Birth seasons of Cercopithecus, Cercocebus and Colobus in Zaire.
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The ecological-constraints model assumes that food items occur in depletable patches and proposes that an increase in group size leads to increased day range due to more rapid patch depletion. Smaller groups become advantageous when an increase in travel costs is not repaid by an increase in energy gained or some other fitness advantage. On the other hand, we also know that group size can be influenced by social factors. Here we contrast the diet and group size of red colobus (Procolobus badius) and black-and-white colobus (Colobus guereza) in Kibale National Park, Uganda to consider how ecological and social factors are affecting their group sizes. Subsequently, we examine whether the insights gained from this detailed comparison can provide an understanding of why the social organization and group size of mantled howlers (Alouatta palliata) and black howlers (A. pigra) differ. Two groups of red colobus and two groups of black-and-white colobus were studied over 10 months. Red colobus groups were larger (48 and 24) than black-and-white colobus groups (9 and 6). The two groups of red colobus overlap home ranges with the two groups of black-and-white colobus; 75% and 95% of their home ranges were within red colobus's home range. There was a great deal of similarity in the plant parts eaten by the two species and both species fed primarily on young leaves (red colobus 70%, black-and-white colobus 76%). In terms of the actual species consumed, again there was a great deal of similarity between species. The average dietary overlap among months for the two neighboring groups of red colobus was 37.3%, while the dietary overlap between the red colobus and the black-and-white colobus group that had its home range almost entirely within the home range of the red colobus groups averaged 43.2% among months. If ecological conditions were responsible for the difference in group size between the two colobine species, one would expect the density of food trees to be lower in the home ranges of the black-and-white colobus monkeys, since they have the smaller group size. We found the opposite to be true. Both black-and-white colobus groups had more food trees and the cumulative size of those trees was greater than those in the red colobus's home ranges. We quantify how these differences parallel differences in mantled and black howlers. The average group size for mantled howlers was 12.9 individuals, and for black howlers it was 5.3 individuals. We explore possible social constraints, such as infanticide, that prevent black-and-white colobus and black howlers from living in large groups.
Humans are responsible for massive changes to primate habitats, and one unanticipated consequence of these alterations may be changes in host-parasite interactions. Edges are a ubiquitous aspect of human disturbance to forest landscapes. Here we examine how changes associated with the creation of edges in Kibale National Park, Uganda, alter the parasite community that is supported by two species of African colobines: the endangered red colobus (Piliocolobus tephrosceles) and the black-and-white colobus (Colobus guereza). An analysis of 822 fecal samples from edge and forest interior groups revealed no difference in the richness of parasite communities (i.e., the number of parasite species recovered from the host's fecal sample). However, for both species the proportion of individuals with multiple infections was greater in edge than forest interior groups. The prevalence of specific parasites also varied between edge and forest interior groups. Oesophagostomum sp., a potentially deleterious parasite, was 7.4 times more prevalent in red colobus on the edge than in those in the forest interior, and Entamoeba coli was four times more prevalent in red colobus on the edge than in animals from the forest interior. Environmental contamination with parasites (measured as parasite eggs/gm feces) by red colobus from the edge and forest interior differed in a similar fashion to prevalence for red colobus, but it did not differ for black-and-white colobus. For example, egg counts of Oesophagostomum sp. were 10 times higher in red colobus from the edge than in those from the interior. The less severe infections in the black-and-white colobus relative to the red colobus may reflect the fact that black-and-white colobus raid agricultural crops while red colobus do not. This nutritional gain may facilitate a more effective immune response to parasites by the black-and-white colobus. The fact that animals on the edge are likely not nutritionally stressed raises an intriguing question as to what facilitates the elevated infections in edge animals. We speculate that interactions with humans may be linked to the observed patterns of infections, and hence that understanding the ecology of infectious diseases in nonhuman primates is of paramount importance for conservation and potentially for human-health planning.
Factors that influence proximity and the number and duration of contacts among individuals can influence parasite transmission among hosts, and thus parasite prevalence and species richness are expected to increase with increasing host density. To examine this prediction we took advantage of a unique situation. Following the clearing of a forest fragment that supported red colobus (Piliocolobus tephrosceles) and black-and-white colobus (Colobus guereza), the animals moved into a neighboring fragment that we had been monitoring for a number of years and for which we had described the primate parasite community. After the animals immigrated into the fragment, the colobus populations more than doubled and colobus density became almost twice that found in Kibale National Park, Uganda. Despite this increase in host density, the richness of the parasite community did not increase. However, in both colobus species the prevalence of Trichuris sp., the only commonly occurring gastrointestinal parasite, increased. Over the next 5 years the prevalence and intensity of infection of Trichuris sp. in red colobus declined and their population numbers slowly increased. In contrast, the prevalence and intensity of infection of Trichuris sp. increased in black-and-white colobus and remained high following the immigration, and their population size declined. While Trichuris sp. infections are typically asymptomatic, we consider it a possibility that they contributed to the decline of the black-and-white colobus, and that the red colobus may be serving as a reservoir for Trichuris, thereby increasing the infection risk for black-and-white colobus.
The size sequence of the molar teeth in three genera, including seven species, of the Cercopithecidae was examined on the basis of mesiodistal and buccolingual crown diameters, and rectangle measurements (mesiodistal d. X buccolingual d.). The determination of molar size order was estimated by the three methods of mean values, size sequences and reductive indices (M2/M1, M3/M1). The results obtained are summarized as follows: 1. The molar size sequence of the seven species. The pattern of sequence in the molars of the Cercopithecidae was divided into six types: (I) M1 less than M2 less than M3 or (M1M2) less than M3, (II) M1 less than (M2M3), (III) M1 less than M3 less than M2, (IV) (M1M2M3), (V) (M3M1) less than M2, and (VI) M3 less than M1 less than M2 or M3 less than (M1M2). 2. The upper molars. i) The most frequent pattern of mean values found was type II for all the species of the Macaca and Colobus. The two species of Cercopithecus showed predominantly the characteristics of types III, IV and V. ii) The molar size sequence individual data revealed that in the mesiodistal crown diameter types I and II were distributed frequently in the Macaca, types IV and V in the Colobus, and type VI in the Cercopithecus. The reduction of the third molar, therefore, seemed to be more pronounced in the Cercopithecus when compared with the other two genus, Macaca and Colobus, because type VI is the type with more size reduction in the third molar. In the case of the buccolingual crown diameters, the most common size sequences were found in types II and III for the Macaca and Colobus. In the Cercopithecus, however, type V exceeded other types in frequence. iii) The results of the reductive indices showed that the second molars in the Colobus were greatly reduced compared to the first, followed by those of the Cercopithecus and Macaca. In the third molar reduction, the Cercopithecus had a relatively small third molar compared to the first, following the Colobus and Macaca. 3. The lower molars. i) The pattern in the mean values were frequently found to be type I for the Macaca and Colobus, and types III and II for the Cercopithecus. ii) The size sequence in the frequency of the mesiodistal crown diameter was well presented in type I for the Macaca and Colobus, while types IV and V were distributed frequently for Cercopithecus in which the reduction of the third molar was noticeable.(ABSTRACT TRUNCATED AT 400 WORDS)
From March 1999 through August 2000, 511 stool samples collected from 11 different primate species in 10 geographically distinct locations in Kenya, East Africa, were screened for the presence of Cyclospora spp. oocysts. Positive samples (43/102, 42%) were identified in vervet monkeys (Cercopithecus aethiops) in 4 of 4 locations; 19/206 (9%) in yellow and olive baboons (Papio cynocephalus, P. anubis, respectively) in 5 of 5 locations; and 19/76 (25%) in black and white colobus monkeys (Colobus angolensis, C. guereza, respectively) from 2 of 3 locations. DNA sequences obtained from 18 S rRNA coding regions from respective subsets of these positive samples were typed as Cyclospora cercopitheci (samples from Cercopithecus aethiops). Cyclospora papionis (samples from Papio cynocephalus and P. anubis), and Cyclospora colobi (samples from Colobus angolensis and C. guereza). Cyclospora oocysts were not detected in samples collected from patas, highland sykes, lowland sykes, blue sykes, DeBrazza, or red-tailed monkeys. A coded map showing the geographic location of the collected samples is given. Stool samples from 1 troop of vervet monkeys were collected over a 12-mo period. Positive samples ranged between 21 and 63%. These results suggest that there is no strongly marked seasonality evident in Cyclospora infection in monkeys as has been noted in human infection. This is further confirmed by the recovery of positive samples collected from vervet monkeys, baboons, and colobus monkeys at all times of the year during this survey. This absence of seasonality in infection is especially notable because of the extreme weather patterns typical of Kenya, where marked rainy and dry seasons occur. A second noteworthy observation is that the striking host specificity of the Cyclospora species initially described was confirmed in this survey. Baboons were only infected with C. papionis, vervet monkeys with C. cercopitheci, and colobus monkeys with C. colobi, despite geographic overlaps of both the monkey and parasite species and wide geographic distribution of each parasite and monkey host.
Understanding the determinants of animal abundance has become more vital as ecologists are increasingly asked to apply their knowledge to the construction of informed management plans. However, there are few general models are available to explain variation in abundance. Some notable exceptions are studies of folivorous primates, in which the protein-to-fiber ratio of foods has been shown to predict biomass. Here we examine the generality of Milton's [American Naturalist 114:363-378, 1979] protein/fiber model by providing a detailed analysis of diet selection in black-and-white colobus monkeys (Colobus guereza), and applying the model to populations shown to be stable; an assumption not previously examined. Based on observations of two groups of black-and-white colobus in Kibale National Park, Uganda, and one group in a forest fragment, we documented that the animals selected young leaves that had more protein, were more digestible, and had a higher protein-to-fiber ratio than mature leaves. The mature leaves did not differ from young leaves with respect to secondary compounds or mineral content (with the exceptions of copper and zinc). All of the colobus groups selected foods with a high protein-to-fiber ratios. However, one group also selected more digestible foods, and in another group, foraging efforts were positively related to zinc and negatively related to potassium. Previous studies that examined Milton's protein/fiber model did not demonstrate that the study populations were stable. If some populations were not at carrying capacity, then the correlations drawn between food availability and/or quality and folivore biomass may have been spurious. To address this issue, we censused a series of forest fragments in 1995 and again in 2000. We found that the populations in these fragments had declined from 165 in 1995 to 119 animals in 2000. However, based on evidence of population stability and lack of forest disturbance, we concluded that five of the original populations were stable. The biomass of these populations was related to the protein-to-fiber ratio of the fragment's trees. Combining our data with published data, we demonstrate that the protein-to-fiber ratios of mature leaves available to these folivorous primates accounted for 87% of the variance in their biomass.