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The Taxonomic Diversity of the Colobinae of Africa

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The Taxonomic Diversity of the Colobinae of Africa JASs Invited Reviews Journal of Anthropological Sciences Vol. 85 (2007), pp. 7-34 The taxonomic diversity of the Colobinae of Africa Colin P. Groves School of Archaeology and Anthropology, Australian National University, Canberra, ACT 0200, Australia e-mail: [email protected] Summary - Th e colobine monkeys of Africa are much more diverse than is often realized; in the case of Red Colobus (genus Piliocolobus), this is because the species replace each other geographically, and there is even some interbreeding where their ranges meet. It is not possible to make a modern taxonomic revision of Black-and-white Colobus (genus Colobus), in the absence of a thorough modern study to determine consistency of the diff erences between the taxa deemed to be subspecies of the two polytypic species, C. angolensis and C. guereza. Red Colobus, in particular, are by their behaviour extremely vulnerable, and one species is presumed extinct, and another has not been seen for some years. Keywords - Colobus, Taxonomy, Biogeography, Piliocolobus, Procolobus, Africa, Cercopithecoidea. Introduction the mouth of the Saccus, but its exact function is unknown. Liquid ingesta, especially fruit Th e Old World monkeys are divided into two juices, bypass the fermentation chamber, going subfamilies: Cercopithecinae and Colobinae. Th e directly via a groove (which is closed off into a Colobinae are commonly known as ‘leaf eaters’, tube by longitudinal muscle bands) into the Pars but this is too sweeping: while it is true that they Pylorica. Th e colobine multipartite stomach are capable of subsisting on leaves to a much has been likened to the ruminant stomach, greater degree than probably any other primate, although structurally and probably functionally detailed fi eld study has shown that many species the stomach of Macropodidae (kangaroos and are in fact specialized seed eaters as much as leaf wallabies) is probably a better comparison. In all eaters. Th e stomach has been described by Kuhn three cases, however, the system enables the animal (1964). In all of the Colobinae it consists of to extract much more energy from its food than three (or four) parts. Th e most conspicuous part can other herbivores, by making complete use is the Saccus Gastricus, an enormous blind sac of the structural carbohydrates in cellulose and in which symbiotic bacteria attack (and possibly hemicellulose. Th is is the secret to ‘leaf eating’. detoxify) solid ingesta, fermenting cellulose Th e stomach, when full, extends the belly and hemicellulose, releasing short-chain fatty and accounts for between 10.8 and 24.9% of acids which are directly absorbed into the the body weight in records of 14 specimens bloodstream. Th e bacterial pap, now enriched (10 Colobus guereza, three C. angolensis, in protein, moves into the next part, the Tubus one Piliocolobus tephrosceles) in the Kenya Gastricus, where pH gradually drops until the National Museum, Nairobi. Th e molar and fi nal part, the Pars Pylorica, which corresponds premolar teeth of Colobinae are distinguished to the simple stomach of other primates and from those of Cercopithecinae by their high where proper digestion begins. Some genera, crowns and occlusal relief, and the shortened including Procolobus and Piliocolobus, have a trigonids. Skeletally, they are characterised by fourth stomach compartment, the Praesaccus, at thumbs being short or absent, and somewhat the JASs is published by the Istituto Italiano di Antropologia www.isita-org.com 8 African colobines shortened second toes on the foot. Strasser Systematic account (1994) detailed other specialisations of the foot morphology. In the skull, mandibular Colobus. Black-and-White Colobus. corpus deepens posteriorly, and generally the Diagnosis: Skull prognathous but supraorbital interorbital pillar is very broad, although in two torus not strongly developed; sagittal crest hardly Asian genera, Nasalis and Simias, it is narrow as ever developed; no suborbital fossa; borders of in the Cercopithecinae. Th e African genera of pyriform aperture rounded; no anterior nasal the Colobinae (which are all known popularly spine; posterior palatal canals not in deep fossae; as colobus monkeys) can be distinguished from choanae and pterygoid fossa narrow, basisphenoid the Asian genera (which are broadly known as crested, laterally guttered. langurs) by lacking a thumb (Fig. 1). In some In mandible, digastric fossae poorly imprinted; individuals, a tiny nubbin of a thumb is present, lower margin of jaw curved upward especially at but there is never more than a rudimentary anterior end; ascending ramus upright; coronoid proximal phalanx. In the past, all colobus process hooked backward; no symphyseal foramen. were commonly put together in a single genus, Maxillary incisors elongated, narrow, with Colobus; Verheyen (1962) retained Black-and- no lingual cingulum or tubercle; tips of lateral white and Red Colobus together in this genus pair point mediosagittally. Postcanine dentition but split off Olive Colobus into a separate genus heavy. Th e larynx is enlarged, measuring in a male Procolobus; Kuhn (1967), followed by Grubb et C. vellerosus 59mm long by 36mm dorsoventrally al. (2003), transferred Red Colobus to Procolobus, and 34mm wide (Hill & Booth, 1957). External but kept them apart from Olive Colobus in a nose convex, its septum extending onto upper lip. separate subgenus, Piliocolobus; while Groves Ischial callosities in male fused across midline. (2001) recognised all three as separate genera. No perineal swellings. Fur black, with white areas I continue to regard all three as separate genera. in four out of the fi ve species; infant coat white. Fig. 1 - African colobines, such as this Colobus polykomos, lack thumbs (photo S. Gippoliti). C.P. Groves 9 Procolobus. Olive Colobus. populations would or would not interbreed Diagnosis: Skull not very prognathous, were their ranges to meet, or according to one’s premaxillae vertical; supraorbital arcades thin, hypothesis of interrelationships: we should curved; sagittal crest develops in adult males; classify species according to how things are suborbital fossa present; pyriform aperture with observed to be, given the available evidence, not sharp margin, often an incipient anterior nasal spine; according to how we think things ought to be! posterior palatal canals in deep fossae; choanae Th ere are two drawbacks to this way of viewing and pterygoid fossa broad, basisphenoid fl at. In species, both of them illustrated by colobus mandible, digastric fossae deep, well separated; taxonomy. First is that diff erent species may upper and lower margins of mandible straight, indeed interbreed where their ranges meet, and parallel; ascending ramus slopes back; coronoid even form an inextricable hybrid swarm. We process rounded; a labial symphyseal foramen. just have to acknowledge this as a fact of nature, Maxillary incisors relatively broad, low, and we must treat hybrid swarms as valuable with a lingual cingulum and tubercle; lateral populations for conservation purposes in their incisors somewhat caniniform, their points own right, as illustrating the operation of directed downward. Postcanine dentition not as natural processes. Th e second drawback is that massive as Colobus, but M3 with a tuberculum the state of research in diff erent genera may be sextum. Th e larynx is not enlarged: 20 x 12 x at very diff erent stages of understanding; this is 14 mm (Hill & Booth, 1957). External nose the case for the African Colobinae. I have myself simple. Ischial callosities separate in both carefully studied available specimens of Red sexes. Perineal swellings developed periodically Colobus in European and some other museums, in adult females and temporarily in subadult and feel confi dent that I have identifi ed the males. Pelage predominately phaeomelanic; units, whereas such study as I have made on infant coat not strikingly diff erent from adult's. Black-and-white Colobus is incomplete (with the exception of the West African taxa), and Piliocolobus. Red Colobus. I have no option but to accept the revisions Skull generally prognathous, with good made, using diff erent species concepts, by supraorbital development; supraorbital arcades Colyn (1991) and by David Hull (personal straight, robust; sagittal crest develops in adult communication). If this results in a rather males; suborbital fossa present; borders of unbalanced treatment, it is to be regretted but I pyriform aperture rounded; no anterior nasal feel I have no option, as I have no evidence as to spine; posterior palatal canals not in deep fossae; what diagnosable units (phylogenetic species) choanae and pterygoid fossa narrow, basisphenoid might exist within Colobus angolensis or Colobus crested, laterally guttered. Mandible more as in guereza as presently recognised. Procolobus but with no symphyseal foramen. Dentition more as Colobus but with no incisor cingulum or tubercle; no tuberculum sextum. Genus COLOBUS Larynx is barely larger than in Procolobus: Illiger, 1811. 24 x 16 x 15mm (Hill & Booth, 1957). External characters in general as in Procolobus. Colobus Illiger, 1811, Prodr. Syst. Mamm. Avium, 69. As for species, I employ the Phylogenetic Colobolus Gray, 1821, London Med.Repos.15: Species Concept (Cracraft, 1983), as argued 298. in detail in Groves (2001). In this view, species Guereza Gray, 1870, Cat. Monkeys, Lemurs & are units essentially diagnosed by the possession Fruit-eating Bats, Brit.Mus.,V and 19. of fi xed heritable diff erences from other such Stachycolobus Rochebrune, 1887,
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