Niche overlap and polyspecific associations among sympatric cercopithecids in the campo animal reserve,Southwestern Cameroon

The community structure and polyspecific associations found in five sympatric resident primates (Cercopithecus nictitans, C. cephus, C. pogonias, Cercocebus torquatus, andC. albigena) and a temporary visitor (Mandrillus sphinx) were studied within the Campo Animal Reserve, a tropical rain forest in southwestern Cameroon. Several ecological variables of the six species, e.g. the number of foraging groups in the study area, group size, home range size, biomass density, and individual density, were estimated by undertaking a systematic census and by other means, e.g. tracing.C. nictitans was the most abundant species in terms of its foraging groups, biomass, and individual densities. The foraging biomass varied significantly among the six species. The five resident species frequently formed polyspecific groups.M. sphinx also associated with them when it visited the area. The food items fed on by the five resident species were also checked. An index for synecological analysis revealed that the five species shared similar food niches. The five species also utilized similar foraging areas and strata, possibly depending on the availabilities of common foods. The above results cannot be explained by conventional equilibrium-competition models. For understanding the polyspecific associations, it is proposed that the primates form polyspecific foraging groups in order to optimize their foraging biomass. This idea leads to a model that can also explain certain of the grouping behaviors, such as group fission-fusion, found in large body size primates.     

Census,habitat preference,and polyspecific associations of six monkeys in the Lomako Forest,Zaire

This paper reports preliminary data on a number of poorly known cercopithecids inhabiting the Lomako Forest, Zaire. Data include those on density, biomass, group size, the relationship between group type (mono vs. polyspecific) and forest type (mixed primary, secondary, monodominant primary, and swamp), and vertical stratification. Species examined are Colobus angolensis, Cercocebus aterrimus, Cercopithecus wolfi, Cercopithecus ascanius, Cercopithecus neglectus, and Allenopithecus nigroviridis. The density and biomass for the Lomako Forest anthropoids (excluding Cercopithecus neglectus and A. nigroviridis) are 165 ind/km² and 1,034 kg/km², respectively. Cercopithecus wolfi and Colobus angolensis are found in group sizes similar to those reported from other sites while Cercopithecus ascanius, Cercocebus aterrimus, and A. nigroviridis display group sizes unique to the Lomako. While Colobus angolensis, Cercocebus aterrimus, Cercopithecus ascanius and Cercopithecus wolfi were found, to varying extents, in all four forest types, Cercopithecus neglectus and A. nigroviridis inhabited only swamp forest. The different species sort out along vertical continua both within and between the four forest types. © 1994 Wiley-Liss, Inc.     

Associations between the drill (Mandrillus leucophaeus) and sympatric monkeys in Korup National Park,Cameroon

Studies of polyspecific associations among African forest primates have primarily focused on arboreal Cercopithecus and Procolobus/Colobus species. We examined the association frequency of the terrestrial drill (Mandrillus leucophaeus) with six sympatric monkey species in Korup National Park, Cameroon, testing reports that Mandrillus associations are infrequent and transient. We conducted 3,284 km of trail walks for 12 months (February–June 2006; July 2007 to January 2008), recording species composition in 612 primate clusters. Using a Markov chain Monte Carlo test, we compared the observed frequency of dyadic associations against null models of “no association.” A novel conservative statistical approach which addresses possible dependence of observations close in time was also used, further strengthening confidence in our findings. Drills associated with all monkeys throughout the study period, and were with at least one other species (range 1–5) in half of the encounters. The association frequency of drills with red‐capped mangabeys (Cercocebus torquatus) was greater than expected by chance, which is interesting given the morphological adaptation of the Mandrillus‐Cercocebus clade for the exploitation of the same dietary niche, hard seeds. The difference we observed in the use of forest strata by drills and mangabeys may reflect a strategy to reduce food competition while in association. The nature and duration of observed drill associations varied. Although some associations seemed to be chance encounters, others lasted for hours with the involved species foraging together. Am. J. Primatol. 73:127–134, 2011. © 2010 Wiley‐Liss, Inc.     

Note on drill (Mandrillus leucophaeus) ecology and conservation status in Korup National Park, Southwest Cameroon

We report preliminary findings on drill population ecology, feeding ecology, primate associations and conservation status in Korup National Park, Cameroon, based on analysis of data collected during 1,346 km (620 field hours) of trail patrols from February to June 2006. We encountered drills on 25 occasions and collected 304 fecal samples. Group size was estimated on four occasions (> or =25, > or =40, > or =67, > or =77), and multiple males were heard emitting the characteristic two-phase grunt of mature male drills in two of these groups. We saw a solitary male drill once. Examination of fecal samples indicated a seasonally variable diet consisting mainly of seeds, fruit pith, leaves and insects (especially ants and termites). Irvingia gabonensis and Musanga cecropioides fruits showed the highest percentage of occurrence in May (96%) and March (75%) samples, respectively, and could be a major food source for drills at that time. Drills were in association with at least one additional primate species during 57.9% of observations. These associations involved all of the diurnal primates found in Korup (Cercopithecus mona, Cercopithecus nictitans, Cercopithecus erythrotis, Cercopithecus pogonias, Piliocolobus preussi and Cercocebus torquatus) except chimpanzees (Pan troglodytes). We discuss our findings in terms of existing knowledge about drill ecology and highlight the urgent need for the protection of the significant drill population in Korup National Park.     

Cheek pouch use in relation to interspecific competition and predator risk for three guenon monkeys (Cercopithecus spp.)

Forest guenons (Cercopithecus spp.) are often found in polyspecific associations that may decrease predator risk while increasing interspecific competition for food. Cheek pouch use may mitigate interspecific competition and predator risk by reducing the time spent in areas of high competition/predator risk. I investigated these ideas in three forest guenons: Campbell’s monkey (Cercopithecus campbelli), spot-nosed monkey (C. petaurista), and Diana monkey (C. diana). I present 13 months of scan sample data from Taï Forest, Côte d’Ivoire, including 3,675, 3,330, and 5,689 records of cheek pouch distention, to quantify cheek pouch use, for Campbell’s, spot-nosed, and Diana monkeys, respectively. Cheek pouches are often used to hold fruit, so I first predicted that the most frugivorous species, Diana monkeys, would have the most cheek pouch distension. Spot-nosed monkeys ate the least amount of fruit over the study period and had the least distended cheek pouches, suggesting the importance of frugivory in relation to cheek pouch distension for this species. This was not a sufficient explanation for Campbell’s monkeys; Campbell’s ate fruit less than Diana monkeys, but had more distended cheek pouches, suggesting that cheek pouch use was not simply a reflection of high frugivory. From the interspecific competition hypothesis, I predicted that Campbell’s monkeys would have more distended cheek pouches than Diana and spot-nosed monkeys, and more distended cheek pouches when associated with Diana because Campbell’s monkeys have the highest potential for interspecific competition with dominant Diana monkeys. From the predator risk hypothesis, I predicted that Campbell’s would have more distended cheek pouches when not associated with highly vigilant Diana monkeys. Campbell’s monkeys had the most distended cheek pouches overall, but had more distended cheek pouches when not in association with Diana, suggesting the greater importance of predator risk rather than interspecific competition in Campbell’s cheek pouch use.     

Mixed-species primate groups in the kibale forest: Ecological constraints on association

Five species of diurnal primates in the Kibale Forest of western Uganda— red colobus (Colobus badius),black- and- white colobus (Colobus guereza),redtail monkeys (Cercopithecus ascanius),blue monkeys (Cercopithecus mitis),mangabeys (Cercocebus albigena)-often associate in mixed- species groups that vary in size and composition from day to day. Across this range of species, we found no consistent effect of association on feeding rate. In addition, there is no systematic difference between the species- specific individual feeding rates when animals were in mixed- species groups feeding in a specific tree on one day and when individuals of one of the same species were feeding in the same individual tree on a subsequent day. If associating in a mixed- species group lowers the risk of predation, one might expect that the number of vigilant events would decrease in mixed- species groups. However, the only species to exhibit a consistent decrease in vigilant behavior when in association was the red colobus. Redtail monkeys were more vigilant when in association. We predicted that the density and distribution of food resources would both constrain the frequency of association and the size of mixed- species groups. Based on 22 months of data on food resources and bimonthly censuses, we found no relationship between the frequency of association (except mangabeys) or mean mixed- species group size and the density and distribution of food resources for all species. Finally, we examined the behavior of the monkeys in and out of association before and after the playback of a crowned hawk eagle call (Spizaetus coronatus),a known predator. When more species were in association, the amount of time they spent being vigilant following the playback was greater and the response more intense than when fewer species were in association or when the group was alone. The results of this study illustrate that the nature of the costs and benefits of polyspecific associations for these different monkey species are complex and vary from species to species.     

Ecological Partitioning of Cercopithecus campbelli, C. petaurista, and C. diana in the Taï Forest

I determined the degree of ecological partitioning among 3 species of guenons (Cercopithecus campbelli, C. petaurista, and C. diana) in the Taï Forest, Côte d’Ivoire and used the partitioning data to understand competitive relationships among them. Over a 13-mo period, I measured ecological partitioning in terms of food and canopy stratum use for 2 habituated groups of each guenon species and also collected data on food availability. I found that the study species diverged primarily in food items consumed and vertical strata occupied. Cercopithecus petaurista ate much more foliage than the other species did and used mostly the middle strata (5–20 m). Cercopithecus diana ate primarily fruit and used mostly the upper strata (>20 m). Cercopithecus campbelli ate mostly fruit together with large amounts of animal matter and primarily occupied the ground and low strata (<5 m). Of the specific pairs, the diets of Cercopithecus campbelli/C. diana overlapped the most overall and decreased during the season of low fruit availability. Cercopithecus campbelli and C. diana age/sex classes also overlapped more than the age/sex classes of other species pairs. The results suggest that the potential for competition was more intense for Cercopithecus campbelli/C.diana relations than it is for other species pairs. I compare my results from Taï with those from other primate and guenon communities and demonstrate that dietary overlaps and seasonal dietary divergence are lower in Taï than in most other guenon communities.     

Primate Community Structure in Guyana: A Biogeographic Analysis

Studies of primate community structure increase our understanding of behavior, adaptation, and evolution. However, there are few biogeographic data on specific composition and association patterns in primate communities. I conducted a biogeographic analysis of the community structure of primate species at 16 sites in Guyana. I used data from 1725 km of line-transect censuses to determine specific composition and association patterns of 220 primate groups . Of the 18 polyspecific groups, 94.1% (N = 16) included squirrel monkeys. There was an overall trend towards positive specific associations among Guyanese primates. The only species that exhibited a negative pattern of interspecific associations were brown and wedge-capped capuchins. The sighting rate for wedge-capped capuchins at sympatric sites was particularly depressed compared to that for brown capuchins. Low plant productivity in Guyanese forests may reduce the diversity of feeding niches and result in a low incidence of polyspecific associations and scramble competition between wedge-capped and brown capuchins.     

Characterization of animal communities involved in seed dispersal and predation of Guibourtia tessmannii (Harms) J.Léonard,a species newly listed on Appendix II of CITES

Characterization of the ecology of endangered timber species is a crucial step in any forest management strategy. In this study, we described the animal communities involved in seed dispersal and predation of a high‐value timber species Guibourtia tessmannii (Fabaceae; Detarioideae), which is newly listed on Appendix II of CITES. We compared the animal communities between two forest sites (Bambidie in Gabon and Ma'an in Cameroon). A total of 101 hr of direct observations and 355 days of camera trapping revealed that a primate (Cercopithecus nictitans nictitans) and a hornbill (Ceratogymna atrata) were important seed dispersers in Gabon. Conversely, a greater presence of a rodent (Cricetomys emini), which could act both as predator and disperser, was observed in Cameroon. This study suggests that animal communities involved in seed dispersal of G. tessmannii may vary depending on environmental conditions and anthropogenic impacts. However, further studies are needed to properly identify the factors involved in seed dispersal and predation of G. tessmannii.     

Despotic wild patas monkeys (Erythrocebus patas) in Kala Maloue, Cameroon

The socio-ecological model predicts that the quality, distribution, and patch size of food resources determines the dominance hierarchy of female monkeys based on the type of food competition they experience. Comparative studies of closely related species have evaluated the socio-ecological model and confirmed its validity. For example, female patas monkeys in Laikipia, Kenya, form a nonlinear and unstable dominance hierarchy (i.e., egalitarian), whereas females of sympatric, closely related savannah monkeys form a linear and stable dominance hierarchy (i.e., despotic), in accordance with the model's predictions of the characteristics of food resources. I compared agonistic interactions involving food between patas monkeys (Erythrocebus patas) and sympatric savannah monkeys (Cercopithecus aethiops) in Kala Maloue, Cameroon. I found linear dominance hierarchies not only in savannah monkeys, but also in patas monkeys in Kala Maloue. The rates of agonistic interactions during feeding between patas monkeys were equivalent to those between savannah monkeys in Kala Maloue; further, these rates were significantly higher than those of both Laikipia patas and savannah monkeys. The results imply that patas monkeys in Kala Maloue are not egalitarian, but are despotic, similar to savannah monkeys. Disparity in the dominance hierarchies of patas monkeys between Kala Maloue and Laikipia were attributable to the differences in the characteristics of food resources. Although patas monkeys in Laikipia subsist on small and dispersed food resources within a high-density area, those in Kala Maloue subsisted on food resources that were clumped in intermediate-sized patches within a low-density area. This study shows that the socio-ecological model is applicable not only for interspecific comparisons but also for intraspecific comparisons.     

A new species of guenon (genus Cercopithecus) from Gabon

A new species of African forest monkey has been discovered in the lowland tropical rainforest of central Gabon. The previously undescribed monkey is a guenon, a member of the widespread African genus Cercopithecus. and belongs to the Cercopithecus lhoesti superspecies, whose two previously recognized species live in widely separated populations, C. preussi in Cameroon, and C. lhoesti east of the Zaire basin. The new guenon occurs as a small isolated population separated by 600 km and 1600 km, respectively, from the other two. Distinctive external characteristics are its white throat-ruff, extensive chestnut-orange saddle, and yellow-orange tip to the tail. Chromosomal analysis confirms its close phylogenetic affinities with C. preussi and C. Ihoesti, but reveals a unique karyotype. The guenon is described as a new species, Cercopithecus solatus, the sun-tailed monkey.     

Influence of Chimpanzee Predation on Associations Between Red Colobus and Red-tailed Monkeys at Ngogo,Kibale National Park,Uganda

Colobines often associate with cercopithecines at various African sites. Such polyspecific associations presumably have an antipredation function. At Ngogo, Kibale National Park, Uganda, red colobus (Procolobus rufomitratus) spend considerable time in association with red-tailed monkeys (Cercopithecus ascanius), and they are also heavily hunted by chimpanzees (Pan troglodytes). I conducted behavioral observations and playback experiments to test the hypothesis that red colobus and red-tailed monkeys obtain mutual protection and predator-related benefits by associating. Despite high chimpanzee hunting pressure on red colobus and much lower hunting pressure on red-tailed monkeys, red-tailed monkeys initiate, maintain, and terminate the associations. The results suggest that rather than providing red colobus with protection against chimpanzees, the associations occur mostly because they protect red-tailed monkeys against predation by eagles.     

Intrasexual competition and body weight dimorphism in anthropoid primates

Body weight dimorphism in anthropoid primates has been thought to be a consequence of sexual selection resulting from male-male competition for access to mates. However, while monogamous anthropoids show low degrees of weight dimorphism, as predicted by the sexual selection hypothesis, polygynous anthropoids show high variation in weight dimorphism that is not associated with measures of mating system or sex ratio. This observation has led many to debate the role of other factors such as dietary constraints, predation pressure, substrate constraints, allometric effects, and phylogeny in the evolution of anthropoid weight dimorphism. Here, we re-evaluate variation in adult body weight dimorphism in anthropoids, testing the sexual selection hypothesis using categorical estimates of the degree of male-male intrasexual competition (“competition levels”). We also test the hypotheses that interspecific variation in body weight dimorphism is associated with female body weight and categorical estimates of diet, substrate use, and phylogeny. Weight dimorphism is strongly associated with competition levels, corroborating the sexual selection hypothesis. Weight dimorphism is positively correlated with increasing female body weight, but evidence suggests that the correlation reflects an interaction between overall size and behavior. Arboreal species are, on average, less dimorphic than terrestrial species, while more frugivorous species tend to be more dimorphic than folivorous or insectivorous species. Several alternative hypotheses can explain these latter results. Weight dimorphism is correlated with taxonomy, but so too are competition levels. We suggest that most taxonomic correlations of weight dimorphism represent “phylogenetic niche conservatism”; however, colobines show consistently low degrees of weight dimorphism for reasons that are not clear. Am J Phys Anthropol 103:37–68, 1997. © 1997 Wiley-Liss, Inc.     

Sexual dimorphism,mating system,and effect of phylogeny in De Brazza's monkey (Cercopithecus neglectus)

De Brazza's monkey (Cercopithecus neglectus), like other guenons, shows marked sexual dimorphism in an array of features. While strong sexual dimorphism is generally associated with a polygynous mating system, populations of De Brazza's monkeys in Gabon are reportedly monogamous. An explanation of this unique phenomenon is offered here. Patterns of sexual dimorphism are examined for morphology, growth and development, behavior, and ecology, and field and captive studies on the social organization and mating system of De Brazza's monkey and congeneric guenon species are reviewed. Based on the findings, it is postulated that 1) De Brazza's monkeys are not strictly monogamous, but exhibit interpopulational variation in their mating system, from facultative monogamy to mild polygyny; 2) marked sexual dimorphism most likely reflects the effect of the historical-phylogenetic factor; ie, it represents a holdover of a degree of dimorphism established earlier in evolutionary history when the degree of polygyny Was higher; and 3) lessening in the degree of polygyny and a tendency toward monogamy represents a consequence of selection toward small group size. Small group size, a unique antipredator strategy, and failure to form polyspecific associations are ultimately most likely the result of intragroup and interspecific competition and predation pressure.     

Cooperation and competition in two forest monkeys

Putty-nosed monkeys, Cercopithecus nictitans stampflii, occurat various sites in West Africa, particularly in the transitionzone between rainforest and savannah. The species is sometimesseen in primary rainforest, although at a curiously low densitycompared with that of other monkey species. We conducted a 24-monthfield study in the tropical rainforest of Taï NationalPark, Ivory Coast, and found that putty-nosed monkeys requirean ecological niche almost identical to that of the Diana monkeys,Cercopithecus diana diana. Moreover, the niche breadth of putty-nosedmonkeys was significantly decreased in the presence of Dianamonkeys, suggesting that feeding competition with Diana monkeyskept putty-nosed monkeys from successfully colonizing a rainforesthabitat. However, contrary to the interspecies competition hypothesis,groups of both species almost completely overlapped in homeranges and formed near-permanent mixed-species associations,rather than avoiding each other. We hypothesized that Dianamonkeys tolerated immigrating putty-nosed monkeys and formedmixed-species groups with them, despite high levels of competition,because of their merit in predation defense. Direct observationsand a series of field experiments confirmed that male putty-nosedmonkeys play a vital role in defense against crowned eagles,suggesting that putty-nosed monkeys obtain access to feedingtrees by offering antipredation benefits to Diana monkeys. Wediscuss these findings in light of biological market theory.     

The formation of red colobus-diana monkey associations under predation pressure from chimpanzees.

It is generally assumed that most primates live in monospecific or polyspecific groups because group living provides protection against predation, but hard evidence is scarce. We tested the antipredation hypothesis with observational and experimental data on mixed-species groups of red colobus (Procolobus badius) and diana monkeys (Cercopithecus diana) in the Taï National Park, Ivory Coast. Red colobus, but not diana monkeys, are frequently killed by cooperatively hunting chimpanzees. Association rates peaked during the chimpanzees'' hunting season, as a result of changes in the behaviour of the red colobus. In addition, playbacks of recordings of chimpanzee sounds induced the formation of new associations and extended the duration of existing associations. No such effects were observed in reaction to control experiments and playbacks of leopard recordings.     

西双版纳茶林及橡胶林区鸟类调查

1983年5—7月,作者对云南省西双版纳的勐海、勐养和勐腊三个地区的茶林和橡胶林区的鸟类区系组成情况进行了调查,其结果若与开发前（郑作新、郑宝赉,1961,1962;A.И.伊万诺夫,1961）的记录对照,在季节时间上有所不同,但该地区的热带雨林和季雨林被开垦种植为茶林、橡胶林及其它经济作物之后,反映出各林区现在的繁殖鸟种类减少很多,鸟类区系组成情况有显著的变化。     

Association of Terrestrial Mangabeys (Cercocebus atys) with Arboreal Monkeys: Experimental Evidence for the Effects of Reduced Ground Predator Pressure on Habitat Use

We examined the behavior of two arboreal monkeys—Piliocolobus badius (western red colobus) and Cercopithecus diana (Diana monkey)—in the presence and absence of a third, predominately terrestrial monkey, Cercocebus atys (sooty mangabey) in the Ivory Coast's Taï Forest. Via experimental data, we show that sooty mangabeys are effective sentinels for ground predators. Then we compared strata use of red colobus and Diana monkeys in the presence and absence of mangabeys. Our data indicate that red colobus and Diana monkeys descend to low forest levels and to the forest floor significantly more often when mangabeys are spatially intermingled with them, i.e. in polyspecific associations. The niche extension may provide some foraging advantage, especially for red colobus. We identified the specific causal agent—relaxed ground predator pressure—of a temporary shift in habitat use. In this instance, the presence of one species (Cercocebus atys) leads to the temporary expansion of a niche normally avoided by others Piliocolobus badius and Cercopithecus diana.     

Effects of habitat disturbance and food supply on population densities of three primate species in the Kakamega Forest, Kenya

While habitat disturbance and food availability are major factors thought to determine the abundance of primates, evidence for their importance is uneven. We assessed the effects of these factors on three monkey species, guerezas ( Colobus guereza ) , blue monkeys ( Cercopithecus mitis ) and redtails ( Cercopithecus ascanius ), in four areas of the Kakamega Forest, Kenya. Group densities of guerezas and blue monkeys were higher in areas where disturbance levels were also higher. Food availability measured as basal area density of food trees did not correlate significantly with the group densities of any of the three monkeys. The diversity of food trees, another potential measure of food abundance did, however, correlate with group densities of guerezas and blue monkeys suggesting that food availability may positively influence monkey density, and may sometimes increase with disturbance. Group densities of redtails did not correlate with any habitat variable examined, suggesting that factors other than those we considered may have influenced the abundance of this species particularly.     

Ranging behaviour and its relation to diet selection in Tana River Red colobus (Colobus badius rufomitratus)

Ranging behaviour of a group of Red colobus monkeys, Colobus badius rufomitratus, on the Tana River, Kenya, is described. The main study group had a mean day journey of 603 in and an annual range area of 9 ha (measured in 0.25 ha quadrats). Relationships between ranging patterns and diet selection are sought on both spatial and temporal dimensions. Differential use of the annual range is related to the distribution of major food species, while the diversity of monthly ranging patterns is inversely related to the availability of young growth in the canopy. These findings lead to a discussion of why range areas at the Tana are much smaller than those reported from rain-forest sites.