koich and Kenga Rivers. A town called Koich is located at 02°35′N, 32°14′E (Times Atlas, 1992) and Koich stream is at 03°28′N, 31°00′E to 03°23′N, 31°30′E (Pohill, 1988). Kenga River could not be located.
West Nile Dist., Koboko Co., 03°24′N, 30°58′E (Times Atlas, 1992), Godea—not located.
West Nile Dist., Koboko Co., 03°24′N, 30°58′E (Times Atlas, 1992), Ludara—not located.
West Nile Dist., Koboko Co., 03°24′N, 30°58′E (Times Atlas, 1992), Utukiliri—not located.
Rift Valley System
Ethiopia
Arussi, Mt. Ansha. The district of Arussi is now called Arsi. Mount Ansha is located north of Mount Cacca (Kaka), 07°45′N, 39°15′E (Yalden et al., 1980), Arussi Mountains, 07°30′N, 39°20′E (Yalden et al., 1977). Kaka Mountains are marked on Carter's expedition map (no. 4520, AMNH map file); however, Mount Ansha is not.
Lake Stephanie, 04°26′N, 36°56′E to 05°00′N, 36°45′E (Chapin, 1954).
Kenya
Aberdare Range, 00°07′S, 36°33′E to 00°58′S, 36°39′E (Chapin, 1954).
Elgeyo Forest, 7,000 ft, 00°18′N, 35°33′E to 01°20′N, 35°38′E (Pohill, 1988).
Kijabe, 00°56′S, 36°34′E (Times Atlas, 1992).
Kikuyu. The village of Kikuyu is at 01°14′S, 36°40′E (Chapin, 1954).
Kikuyu Escarpment, 00°56′S, 36°35′E to 01°06′S, 36°35′E (Chapin, 1954), Theita (Thika) River, 00°44′S, 36°47′E to 00°54′S, 37°28′E (Chapin, 1954).
Nyeri Dist., South Laikipia Forest, 1 mi north of Bellevue, 7,900 ft. The coordinates for Bellevue are given in Pine's field notes as 00°12′S, 36°43′E.
Tana River, 00°42′S, 37°14′E to 02°33′S, 40°31′E (Chapin, 1954).
Uasin—Gishu District, 00°10′N, 34°50′E to 00°56′N, 35°32′E (Chapin, 1954).
Tanzania
Rungwe, 5,650 ft, 09°10′S, 33°35′E (Times Atlas, 1992).
Acknowledgments
I thank Darrin Lunde and Esteban Sarmiento for advise on this work throughout the process. I am grateful to John Oates and Christopher Norris for helpful suggestions and criticism of an early draft of the manuscript. Eric Stiner provided essential assistance with the map and my brother, Kevin O'Leary checked locality coordinates for the gazetteer. Finally, I thank Rob Voss and two anonymous reviewers for their valuable comments.
REFERENCES
Fig. 1.
Collection localities in the Congo Basin. ▪ represent localities where black-and-white colobus were collected, □ represent localities where red colobus were collected. Localities where both red, and black-and-white colobus were collected are marked with a ;rq. The localities shown on the map are as follows: Congo (Brazzaville): 1. Makoua, 2. Ouesso. Democratic Republic of the Congo: 3. Abawe, 4. Akenge, 5. Angumu, 6. Avakubi, 7. Bafuka, 8. Bafwabaka, 9. Beni, 10. Bolobo, 11. Faradje, 12. Gamangui, 13. Kananga, 14. Lukolela, 15. Medje, 16. Niapu, 17. Poko, 18. Risimu, 19. Rutshuru, 20. Ukaturaka, 21. Vankerckhovenville, 22. Yakuluku. Uganda: 23. Kibale Forest, Kanyawara, 24. Kibale Forest, near Dubona camp, 25. Malabigambo Forest
