Identification of a new simian immunodeficiency virus lineage with a vpu gene present among different cercopithecus monkeys (C. mona, C. cephus, and C. nictitans) from Cameroon

During a large serosurvey of wild-caught primates from Cameroon, we found 2 mona monkeys (Cercopithecus mona) out of 8 and 47 mustached monkeys (Cercopithecus cephus) out of 302 with human immunodeficiency virus (HIV)-simian immunodeficiency virus (SIV) cross-reactive antibodies. In this report, we describe the full-length genome sequences of two novel SIVs, designated SIVmon-99CMCML1 and SIVmus-01CM1085, isolated from one mona (CML1) and one mustached (1085) monkey, respectively. Interestingly, these viruses displayed the same genetic organization (i.e., presence of a vpu homologue) as members of the SIVcpz-HIV type 1 lineage and SIVgsn isolated from greater spot-nosed monkeys (Cercopithecus nictitans). Phylogenetic analyses of SIVmon and SIVmus revealed that these viruses were genetically distinct from other known primate lentiviruses but were more closely related to SIVgsn all across their genomes, thus forming a monophyletic lineage within the primate lentivirus family, which we designated the SIVgsn lineage. Interestingly, mona, mustached, and greater spot-nosed monkeys are phylogenetically related species belonging to three different groups of the genus Cercopithecus, the C. mona, C. cephus, and Cercopithecus mitis groups, respectively. The presence of new viruses closely related to SIVgsn in two other species reinforces the hypothesis that a recombination event between ancestral SIVs from the family Cercopithecinae is the origin of the present SIVcpz that is widespread among the chimpanzee population.     

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.     

Sex chromosome phylogenetics indicate a single transition to terrestriality in the guenons (tribe Cercopithecini)

This is the first molecular study to trace the evolutionary transition in substrate preference across a primate radiation. We surveyed 20 guenons (tribe Cercopithecini) and 4 outgroup taxa for two Y-chromosomal genes, TSPY ( approximately 2240 bp) and SRY ( approximately 780 bp), and one X-chromosomal intergenic region ( approximately 1600 bp) homologous to a fragment of human Xq13.3. Parsimony and maximum likelihood analyses of the sex chromosomal datasets consistently cluster the three terrestrial taxa, Cercopithecus aethiops, Cercopithecus lhoesti, and Erythrocebus patas, into a group that is reciprocally monophyletic with a clade of arboreal Cercopithecus spp. Given that the common ancestor of the two clades was most likely an arboreal taxon, this phylogenetic pattern suggests the transition to terrestriality occurred only once among the extant guenons. This pattern also indicates that the genus Cercopithecus is paraphyletic, as presently defined, and calls for taxonomic revision so that the nomen describes a strictly monophyletic group. We outline four acceptable taxonomic schemes and suggest that the most appropriate is to reassign C. aethiops, C. lhoesti, and E. patas to the resurrected genus Chlorocebus. Finally, while the phylogeny and taxonomy of terrestrial guenons were the focus of this study, the X-chromosome sequences presented here represent the first molecular evidence to unambiguously place Allenopithecus nigroviridis as the basal lineage of the tribe Cercopithecini.     

Ranging Patterns in Relation to Seasonality and Frugivory Among Cercopithecus campbelli, C. petaurista, and C. diana in the Taï Forest

My objective is to better understand the influences of seasonality and frugivory on ranging patterns for 3 guenon species of the Ta? Forest: Cercopithecus campbelli, C. petaurista, and C. diana). Over a 17-mo period, I gathered data on the daily path length, home range size, and home range use for 2 habituated groups of each species. The ranging patterns of the 3 species were very similar to each other and across seasons. Further, the ranging patterns were not closely related to fruit abundance or consumption. Each species had a long-ranging strategy with long daily ranges relative to home range size and little repeated use of areas on successive days, which may relate to territory and boundary patrols, especially for Cercopithecus diana. I compare them with other guenon communities and demonstrate that the ranging patterns of the 3 species are more similar than the ranging patterns of sympatric guenons in other communities. I discuss the results in relation to the association of Cercopithecus campbelli and C. petaurista with C. diana for antipredator benefits.     

Diet and serum lipid responses of seven species ofCercopithecus monkeys: A preliminary study

Four social groups ofCercopithecus monkeys, includingaethiops, cephus, diana, mitis, mona, neglectus, andpetaurista (48 individuals), were fed a diet containing 0.68 mg cholesterol per kcal. Blood samples were analyzed for serum cholesterol and triglycerides. Elevations in serum cholesterol appear to be individualistic and exhibit only slight differences between species and social groups.     

Gastrointestinal parasites of the guenons of western Uganda

From January 1998 to December 2002, we collected 293 fecal samples from free-ranging individuals of the 4 guenon species of western Uganda, i.e., redtail guenons (Cercopithecus ascanius), blue monkeys (Cercopithecus mitis), l'hoesti monkeys (Cercopithecus lhoesti), and vervet monkeys (Cercopithecus aethiops), to quantify the prevalence of gastrointestinal parasites. Helminth eggs, larvae, and protozoan cysts were isolated by sodium nitrate flotation and fecal sedimentation. Helminth parasites were identified, and infection prevalence was determined for all 4 guenon species. Coprocultures facilitated identification of strongylate nematodes. For the most common species, the redtail guenon, we documented prevalence of protozoan parasites and examined the effect of season and host sex on infection prevalence. Six nematodes (Strongyloidesfulleborni, Oesophagostomum sp., unidentified strongyle, Trichuris sp., Streptopharagus sp., and Enterobius sp.), 1 cestode (Bertiella sp.), 1 trematode (Dicrocoeliidae), and 5 protozoans (Entamoeba coli, Entamoeba histolytica, lodameoba butschlii, Giardia lamblia, and Chilomastix mesnili) were detected. Seasonal patterns of infection were not readily apparent for any parasite species infecting redtail guenons. Although prevalence never differed between male and female guenons, only adult females were infected with Oesophagostomum sp. and S. fulleborni.     

New simian immunodeficiency virus infecting De Brazza's monkeys (Cercopithecus neglectus): evidence for a cercopithecus monkey virus clade

Nearly complete sequences of simian immunodeficiency viruses (SIVs) infecting 18 different nonhuman primate species in sub-Saharan Africa have now been reported; yet, our understanding of the origins, evolutionary history, and geographic distribution of these viruses still remains fragmentary. Here, we report the molecular characterization of a lentivirus (SIVdeb) naturally infecting De Brazza's monkeys (Cercopithecus neglectus). Complete SIVdeb genomes (9,158 and 9227 bp in length) were amplified from uncultured blood mononuclear cell DNA of two wild-caught De Brazza's monkeys from Cameroon. In addition, partial pol sequences (650 bp) were amplified from four offspring of De Brazza's monkeys originally caught in the wild in Uganda. Full-length (9068 bp) and partial pol (650 bp) SIVsyk sequences were also amplified from Sykes's monkeys (Cercopithecus albogularis) from Kenya. Analysis of these sequences identified a new SIV clade (SIVdeb), which differed from previously characterized SIVs at 40 to 50% of sites in Pol protein sequences. The viruses most closely related to SIVdeb were SIVsyk and members of the SIVgsn/SIVmus/SIVmon group of viruses infecting greater spot-nosed monkeys (Cercopithecus nictitans), mustached monkeys (Cercopithecus cephus), and mona monkeys (Cercopithecus mona), respectively. In phylogenetic trees of concatenated protein sequences, SIVdeb, SIVsyk, and SIVgsn/SIVmus/SIVmon clustered together, and this relationship was highly significant in all major coding regions. Members of this virus group also shared the same number of cysteine residues in their extracellular envelope glycoprotein and a high-affinity AIP1 binding site (YPD/SL) in their p6 Gag protein, as well as a unique transactivation response element in their viral long terminal repeat; however, SIVdeb and SIVsyk, unlike SIVgsn, SIVmon, and SIVmus, did not encode a vpu gene. These data indicate that De Brazza's monkeys are naturally infected with SIVdeb, that this infection is prevalent in different areas of the species' habitat, and that geographically diverse SIVdeb strains cluster in a single virus group. The consistent clustering of SIVdeb with SIVsyk and the SIVmon/SIVmus/SIVgsn group also suggests that these viruses have evolved from a common ancestor that likely infected a Cercopithecus host in the distant past. The vpu gene appears to have been acquired by a subset of these Cercopithecus viruses after the divergence of SIVdeb and SIVsyk.     

Blood markers and genetic evolution in Cercopithecinae

Serum proteins and RBC enzymes were surveyed in 16 species (183 animals) of African guenons (tribe Cercopithecini) in order to determine their genetic polymorphism and to establish dendrograms on the basis of their allele frequencies. The molecular data obtained were compared with those of mangabeys (16 animals tested) and discussed in the light of our results inPapio andMacaca. The species surveyed wereCercopithecus neglectus, C. hamlyni, C. l'hoesti (C. l'h. l'hoesti, C. l'h. preussi, andC. l'h. solatus), C. nictitans, C. mitis (C. m. kolbi, C. m. albotorquatus, C. m. stuhlmanni, andC. m. albogularis), C. cephus, C. ascanius, C. erythrotis, C. petaurista, C. mona, C. pogonias, C. wolfi, andC. aethiops, Miopithecus talapoin, Allenopithecus nigroviridis andErythrocebus patas, Lophocebus albigena, andCercocebus torquatus. Eleven loci (ten systems) were studied in red blood cell enzymes and the Gc, Gm, Km, and Bm systems in DBP and immunoglobulin serum proteins. Most of the loci were polymorphic. Similar and different polymorphisms occur in closely related species or subspecies, particularly inCercopithecus. Guenons have phenotypes clearly distinct from mangabeys.     

Characterization of a novel vpu-harboring simian immunodeficiency virus from a Dent's Mona monkey (Cercopithecus mona denti)

We report the identification of a new simian immunodeficiency virus (SIV), designated SIVden, in a naturally infected Dent's Mona monkey (Cercopithecus mona denti), which was kept as pet in Kinshasa, capital of the Democratic Republic of Congo. SIVden is genetically distinct from the previously characterized primate lentiviruses. Analysis of the full-length genomic sequence revealed the presence of a vpu open reading frame. This gene is also found in the virus lineage of human immunodeficiency virus type 1 (HIV-1) and chimpanzee immunodeficiency virus (SIVcpz) and was recently described in viruses isolated from Cercopithecus nictitans, Cercopithecus mona, and Cercopithecus cephus. The SIVden vpu coding region is shorter than the HIV-1/SIVcpz and the SIVgsn, SIVmon, and SIVmus counterparts. Unlike Pan troglodytes schweinfurthii viruses (SIVcpzPts) and Cercopithecus monkey viruses (SIVgsn, SIVmon, and SIVmus), the SIVden Vpu contains the characteristic DSGXES motif which was shown to be involved in Vpu-mediated CD4 and IkappaBalpha proteolysis in HIV-1 infected cells. Although it harbors a vpu gene, SIVden is phylogenetically closer to SIVdeb isolated from De Brazza's monkeys (Cercopithecus neglectus), which lacks a vpu gene, than to Cercopithecus monkey viruses, which harbor a vpu sequence.     

Variation in guenon skulls (I): species divergence, ecological and genetic differences

Guenons are the most diverse clade of African monkeys. They have varied ecologies, include arboreal and terrestrial species, and can be found in nearly every region of sub-Saharan Africa. Species boundaries are often uncertain, with a variable number of species and subspecies mostly recognised on the basis of their geographic distribution and pelage. If guenon soft tissue patterns show high variability, the same does not seem to hold for skull morphology. Guenon skulls are traditionally considered relatively undifferentiated and homogeneous. However, patterns of variation in skulls have never been examined using a large number of specimens sampled across the breadth of species diversity. Thus, in the present study, skulls of adult guenons and two outgroup species are analysed using three-dimensional geometric morphometrics. Three-dimensional coordinates of 86 anatomical landmarks were measured on 1,315 adult specimens belonging to all living guenon species except Cercopithecus dryas. Species are well-discriminated using shape but the best discrimination occurs when species have either a long evolutionary history (e.g., Allenopithecus nigroviridis) or represent extremes of size variation (Miopithecus sp. and Erythrocebus patas). Interspecific phenetic relationships reflect size differences. Four main clusters are found that mainly correspond to four size groups: the smallest species (Miopithecus sp.), the largest species (E. patas plus the study outgroups), a group of medium-small arboreal guenons, and a group of medium-large arboreal and terrestrial guenons. Correlations between interspecific shape distances and interspecific differences in size are higher than between shape distances and genetic distances. However, if only the component of interspecific shape variation which is not correlated to evolutionary allometry is used in the comparison with genetic distances, correlations are up to 1.4 times larger than those including allometric shape. The smallest correlations are those between shape and ecological distances, which is consistent with the lack of clusters clearly reflecting broad ecological specialisations (e.g., arboreality versus terrestriality). Thus, size, which is generally considered more evolutionarily labile than shape, seems to have played a major role in the evolution of the guenons. The incongruence between interspecific shape differences and phylogeny might be explained by a large proportion of shape changes having occurred along allometric trajectories that tend to be conserved within this clade.     

Group size and group composition of the mona monkey (Cercopithecus mona) on the Island of Grenada,West Indies

Cercopithecus, the genus of guenons, is the largest of the African primate genera, and yet more than half of the species belonging to this group have never been the focus of a long-term field study. In this paper, I present data on group size and composition for a previously unstudied population of guenons on the Caribbean island of Grenada. The mona monkey, Cercopithecus mona, was introduced to Grenada from Africa approximately 200–300 years ago. Two types of social groups were found for Cercopithecus mona on Grenada: all-male groups consisting of two to four individuals and bisexual groups containing 5–32 individuals. All-male groups of Grenada mona monkeys contained any combination of juveniles, subadults, and/or adults. All-male groups were a common occurrence on Grenada but have never been reported for African C. mona and have been reported only in two other forest Cercopithecusspecies. Bisexual groups appeared to consist of one adult male, one to six adult females, subadult females, and juveniles and infants of both sexes. Even though no more than one adult male was ever seen in each bisexual group of monas on Grenada, other males were heard giving copulation calls simultaneously with resident adult male loud calls, suggesting that other males occasionally infiltrate bisexual groups. Am. J. Primatol. 43:167–173, 1997. © 1997 Wiley-Liss, Inc.     

Primate genus Miopithecus: evidence for the existence of species and subspecies of dwarf guenons based on cellular and endogenous viral sequences

Sequence data from the mitochondrial 12S rRNA gene were combined with endogenous retrovirus sequences to study the position of the genus Miopithecus in the primate tree. The mitochondrial sequences indicated that Miopithecus is a true genus distinct from Cercopithecus, although talapoin monkeys are commonly referred to as dwarf guenons. The existence of two species of dwarf guenons, suggested by differences in coat color, pigmentation, and geographic location, was supported by substantial mitochondrial 12S rRNA gene divergence. In line with the informal proposal of J. Kingdon (1997, "The Kingdon Field Guide to African Mammals," Academic Press, London), we use the names Miopithecus talapoin for the southern, darker species and Miopithecus ougouensis for the northern, lighter-colored monkeys. Different 12S rRNA gene haplotypes found in M. ougouensis individuals suggest the possible existence of additional subspecies. Simian endogenous retrovirus (SERV) strain 23. 1 proviruses were introduced in the primate germ-line after the Cercopithecinae split from the Colobinae, estimated at around 9-14 million years ago. SERV sequences were used for timing of divergence events in Cercopithecinae and confirmed the close relationship between the genera Cercopithecus and Miopithecus, which was only weakly supported by the more variable mtDNA sequences in a distance analysis, demonstrating the utility of these pseudogenes in phylogenetic grouping.     

The evolution of the Cercopithecini: a (post)modern synthesis

The Cercopithecini, or African guenon monkeys, are one of the most diverse clades of living primates and comprise the most species‐rich clade of Catarrhini. Species identity is announced by flamboyant coloration of the facial and genital regions and, more cryptically, by vigorous chromosomal rearrangements among taxa. Beneath the skin, however, these animals are skeletally conservative and show low levels of genetic sequence divergence consonant with recent divergence between congeneric species. The guenons clearly demonstrate that morphological, cytogenetic, and reproductive differentiation proceed at different rates during speciation. We review diverse kinds of data in an effort to understand this conundrum.     

Red-Tailed Guenons (Cercopithecus ascanius) and Strychnos mitis: Evidence for Plant Benefits Beyond Seed Dispersal

I report data collected on red-tailed guenon (Cercopithecus ascanius schmidti) fruit processing behaviors between June 1993 and April 1994. Red-tailed guenons consumed the fruit of Strychnos mitis in 542 of 2,930 fruit-eating events (FEEs). The monkeys spat out cleaned seeds of Strychnos mitis in a majority of these records (477/542; 88%); seeds were occasionally swallowed whole, but only when pulp was unripe (69/542; 12%). In 83% of the FEEs on Strychnos, the red-tailed guenons spat out seeds within 10 m of the removal site; they typically stayed in the same tree while processing fruit, and in 56% of the FEEs, they moved <1 m before spitting seeds. I monitored spat seeds to evaluate the impact of monkey fruit processing on seed fate. Results indicate that 83% of seeds spat out by the red-tailed guenons germinated, while only 12% of unprocessed seeds survived to germination (p < 0.01). Of the processed seeds that germinated, 60% survived to germination and seedling establishment, while only 5% of unprocessed seeds survived to seedling establishment (p < 0.01). Unprocessed seeds were also more likely to be attacked by seed predators (p < 0.01) and fungus (p < 0.01). Although there is generally high mortality in seeds/seedlings, mature trees of Strychnos mitis are found in groves of adults, under which dense populations of seedlings and saplings can occur. These data suggest that Strychnos mitis does not conform to expectations of the Janzen-Connell model of seed escape from parent trees. Instead, I suggest that by removing pulp, a process that results in a reduction of fungal pathogen attack, red-tailed guenons positively effect the seed survivorship of Strychnos mitis. Although this effect has been observed in pulp-cleaning ant species, it is a hitherto undescribed effect of primates on their fruit resources.     

Time Constraints Do Not Limit Group Size in Arboreal Guenons but Do Explain Community Size and Distribution Patterns

To understand how species will respond to environmental changes, it is important to know how those changes will affect the ecological stress that animals experience. Time constraints can be used as indicators of ecological stress. Here we test whether time constraints can help us understand group sizes, distribution patterns, and community sizes of forest guenons (Cercopithecus/Allochrocebus). Forest guenons typically live in small to medium sized one-male–multifemale groups and often live in communities with multiple forest guenon species. We developed a time-budget model using published data on time budgets, diets, body sizes, climate, and group sizes to predict maximum ecologically tolerable group and community sizes of forest guenons across 202 sub-Saharan African locations. The model correctly predicted presence/absence at 83% of these locations. Feeding-foraging time (an indicator of competition) limited group sizes, while resting and moving time constraints shaped guenon biogeography. Predicted group sizes were greater than observed group sizes but comparable to community sizes, suggesting community sizes are set by competition among guenon individuals irrespective of species. We conclude that time constraints and intraspecific competition are unlikely to be the main determinants of relatively small group sizes in forest guenons. Body mass was negatively correlated with moving time, which may give larger bodied species an advantage over smaller bodied species under future conditions when greater fragmentation of forests is likely to lead to increased moving time. Resting time heavily depended on leaf consumption and is likely to increase under future climatic conditions when leaf quality is expected to decrease.     

Comparative postcranial body shape and locomotion in Chlorocebus aethiops and Cercopithecus mitis

Body weight and length, chest girth, and seven postcranial limb segment lengths are compared between two guenon species, Chlorocebus (Cercopithecus) aethiops (vervets) and Cercopithecus mitis (blue monkeys), exhibiting different habitual locomotor preferences. The subjects, all adults, were wild caught for a non-related research project (Turner et al. [1986] Genetic and morphological studies on two species of Kenyan monkeys, C. aethiops and C. mitis. In: Else JG, Lee PC, editors. Primate evolution, proceedings of the Xth International Congress of Primatology, Cambridge. London). The morphological results are interpreted within the context of previously published observations of primate locomotion and social organization. The sample is unique in that the body weight of each individual is known, allowing the effects of body-size scaling to be assessed in interspecific and intersexual comparisons. C. mitis has a significantly (P < 0.05) greater body weight and trunk length than C. aethiops. A shorter trunk may function to reduce spinal flexibility for ground-running in the latter. Proximal limb segments (arm and thigh) are significantly greater in C. mitis, reflecting known adaptations to committed arboreal quadrupedal locomotion. By contrast, relative distal limb segments (forearm, crus, and foot) are significantly longer in C. aethiops, concordant with a locomotor repertoire that includes substantial terrestrial quadrupedalism, in addition to arboreal agility, and also the requisite transition between ground and canopy. Although normally associated with arboreal monkeys, greater relative tail length occurs in the more terrestrial vervets. However, because vervets exploit both arboreal and terrestrial habitats, a longer tail may compensate for diminished balance during arboreal quadrupedalism resulting from the greater "brachial" and "crural" indices that enhance their ground quadrupedalism. Most interspecific differences in body proportions are explicable by differences in locomotor modalities. Some results, however, contradict commonly held "tenets" that relate body size and morphology exclusively to locomotion. Generally associated with terrestriality, sexual dimorphism (male/female) is greater in the more arboreal blue monkeys. A more intense, seasonal mating competition may account for this incongruity.     

Locomotion and habitat use of Stampflii's putty-nosed monkey (Cercopithecus nictitans stampflii) in the Taï National Park, Ivory Coast

Stampflii's putty-nosed monkeys Cercopithecus nictitans stampflii are large, rare, poorly known guenons with a discontinuous distribution in West Africa. Putty-nosed monkeys occur at low densities in Ivory Coast's Tai National Park where they are believed to have migrated from less forested regions north of the park (Eckardt and Zuberbühler: Behav Ecol 15 (2004) 400-411). In this article, we compare the positional behavior of Putty-nosed monkeys to that of three other guenon species in the Tai Forest and provide an additional test of the relationship among locomotion, body size, maintenance activity, and habitat use in generalized arboreal quadrupeds. Our results indicate that putty-nosed monkeys confine the great majority of their locomotion to the main canopy where most movement occurs on boughs and branches. Comparison of overall locomotor frequencies across the four Tai guenon species reveals a general pattern; quadrupedalism accounts for at least 70% of the total profile, climbing ranges between 15 and 20%, and leaping accounts for 5-10% of total movement. The overall locomotor consistency among the taxa, especially between putty-nosed and Diana monkeys, is significant considering marked interspecific differences in support preference and strata use.     

Molecular phylogeny of Old World monkeys (Cercopithecidae) as inferred from gamma-globin DNA sequences

DNA sequence data of the nuclear-encoded gamma1-gamma2-globin duplication region were used to examine the phylogenetic relationships of 16 cercopithecid (Old World monkey) species representing 12 extant genera. Morphology- and molecular-based hypotheses of Old World monkey branching patterns are generally congruent, except for generic relationships within the subtribe Papionina. The cercopithecids divide into colobines (leaf-eating monkeys) and cercopithecines (cheek-pouched monkeys). The colobines examined by the DNA data divide into an Asian clade (Nasalis, proboscis monkeys; Trachypithecus, langurs) and an African clade (Colobus, colobus monkeys). The cercopithecines divide into tribes Cercopithecini (Erythrocebus, patas monkey; Chlorocebus, green monkeys; Cercopithecus, guenons) and Papionini. Papionins divide into subtribes Macacina (Macaca, macaques) and Papionina (Papio, hamadryas baboons; Mandrillus, drills and mandrills; Theropithecus, gelada baboons; Lophocebus, arboreal mangabeys; Cercocebus, terrestrial mangabeys). In a morphologically based classification, Mandrillus is a subgenus of Papio, whereas Lophocebus is a subgenus of Cercocebus. In contrast, the molecular evidence treats Mandrillus as a subgenus of Cercocebus, and treats both Theropithecus and Lophocebus as subgenera of Papio. Local molecular clock divergence time estimates were used as a yardstick in a "rank equals age" system to propose a reduction in taxonomic rank for most clades within Cercopithecidae.     

Examining deep litter as environmental enrichment for a family group of wolf's guenons,Cercopithecus wolfi

Manipulable substrates promote species‐typical behavior and decrease abnormal behavior in a variety of primate species. However, the effects of providing litter to arboreal primates are not as well studied, and there is little information specifically concerning enrichment for guenons. To inform the captive management of an under‐studied species, we evaluated deep litter substrate as environmental enrichment for a family group of Wolf's guenons, Cercopithecus wolfi. We expected it to promote species‐typical behavior and to act as an intervention by reducing aggressive behaviors targeted toward a juvenile group member by his parents. We compared the guenons' behavior during baseline periods in which normal husbandry routines were followed to periods when the entire floor of their enclosure was covered with 30 cm of straw or wood wool. We then evaluated the group's preference between litters by comparing their relative use. The guenons spent more time feeding and were more active during both litter conditions, but relative to their respective baselines, they spent more time examining straw and less time examining wood wool. Straw, but not wood wool, promoted some affiliative behavior as well as greater tolerance of the juvenile's social proximity to others. However, the addition of deep litter did not ameliorate patterns of agonistic behavior among our subjects. Our results suggest that straw conferred greater behavioral benefits than wood wool as a deep litter substrate for this guenon group and may constitute a form of environmental enrichment for this species. Zoo Biol 29:626–632, 2010. © 2009 Wiley‐Liss, Inc.     

Better to Live with Allogenerics Than to Live Alone? The Case of Single Male Cercopithecus pogonias in Troops of Colobus satanas

We report the integration of single male crowned guenons (Cercopithecus pogonias) into troops of black colobus (Colobus satanas). We observed one male Cercopithecus pogonias in three troops of Colobus satanas on 30% of observation days (n = 231). Activities of single males guenons did not differ significantly from those of the colobus with which they associated. Moreover, both species performed simultaneously the same activities more often than expected by chance. Interspecific grooming occurred on several occasions. Furthermore, single male guenons spent as much in time social activities when part of a colobus troop, as they typically do when part of a conspecific group. Unlike solitary male crowned guenons, which are silent, a male that is integrated into a troop of colobus is vocal and emits social alarm calls to which colobus monkeys respond. During the single file movements of colobus troops, single male crowned guenons were integrated in the core of the troop and used the same branches at the same height with the colobus. Thus, the life of a single male crowned guenon with black colobus was social. We suggest that the main benefits that he gained is the possibility to live in a social context. Social interactions could be the key element to explain why single males Cercopithecus pogonias join troops of monkeys so different from their natal groups.