John F. Oates

DETERMINANTS OF VARIATION IN TROPICAL FOREST PRIMATE BIOMASS: NEW EVIDENCE FROM WEST AFRICA'

To explore sources of variation in tropical forest primate biomass, and, in particular, to test the hypothesis that soil conditions are a major ultimate determinant of the biomass of colobine monkeys and other primates, we compared data on the soils, vegetation, and primate community at a site in West Africa (Tiwai Island, Sierra Leone) with information from other sites, especially two other African sites (Douala-Edea in Cameroon, and Kibale Forest in Uganda). The biomass of eight anthropoid primate species in old secondary high forest on Tiwai was estimated from data on population densities assessed by transect samples combined with data on social group densities and individual body masses. Samples of soil and tree foliage were collected at the same site, and subjected to a variety of chemical and mechanical analyses. Our estimate of anthropoid biomass at Tiwai is 1229-1529 kg/kM2, including 682-889 kg/km2 of colobines. This is one of the highest primate biomasses recorded anywhere. The soils at Tiwai were found to be relatively high in sand content and low in pH, and to have low levels of mineral nutrients. Levels of condensed tannins in the mature foliage of the trees comprising a major part of the forest canopy were higher than at other sites, but the ratio of protein to fiber in this foliage was also higher than at any other site except Kibale. It is argued that a wide range of environmental factors affect primate population densities, and that nutrient-poor soils and high tannin levels in tree foliage do not necessarily produce a low primate (or colobine) biomass, as some earlier studies had suggested. Fur-

Assessment of the diversity of African primates

This account of the systematics of African primates is the consensus view of a group of authors who attended the Workshop of the IUCN/SSC Primate Specialist Group held at Orlando, Florida, in February 2000. We list all species and subspecies that we consider to be valid, together with a selected synonymy for all names that have been controversial in recent years or that have been considered to be valid by other authors in recent publications. For genera, species-groups or species, we tabulate and discuss different published systematic interpretations, with emphasis on more recent publications. We explain why we have adopted our taxonomic treatment and give particular attention to cases where more research is urgently required and in which systematic changes are most likely to be made. For all taxa, from suborder to subspecies, we provide English names.

Food selection by the South Indian leaf-monkey, Presbytis johnii, in relation to leaf chemistry

SummaryThe leaf-monkey Presbytis johnii has been found to exhibit considerable selectivity in its dietary utilization of mature foliage in a rain-forest habitat. To investigate the basis of this selectivity and to examine the hypothesis that the observed selection is related to the digestibility and toxicity of the available foliage, chemical analyses have been made on 16 of the most important tree species in the monkeys habitat. It has been found that the most heavily used items, which form a staple part of the diet of P. johnii, are characterized by a low fibre content and a very low condensed tannin content. However, neither class of compound is an absolute feeding deterrent as minor, but still significant, mature leaf food items contain considerable amounts of both. It is suggested that the feeding deterrents in these minor items, which may be ingested to supply specific dietary requirements, can perhaps be tolerated because of their dilution in the gut by the dietary staples. An examination of the pepsin/cellulase digestibility of available mature foliage showed that the staple food items tended to be highly digestible. Little correlation has been found between alkaloid content and food selection and it is suggested that the colobine forestomach microflora has the ability to detoxify at least some alkaloids. Additional data on mature leaf petioles and young leaves suggest that the high ratio of cell-sap to cell-wall in these items, and their related high digestibility, explain the high relative abundance of these items in the P. johnii diet.

A new west African chimpanzee subspecies

We have sequenced a portion of the mitochondrial DNA control region from the hairs of Nigerian chimpanzees living on both sides of the Niger River, which is proposed as a biogeographic boundary. Our data suggest that a previously unrecognized type of chimpanzee may be present in Nigeria and adjacent parts of Cameroon, and that the zoogeographic barriers separating chimpanzees are different from those previously proposed.

Taxonomy and phylogeny of black-and-white colobus monkeys. Inferences from an analysis of loud call variation.

Field recordings of male loud calls (or roars) from each major form of black-and-white colobus monkey have been analyzed spectrographically, and features of tempo and pitch measured. Considered together with data on cranial dimensions, coat pattern, and geographical distribution, the results of this analysis suggest that there are five species of black-and-white colobus: Colobus angolensis, C. guereza, C. polykomos, C. satanas, and C. vellerosus. C. guereza and C. vellerosus may have differentiated most recently during a major arid event prior to the last Pleistocene glacial maximum; they have an identical low-pitched roar which we consider to be a shared, derived character. The other species, of which C. satanas has the most distinct roar, may belong to older lineages.

The dangers of conservation by rural development – a case-study from the forests of Nigeria

The Okomu Forest Reserve in south-west Nigeria contains a 114-sq-km wildlife sanctuary that is an important refuge for several threatened species, including the white-throated guenon Cercopithecus erythrogaster. A conservation project that started in Okomu in 1987 focused initially on protection, but the emphasis recently shifted to a programme of agricultural development assistance to migrant farmers in the reserve. This approach, which appears to follow the philosophy espoused in IUCN/UNEP/WWFs Caring for the Earth, may hasten rather than prevent the destruction of this remnant tropical forest and its wildlife .

Patterns of Frugivory in Three West African Colobine Monkeys

We studied the comparative feeding ecology of three species of colobus (Procolobus badius, Procolobus verus, and Colobus polykomos) on Tiwai Island, Sierra Leone. We collected dietary data on each species by scan-sampling habituated groups. Because these groups were observed in the same study area during overlapping time periods, the confounding effects of temporal and spatial variability in food availability were reduced. Our results show that the annual diets of the two larger species (Procolobus badius and Colobus polykomos) include roughly equal proportions of fruits (including seeds), young leaf parts, and mature leaf parts, although P. badius had a greater intake of floral parts. Procolobus verus consumed almost no mature leaf parts, few fruits and seeds, and many young leaf parts. Colobus polykomos commonly fed from lianas. Seeds were the dominant fruit item eaten by all three colobus, and the fruits they selected were generally dull and non-fleshy, in contrast to the brightly-colored, pulpy fruits eaten by guenons. Leguminous plants contributed substantially to the diets of both the larger species, but comparisons with other African forest sites indicate that colobine biomass is not closely correlated with the abundance of leguminous trees in the forest.

New Genetic Evidence on the Evolution of Chimpanzee Populations and Implications for Taxonomy

Primatologists widely recognize chimpanzees as belonging to a single species, Pan troglodytes, which they traditionally have further divided into 3 subspecies: west African P. t. verus, central African P. t. troglodytes, and east African P. t. schweinfurthii. Previously, we suggested that the phylogeographic history of chimpanzees may be different from that implied by the widely used taxonomy of the species. We based the suggestion on only a limited sample of haplotypes from the first hypervariable region (HVRI) of mitochondrial (mt)DNA from chimpanzees in Nigeria. We have now compiled a more geographically comprehensive genetic database for chimpanzees, including samples obtained near the Niger and Sanaga Rivers. Our database is composed of 254 HVRI haplotypes from chimpanzees of known geographic origin, including 79 unique HVRI haplotypes from chimpanzees living in Nigeria and Cameroon. The genetic data provide clear evidence that a major phylogeographic break between chimpanzee lineages occurs near the Sanaga River in central Cameroon and suggest the need for a reclassification of chimpanzees.

African primates : status survey and conservation action plan

Effective habitat conservation is essential if wild populations of primates are to survive. This revised edition of the African Primate Action Plan (1986) provides a summary review of progress on projects originally recommended and gives recommendations for new projects in areas and on populations not previously included

The Bushmeat Trade in Southwestern Nigeria: A Case Study

Hunters and bushmeat retailers in Bendel State, Nigeria were interviewed from May through August 1982. Hunting was found to take place in all months of the year, but became more intense during the dry season. Twentyseven species of mammal were reported to be hunted, while 22 species of mammal and five species of reptile were found on sale. Retailers reported that the grasscutter (Thryonomys swinderianuswas the most popular species among their customers, followed by Maxwells duiker, the brushtailed porcupine, and the bushpig. The grasscutter was also the most widely sold species, followed by the giant rat, monkeys, and Maxwells duiker. Game animals were found to be shot and sold with little regard to existing laws, with the result that the larger mammals have become rare in Bendel and populations of the smaller ones have come under severe pressure. The bushmeat trade is highly commercialized, and substantial profits accrue to middlemen. Recommendations are made for the conservation of a natural resource, wildlife, that not only provides a significant amount of animal protein but also supports a large rural economy.