Oral Processing Behavior of Diana Monkeys (Cercopithecus diana) in Taï National Park, Côte d'Ivoire.

Primates are hypothesized to "fall back" on challenging-to-process foods when preferred foods are less available. Such dietary shifts may be accompanied by changes in oral processing behavior argued to be selectively important. Here, we examine the oral processing behavior of Diana monkeys (Cercopithecus diana) in the Taï Forest across their dietary breadth, testing relationships among food intake, fruit availability, preference, and oral processing behaviors including those involved in food ingestion and breakdown. We conducted 1,066 focal follows from April 2016 to September 2017 documenting frequencies of incisor, canine, and cheek tooth mastications (i.e., chews) per ingestive action (n = 11,906 feeding events). We used phenological survey and scan sample data collected between 2004 and 2009 to examine dietary preference and food availability. Our analyses show that Diana monkeys processed foods in significantly different ways (H2 = 360.8, p < 0.001), with invertebrates requiring the least oral processing, fruit requiring intermediate amounts, and leaves requiring the most oral processing. There was no relationship between fruit availability and consumption of preferred or nonpreferred fruits (p > 0.05); however, preferred fruits were processed with significantly fewer mastications (i.e., less chewing) than nonpreferred fruits (U = 6,557, p < 0.001). We thus demonstrate that, when preferred foods are scarce, Diana monkeys do not fall back on difficult-to-process foods. Changes in processing profiles occurred throughout the year and not solely when preferred foods were in short supply. Though preferred fruits required less processing than nonpreferred fruits, we found no relationship between fruit preference and fruit availability. Diana monkeys' lack of readily identifiable fallback foods may be attributable to the relatively high tree diversity and productivity of the Taï Forest. We conclude that Diana monkeys engage in resource switching, consuming a relatively easy-to-process diet year-round.

Influence of food availability, plant productivity, and indigenous forest use on ranging behavior of the endangered samango monkey (Cercopithecus albogularis schwarzi), in the Soutpansberg Mountains, South Africa.

Understanding the determinants of ranging patterns in species susceptible to habitat fragmentation is fundamental for assessing their long-term adaptability to an increasingly human-dominated landscape. The aim of this study was to determine and compare the influence of ground-based food availability, remotely sensed plant productivity, and indigenous forest use on the ranging patterns of the endangered samango monkey (Cercopithecus albogularis schwarzi). We collected monthly ranging data on two habituated samango monkey groups, from February 2012 to December 2016, from our field site in the Soutpansberg Mountains, South Africa. We used linear mixed models to explore how food availability, plant productivity, and indigenous forest use influenced monthly ranging patterns, while controlling for group size, number of sample days and day length. We found that as more areas of high plant productivity (derived from remotely sensed EVI) were incorporated into the ranging area, both total and core monthly ranging areas decreased. In addition, both total ranging area and mean monthly daily path length decreased as more indigenous forest was incorporated into the ranging area. However, we found no effect of either ground-based food availability or remotely sensed plant productivity on ranging patterns. Our findings demonstrate the behavioral flexibility in samango monkey ranging, as samangos can utilize matrix habitat during periods of low productivity but are ultimately dependent on access to indigenous forest patches. In addition, we highlight the potential of using remotely sensed areas of high plant productivity to predict ranging patterns in a small ranging, forest-dwelling guenon, over ground-based estimates of food availability.

Validating a non-invasive technique for monitoring physiological stress in the samango monkey.

The non-invasive monitoring of physiological stress can provide conservation and wildlife managers with an invaluable tool for assessing animal welfare and psychological health of captive and free-ranging populations. A significant decrease in free-ranging primate populations globally and an increase in captive-housed primates have led to a need to monitor the stress and general welfare of these animals. We examined the suitability of three enzyme immunoassays (EIAs) for monitoring stress-related physiological responses in the samango monkey, Cercopithecus albogularis erythrarchus. We conducted an adrenocorticotropic hormone (ACTH) challenge on a male and female at the National Zoological Garden, Pretoria, South Africa. Individual faecal samples were collected 8 days pre- and post-ACTH administration and subsequently analysed for faecal glucocorticoid metabolite (fGCM) concentrations. During the study, biological stressors occurred for both the male and female. Two of the three EIAs tested (11-oxoetiocholanolone I and II) were able to reliably monitor fGCM alterations throughout the study period in both sexes. The 11-oxoetiocholanolone I EIA, however, had the lowest mean deviation from the calculated baseline value and was thus chosen as the preferred assay. Both the physiological activation of the stress response and the biological response to a stressor could be monitored with the chosen assay. The successful establishment of a reliable, non-invasive method for monitoring adrenocortical activity in C. albogularis erythrarchus will now allow conservationists, scientific researchers and wildlife managers to evaluate the level of stress experienced, and general welfare, by animals in captivity as well as free-ranging populations.

Skeletal morphology of the lesula (Cercopithecus lomamiensis) and the evolution of guenon locomotor behavior.

OBJECTIVES: The guenons (tribe Cercopithecini) are a diverse and primarily arboreal radiation of Old World monkeys from Africa. However, preliminary behavioral observations of the lesula (Cercopithecus lomamiensis), a little-known guenon species described in 2012, report it spending substantial amounts of time on the ground. New specimens allow us to present the first description of lesula postcranial morphology and apply a comparative functional morphology approach to supplement our knowledge of its locomotor behavior. MATERIALS AND METHODS: To infer the substrate use preferences of the lesula, 22 postcranial variables correlated with locomotion were assessed in a sample of 151 adult guenon specimens, including two C. lomamiensis. Using multivariate statistical analyses, we predict the amount of time the lesula spends on the ground relative to the comparative sample. RESULTS: Results suggest that the lesula spends nearly half its time on the ground, and the two available individuals were classified as semiterrestrial and terrestrial with strong support. Comparisons with two outgroup cercopithecid taxa (Colobus guereza and Macaca mulatta) demonstrate that, as a group, guenons retain signals of a generalized, semiterrestrially adapted postcranium compared to specialized arboreal cercopithecids. DISCUSSION: These results corroborate preliminary behavioral observations of the lesula as a semiterrestrial to terrestrial primate and imply multiple evolutionary transitions in substrate use among the guenon radiation. A broader view of cercopithecoid evolution suggests that a semiterrestrial ancestor for extant guenons is more parsimonious than an arboreal one, indicating that the arboreal members of the group are probably recently derived from a more semiterrestrial ancestor.

Adapting Methodology Used on Captive Subjects for Estimating Gut Passage Time in Wild Monkeys.

Gut passage time of food has consequences for primate digestive strategies, which subsequently affect seed dispersal. Seed dispersal models are critical in understanding plant population and community dynamics through estimation of seed dispersal distances, combining movement data with gut passage times. Thus, developing methods to collect in situ data on gut passage time are of great importance. Here we present a first attempt to develop an in situ study of gut passage time in an arboreal forest guenon, the samango monkey (Cercopithecus albogularis schwarzi) in the Soutpansberg Mountains, South Africa. Cercopithecus spp. consume large proportions of fruit and are important seed dispersers. However, previous studies on gut passage times have been conducted only on captive Cercopithecusspp. subjects, where movement is restricted, and diets are generally dissimilar to those observed in the wild. Using artificial digestive markers, we targeted provisioning of a male and a female samango monkey 4 times over 3 and 4 days, respectively. We followed the focal subjects from dawn until dusk following each feeding event, collecting faecal samples and recording the date and time of deposition and the number of markers found in each faecal sample. We recovered 6.61 ± 4 and 13 ± 9% of markers from the male and the female, respectively, and were able to estimate a gut passage window of 16.63-25.12 h from 3 of the 8 trials. We discuss methodological issues to help future researchers to develop in situ studies on gut passage times.

Differential responses of non-human primates to seasonal temperature fluctuations.

Non-human primates (NHPs) can adapt to conditions outside of their natural habitat and climatic ranges but this can be influenced by inherent evolutionary traits or plasticity of species that evolved in diverse environmental conditions. In this study, we investigated how five species of NHPs that have natural distributions across a range of climatic conditions responded to seasonal temperature changes in a captive environment. The activity levels of NHPs were affected by temperature changes over the season, where activity levels were generally reduced at the lower and higher temperature ranges. Species that are naturally found within narrower and warmer climatic ranges, compared to those found in colder environments with wider fluctuations in temperature, showed more marked changes in activity levels in response to temperature changes. In lower temperature conditions, three out of five species showed significantly lower activity levels; whereas in higher temperature conditions, the activity levels of all species did not significantly decrease. The frequency of thermoregulation behaviours was higher, and use of artificial thermoregulatory sources lower, for species that did not substantially adjust their activity levels in different temperature conditions. Our results suggest that NHPs largely retained the evolutionary traits related to thermoregulation, according to the different ambient conditions they evolved in and may have low behavioural plasticity in adapting to conditions outside of their natural ranges. These results provide insights for improving conservation and captive management and may have implications for understanding NHP resilience to the increasing impact of global climate change.

Early life maternal sociality predicts juvenile sociality in blue monkeys.

Maternal effects are widespread in living organisms though little is known about whether they shape individual affiliative social behavior in primates. Further, it remains a question whether maternal effects on affiliative behavior differ by offspring sex, as they do in other physiological systems, especially in species with high levels of adult sexual dimorphism and divergence in social niches. We explored how direct and indirect experiences of maternal affiliative behavior during infancy predicted affiliative behavior approximately 1-6 years later during the juvenile period, using behavioral data from 41 wild blue monkey juveniles and their 29 mothers, and controlling for individual age, sex, and maternal rank. Female juveniles spent less time grooming with any partner and with peers the more maternal grooming they received during infancy, whereas males groomed more with any partner and with peers. Similarly, the more that mothers groomed with other adult females during subjects' infancy, female subjects played less with peers, and male subjects played more as juveniles. Further, this maternal effect on social behavior appears specific to early life, as the same aspects of mothers' sociality measured throughout subjects' development did not predict juvenile behavior. Overall, our results suggest that both direct and indirect experience of mother's affiliative behavior during infancy influence an individual's affiliation later in life that sexes respond differently to the maternal affiliation, and that the first year of life is a critical window.

Assessment and preservation of liquid and frozen-thawed Black crested mangabey (Lophocebus aterrimus) spermatozoa obtained by transrectal ultrasonic-guided massage of the accessory sex glands and electroejaculation.

The Black Crested Mangabey (Lophocebus aterrimus) is an African monkey listed as Near Threatened by the IUCN and in captivity the population is limited to 34 males. The aim of this study was to evaluate two Black Crested Mangabey males, maintained in captivity in a zoological garden and suspected of infertility, with a complete examination of their genital tract using ultrasonography, followed by recovery of semen using transrectal ultrasonic massage of the accessory sexual glands (TUMASG) and electroejaculation. One male had small testicular and accessory sex gland sizes indicative of senile hypoplasia. The other male was suspected of infertility. Four semen samples were obtained. Fresh semen was initially evaluated, diluted in Refrigeration Medium Test Yolk buffer, cooled at 15 °C and cryopreserved. Endocrine profiles (testosterone, oestradiol, FSH, LH, cortisol), prostatic specific antigen and semen variables (volume, concentration, motility by CASA, viability and acrosome status using flow cytometry, morphology, morphometry utilising TEM) were evaluated in raw, cooled and cryopreserved samples. There was no detrimental effect of cooling or cryopreservation on sperm viability and acrosomal integrity. Similar percentages of viable and acrosome-intact spermatozoa were present in cooled (for 6 h) and frozen-thawed semen samples (75.1% compared with 69.0%, P > 0.05), while progressive motility was greater in cooled, compared with frozen-thawed samples (81.5% compared with 67.3%). This study was the first in which there was evaluation of sperm variables in this species and, although this study is limited by the number of animals it provides background information for further studies using assisted reproductive technologies.

Food abundance and weather influence habitat-specific ranging patterns in forest- and savanna mosaic-dwelling red-tailed monkeys (Cercopithecus ascanius).

OBJECTIVES: Primates that live in predominantly forested habitats and open, savanna mosaics should exhibit behavioral responses to differing food distributions and weather. We compared ecological constraints on red-tailed monkey ranging behavior in forest and savanna mosaic environments. Intraspecific variation in adaptations to these conditions may reflect similar pressures faced by hominins during the Plio-Pleistocene. METHODS: We followed six groups in moist evergreen forest at Ngogo (Uganda) and one group in a savanna-woodland mosaic at the Issa Valley (Tanzania). We used spatial analyses to compare home range sizes and daily travel distances (DTD) between sites. We used measures of vegetation density and phenology to interpolate spatially explicit indices of food (fruit, flower, and leaves) abundance. We modeled DTD and range use against food abundance. We modeled DTD and at Issa hourly travel distances (HTD), against temperature and rainfall. RESULTS: Compared to Issa, monkeys at Ngogo exhibited significantly smaller home ranges and less variation in DTD. DTD related negatively to fruit abundance, which had a stronger effect at Issa. DTD and HTD related negatively to temperature but not rainfall. This effect did not differ significantly between sites. Home range use did not relate to food abundance at either site. CONCLUSIONS: Our results indicate food availability and thermoregulatory constraints influence red-tailed monkey ranging patterns. Intraspecific variation in home range sizes and DTD likely reflects different food distributions in closed and open habitats. We compare our results with hypotheses of evolved hominin behavior associated with the Plio-Pleistocene shift from similar closed to open environments.

Dietary generalists and nutritional specialists: Feeding strategies of adult female blue monkeys (Cercopithecus mitis) in the Kakamega Forest, Kenya.

Generalist primates eat many food types and shift their diet with changes in food availability. Variation in foods eaten may not, however, match variation in nutrient intake. We examined dietary variation in a generalist-feeder, the blue monkey (Cercopithecus mitis), to see how dietary food intake related to variation in available food and nutrient intake. We used 371 all-day focal follows from 24 adult females (three groups) in a wild rainforest population to quantify daily diet over 9 months. We measured food availability using vegetation surveys and phenology monitoring. We analyzed >700 food and fecal samples for macronutrient content. Subjects included 445 food items (species-specific plant parts and insect morphotypes) in their diet. Variation in fruit consumption (percentage of diet and total kcal) tracked variation in availability, suggesting fruit was a preferred food type. Fruits also constituted the majority of the diet (by calories) and some fruit species were eaten more than expected based on relative availability. In contrast, few species of young leaves were eaten more than expected. Also, subjects ate fewer young leaves (based on calories consumed) when fruit or young leaves were more available, suggesting that young leaves served as fallback foods. Despite the broad range of foods in the diet, group differences in fiber digestibility, and variation that reflected food availability, subjects and groups converged on similar nutrient intakes (grand mean ± SD: 637.1 ± 104.7 kcal overall energy intake, 293.3 ± 46.9 kcal nonstructural carbohydrate, 147.8 ± 72.4 kcal lipid, 107.8 ± 12.9 kcal available protein, and 88.1 ± 17.5 kcal structural carbohydrate; N = 24 subjects). Thus, blue monkeys appear to be food composition generalists and nutrient intake specialists, using flexible feeding strategies to regulate nutrient intake. Findings highlight the importance of simultaneously examining dietary composition at both levels of foods and nutrients to understand primate feeding ecology.

Dietary Variation in Diana Monkeys (Cercopithecus diana): The Effects of Polyspecific Associations.

Establishing dietary central tendencies and extremes remains an important goal of primate research. While habitat differences and spatial discontinuity are well-documented contributors to dietary variation, other factors including polyspecific associations may significantly impact diet through changes in strata use and/or increased feeding competition. Here, we examine polyspecific association with closely related species as a source of dietary variation in a rain forest primate. Diana monkeys (Cercopithecus diana) in Côte d'Ivoire's Taï Forest frequently participate in mixed-species groups. We use data collected over a 5-year period on 4 Diana monkey groups to examine how association affects group diets. Groups exhibited significant differences in association rates with other guenon species, but this minimally influenced diet when food categories (fruit, invertebrates, leaves) were compared: diet overlap of 4 groups across the study period ranges from 90.8 to 98.1%. Examination of species composition within food categories is more revealing: intergroup dietary overlap decreases to 69.8-79.4% across the study period when comparing species contributing to total frugivory and folivory. These data support earlier findings that Diana monkeys maintain fruit-rich diets by competitively excluding sympatric congeners while highlighting that even selective frugivores such as Diana monkeys may exhibit remarkable dietary flexibility. Our study underscores the fact that broad categorical labels can obscure significant dietary differences.

Factors of influence and social correlates of parturition in captive Campbell's monkeys: Case study and breeding data.

How nonhuman primates deal with birth, at the moment of delivery, and during the following days, remains poorly explored because of the unpredictability of this event, particularly for forest-dwelling arboreal species. Available studies highlight intra- and interspecific variation which suggest flexibility of the timing of delivery, of behavior associated with labor contractions and parturition, and the social context and ambient noise surrounding delivery. Here, we present the findings of a two-decade survey of reproduction in a population of captive Campbell's monkeys. Analysis of 34 births (with a female-biased sex ratio) showed that deliveries occurred systematically at night and more frequently during week-ends and in fall (September-November). This suggests the existence of both circadian and infradian biological rhythms. We present the first detailed description of a birth for this species and its short-term social consequences. In line with previous findings for other monkeys and apes, labor (estimated by unusual stretching postures) and parturition were rapid, and delivery occurred in a clear social and vocal context. During the following days, the new mother became the center of attention of for young (kin and non-kin) females and rose through the hierarchy. We discuss socio-ecological factors, notably captivity conditions and the high degree of tolerance in the species' social system, and confirm the existence of both "rigidity" and "flexibility" in the primates' adaptive reproductive strategies.

Persistent Females and Compliant Males Coordinate Alarm Calling in Diana Monkeys.

Sexual dimorphisms in animal vocal behavior have been successfully explained by sexual selection theory (e.g., mammals [1-5]; birds [6, 7]; anurans [8, 9]), but this does not usually include alarm calls, which are thought to be the product of kin or individual selection (e.g., [10, 11]). Here, we present the results of playback experiments with wild Diana monkeys, a species with highly dimorphic predator-specific alarms, to investigate the communication strategies of males and females during predator encounters. First, we simulated predator presence by broadcasting vocalizations of their main predators, leopards or eagles. We found that males only produced predator-specific alarms after the females had produced theirs, in response to which the females ceased alarm calling. In a second experiment, we created congruent and incongruent situations, so that the calls of a predator were followed by playbacks of male or female alarms with a matching or mismatching referent. For congruent conditions, results were the same as in the first experiment. For incongruent conditions, however, the males always gave predator-specific alarms that referentially matched the females' calls, regardless of the previously displayed predator. In contrast, females always gave predator-specific alarms that matched the predator type, regardless of their own male's subsequent calls. Moreover, the females persistently continued to alarm call until their own male produced calls with the matching referent. Results show that males and females attend to the informational content of each other's alarm calls but prioritize them differently relative to an experienced external event, a likely reflection of different underlying selection pressures.

Agonism and dominance in female blue monkeys.

Agonistic behavior features prominently in hypotheses that explain how social variation relates to ecological factors and phylogenetic constraints. Dominance systems vary along axes of despotism, tolerance, and nepotism, and comparative studies examine cross-species patterns in these classifications. To contribute to such studies, we present a comprehensive picture of agonistic behavior and dominance relationships in wild female blue monkeys (Cercopithecus mitis), an arboreal guenon, with data from 9 groups spanning 18 years. We assessed where blue monkeys fall along despotic, tolerant, and nepotistic spectra, how their dominance system compares to other primates, primarily cercopithecines, and whether their agonistic behavior matches socioecological model predictions. Blue monkeys showed low rates of mainly low-intensity agonism and little counter-aggression. Rates increased with rank and group size. Dominance asymmetry varied at different organizational levels, being more pronounced at the level of interactions than dyad or group. Hierarchies were quite stable, had moderate-to-high linearity and directional consistency and moderate steepness. There was clear maternal rank inheritance, but inconsistent adherence to Kawamura's rules. There was little between-group variation, although hierarchy metrics showed considerable variation across group-years. Overall, blue monkeys have moderately despotic, moderately tolerant, and nepotistic dominance hierarchies. They resemble other cercopithecines in having significantly linear and steep hierarchies with a generally stable, matriline-based structure, suggesting a phylogenetic basis to this aspect of their social system. Blue monkeys most closely match Sterck et al.'s [1997] Resident-Nepotistic-Tolerant dominance category, although they do not fully conform to predictions of any one socioecological model. Our results suggest that socioecological models might better predict variation within than across clades, thereby incorporating both ecological variables and phylogenetic constraints. Our findings also highlight the need for clearer definitions of socioecologically relevant dominance categories, which would ideally derive from quantitative measures of dominance behavior. Intraspecific and methodological variation may, however, be a challenge.

Degree of terrestrial activity of the elusive sun-tailed monkey (Cercopithecus solatus) in Gabon: Comparative study of behavior and postcranial morphometric data.

We carried out a multidisciplinary study linking behavioral and morphological data from a little-known guenon species, Cercopithecus solatus, endemic to Gabon. Over a period of 9 months, we documented the pattern of stratum use associated with postural and locomotor behavior by direct observation (650 hrs) of a semi-free-ranging breeding colony. We also conducted a morphometric analysis of the humerus and limb proportions of 90 adult specimens from 16 guenon species, including C. solatus. Field observations indicated that C. solatus monkeys spent a third of their time on the ground, similar to semi-terrestrial guenon species. We detected two patterns of stratum use: at ground level, and in trees, at a height of 3-10 m. The monkeys spent more time on the ground during the dry season than the wet season, feeding mainly at ground level, while resting, and social behaviors occurred more frequently in the tree strata. Our study of humerus size and shape, together with the analysis of limb proportions, indicated morphofunctional adaptation of C. solatus to greater terrestriality than previously thought. We therefore characterize C. solatus as a semi-terrestrial guenon, and propose a new hypothesis for the ancestral condition. By combining behavioral and morphological results, we provide new information about the adaptive strategies of the species, and the evolutionary history of guenons, thus contributing to the conservation of the sun-tailed monkey in the wild.

Social status and helminth infections in female forest guenons (Cercopithecus mitis).

OBJECTIVE: When resource competition within primate social groups is effective, high-ranking individuals generally gain fitness benefits. Contrary to expectations, female Cercopithecus mitis form linear dominance hierarchies without evidence for rank-related variation in fitness-relevant measures, raising questions about the evolution of guenon social structure. Here, we test whether social status predicts gastrointestinal helminth infections, known to influence health and morbidity in other mammalian hosts. In addition, we assess whether infections contribute to stress responses as indicated by fecal glucocorticoid (fGC) levels. METHODS: We quantified infections and hormone levels in 382 fecal samples from 11 adult female Sykes' monkeys (C. m. albogularis) over four months in one wild study group at Gede Ruins, Kenya. Using a generalized estimating equations technique, we modeled the odds of infection, relative infection intensity, and fGC variation. RESULTS: High-ranking females were less likely infected with Trichuris and Trichostrongylus, had lower fecal egg counts for both taxa, and overall lower helminth richness than low-ranking females. An inverse relationship between rank and Trichuris egg counts existed also in a study population of blue monkeys (C. m. stuhlmanni), where we collected comparable data over a shorter period. Regardless of rank, lactating females were more likely than non-lactating females to be infected with Trichuris, and had higher fecal egg counts for both Trichuris and Oesophagostomum. Lastly, we report evidence that Trichuris infections exacerbated energetic stress and that food supplementation by tourists increased infection levels. CONCLUSION: Our findings suggest that high-rank may provide long-term health and energetic benefits for female C. mitis, with potential fitness implications.

Suffixation influences receivers' behaviour in non-human primates.

Compared to humans, non-human primates have very little control over their vocal production. Nonetheless, some primates produce various call combinations, which may partially offset their lack of acoustic flexibility. A relevant example is male Campbell's monkeys (Cercopithecus campbelli), which give one call type ('Krak') to leopards, while the suffixed version of the same call stem ('Krak-oo') is given to unspecific danger. To test whether recipients attend to this suffixation pattern, we carried out a playback experiment in which we broadcast naturally and artificially modified suffixed and unsuffixed 'Krak' calls of male Campbell's monkeys to 42 wild groups of Diana monkeys (Cercopithecus diana diana). The two species form mixed-species groups and respond to each other's vocalizations. We analysed the vocal response of male and female Diana monkeys and overall found significantly stronger vocal responses to unsuffixed (leopard) than suffixed (unspecific danger) calls. Although the acoustic structure of the 'Krak' stem of the calls has some additional effects, subject responses were mainly determined by the presence or the absence of the suffix. This study indicates that suffixation is an evolved function in primate communication in contexts where adaptive responses are particularly important.

Sniffing behaviours in guenons.

In Old World monkeys (OWM), the olfactory sense is thought to be less important than in other primate taxa. However, during the last decade experimental studies have shown that OWM possess much better olfactory capabilities than suspected. Here, we investigate for the very first time sniffing behaviours in three guenon species (Diana, de Brazza's and owl-faced monkeys) held in the Leipzig Zoo, Germany. We recorded frequencies and contexts of sniffing. The sniff index was used to allow comparisons across species. It was found that individuals sniffed 6.12 ± 7.69 times per hour. Most sniffing occurred in food-related contexts. Furthermore, Diana monkeys sniffed less often than de Brazza's and owl-faced monkeys. Thus, natural selection may drive the differences in using the olfactory sense in closely related primate species.

The vocal repertoire of adult male blue monkeys (Cercopithecus mitis stulmanni): a quantitative analysis of acoustic structure.

Vocal signals are key elements in understanding species' behavior, sociality, and evolution. Quantified repertoires serve as foundations for investigating usage and function of particular signals, and also provide a basis for comparative analyses among individuals, populations, and taxa to explore how entire signal systems evolve. This study presents a descriptive catalogue of all vocal signals used by adult male blue monkeys (Cercopithecus mitis stuhlmanni). During 12 months in the Kakamega Forest, Kenya, I observed and digitally recorded vocal behavior of 32 adult males across a variety of socioecological contexts. From recordings, I measured 18 temporal-frequency parameters. Undirected ordination and hierarchical cluster analysis identified six distinct call types regularly used by males: ant, boom, ka, katrain, nasal scream, and pyow. Cross-validated discriminant function analysis supported the classifications. The repertoire is best described as discrete, though some gradation occurs between pyows and ants. Summary of acoustic structure and exemplar spectrograms are provided for each call type, along with preliminary examination of socioecological contexts in which they were produced. Discussion addresses repertoire structure, similarity to other taxa, and potential for functional inferences.

Biogeographic variation in the diet and behaviour of Cercopithecus mitis.

Primate species are characterised by variation in foraging behaviour and dietary composition across their geographic range. Here we examine how ecological conditions account for variation in the behavioural ecology of a widespread arboreal guenon, Cercopithecus mitis. Although substantial variation existed in time budgets, group size, home range and day journey length, clear biogeographic patterns were not apparent. In contrast, dietary variation was correlated with underlying climatic conditions. Temperature seasonality, which tends to increase with latitude, was significantly positively related to the proportion of fruit in the diet and negatively related to the proportion of animal matter. Both dietary components were 'preferred' foods, with the variability between populations reflecting the availability of different food types across their geographic range. Although we found no significant relationships between climate and the proportion of leaves in the diet, the ability for C. mitis to vary its diet to include a diversity of food types, and to incorporate a significant proportion of leaves when preferred sources are scarce, likely underpins its ability to survive across such a large distribution.