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.     

Diet for a small primate: Insectivory and gummivory in the (large) patas monkey (Erythrocebus patas pyrrhonotus)

A 17 month field study of unprovisioned patas monkeys (Erythrocebus patas pyrrhonotus) in Laikipia, Kenya, using both ad libitum and scan sampling techniques, revealed that the diet of patas monkeys consists primarily of gum of Acacia drepanolobium, arthropods (both free-living and concentrated in the swollen thorns of A. Drepanolobium), and other animals. This type of diet is normally found only in smaller-bodied primates. Results from vegetational transects suggest that the larger-bodied patas monkey can subsist on such a diet because gum and arthropods are relatively easily found in their habitat, thereby minimizing search time. Patas monkeys also spend more time moving and less time feeding (while not moving) than other Old World primates. The characteristic long limbs of patas may have evolved in response to feeding on small, nonusurpable, and widely distributed foods, in which access to foods is maximized while time and energy spent in terrestrial travel between food sites are minimized. Am. J. Primatol. 45:381–398, 1998. © 1998 Wiley-Liss, Inc.     

The ontogeny of sex differences in the behavior of patas monkeys

Many studies of sex differences in primates have been based on small experimental groups of peers in which only a limited range of social behavior could be expressed. In addition, the first few months of life are often the focus of such studies, with relatively little attention paid to older juveniles. In this study, 11 male and 9 female juvenile patas monkeys, living in a captive social group with all age-sex classes available, were observed between 1 and 4 years of age. A subset of seven patas monkeys was also observed between birth and 1 year of age. Here, we report the development of sex differences in independence, play, grooming, positioning behavior, and aggression over the juvenile period. Juvenile male patas monkeys played more and in longer bouts than females, but wrestling (rough-and-tumble play) was not more common among males. There were few differences in behaviors directed to male and female juveniles by other group members. Distinct differences emerged only in the behaviors of the juveniles themselves, with females being more active participants in social and aggressive interactions than males. In general, sex differences in patas monkeys show a mixture of patterns, some of which are predictive of adult sex differences and some of which appear to be specific to the particular demands of the juvenile period in this species     

Blood values of free-ranging patas monkeys (Erythrocebus patas)

Free-ranging patas monkeys (Erythrocebus patas) from El Guayacán island, Puerto Rico, were surveyed to establish values for the hemogram, serum biochemicals, calcium, and phosphorus. Results were tabulated for males and nonpregnant/nonlactating, pregnant, and lactating females. A summary of blood values from previous studies on captive patas monkeys was also tabulated for comparison.     

The ecology of the introduced patas monkey (Erythrocebus patas) population of Southwestern Puerto Rico

This paper presents the results of a study on the introduced, free-ranging patas monkey population of Southwestern Puerto Rico (SWPR). It describes information on the population size, social group composition, diet, daily ranging patterns, and patas home range during a 3 year period. The patas monkey population in the study area consisted of approximately 120 individuals in four heterosexual groups and several all-male bands. Within their home ranges (26.8 km²), the population density was 4.47 individuals/km². Home range size among the population's four heterosexual groups varied from 3.72 km² to 15.39 km², and minimum daily distance traveled ranged from 0.8–2.0 km. In general, the social structure and mating system of this population parallels what has been described for African populations. However, habitat use, ranging behavior, and the quality of intergroup interactions suggests that patas of this population exhibit territorial behavior. Am. J. Primatol. 45:351–365, 1998. © 1998 Wiley-Liss, Inc.     

Comparison of responses to alarm calls by patas (Erythrocebus patas) and vervet (Cercopithecus aethiops) monkeys in relation to habitat structure

We studied responses to alarm calls of sympatric patas (Erythrocebus patas) and vervet (Cercopithecus aethiops) monkeys in relation to habitat structure, with the intention of understanding the relationship between the environment and predator avoidance. Patas and vervet monkeys are phylogenetically closely related and overlap in body size. However, while patas monkeys are restricted to nonriverine habitats at our study site, vervets use both nonriverine and riverine habitats, allowing us to "vary" habitat structure while controlling for effects of group size, composition, and phylogeny. Patas monkeys in the nonriverine habitat responded to mammalian predator alarm calls with a greater variety of responses than did vervets in the riverine habitat, but not when compared with vervets in the nonriverine habitat. Ecological measurements confirm subjective assessments that trees in the riverine habitat are significantly taller and occur at lower densities than trees in the nonriverine habitat. Despite the lower density of trees in the riverine habitat, locomotor behavior of focal animals indicates that canopy cover is significantly greater in the riverine than the nonriverine habitat. Differences in responses to alarm calls by the same groups of vervets in different habitat types, and convergence of vervets with patas in the same habitat type, suggest that habitat type can be a significant source of variation in antipredator behavior of primates.     

Microhabitat preference and vertical use of space by patas monkeys (Erythrocebus patas) in relation to predation risk and habitat structure

Habitat structure can be important in determining habitat preference of animals because it is often closely linked to factors that affect survival and reproduction, such as food availability and predation risk. Here we examine the ways in which microhabitat structure and predation risk affect the habitat preference of wild patas monkeys (Erythrocebus patas). Patas monkeys in Kenya are typically restricted to Acacia drepanolobium habitat, but within our study group's home range, there are two distinct microhabitats, one with taller trees ('tall microhabitat') and one with apparently perennially shorter trees ('short microhabitat'). Examination of ranging behavior indicates that the patas monkeys preferred the tall microhabitat. In the tall microhabitat, focal animals climbed into trees that were significantly taller than average, indicating that they preferred tall trees. Female patas monkeys spent more time scanning from tall trees than from short trees and detected predators only from taller than average trees, based on alarm call data. Their use of tall trees may have decreased their predation risk by increasing their ability to detect predators. We found no evidence of increased food availability or reduced predator presence in the tall microhabitat that could contribute to the monkeys' preference for the tall microhabitat.     

Comments on oral grooming by patas monkeys,including a comparison with rhesus macaques

Observations of the motor patterns used by patas monkeys during allogrooming indicate that this species uses oral movements much more than previously believed. Compared to rhesus macaques, patas show more mouthpicks and licks, and fewer handpicks. Despite behavioral and anatomical evidence for good precision gripping, patas usually remove debris from a partner's fur orally rather than manually.     

Life-history parameters of a wild group of West African patas monkeys (Erythrocebus patas patas)

Based on long-term, although intermittent, observations (2 years 4 months of 14 years), we present data on birth seasonality, age at first birth, interbirth intervals, mortality rates, age at first emigration, and population change of a wild population of West African patas monkeys (Etythrocebus patas patas) in northern Cameroon. Birth season was from the end of December until the middle of February, corresponding to the mid-dry season. In spite of large body size, the patas females had the earliest age at first birth (36.5 monthsold) and the shortest interbirth intervals (12 months) compared to the closely related wild forest guenons. Age at first emigration of the males was considered to occur between 2.5 and 4.5 years. The group size of the focal group drastically decreased between 1984 and 1987, and steadily increased until 1994, then decreased again in 1997. The neighboring group also showed a similar trend in group size. The population decreases were likely to be caused by drought over 3 years. Annual crude adult mortality rate was 4% during population increase periods (PIP) between 1987 and 1994. It rose to 22% during all the periods (AP), including drought over 3 years. Despite their smaller body size, the rate of the wild forest guenons (Cercopithecus mitis) (4%) was the same and much lower than those of the patas during PIP and AP, respectively. The annual average juvenile mortality rate was 13% during PIP and it also rose to 37% during AP. That of wild forest guenons (C. ascanius) (10–12%) was a little lower and much lower than those of the patas during PIP and AP, respectively. These findings were consistent with Charnov's theoretical model of mammalian life-history evolution in that patas with high adult and juvenile mortality showed early and frequent reproduction in spite of large body size. Charnov also considered high adult mortality as a selective force and high juvenile mortality as a density-dependent consequence of high fecundity. Our results support the former but not the latter research findings.     

Patterns of mating among male patas monkeys (Erythrocebus patas) in Kenya

An habituated group of wild patas monkeys was observed in Kenya for 550 h in 1984. Observations were made primarily during an interval that, as previous studies at the same site had demonstrated, coincided with the annual mating and conception periods. Earlier field studies of patas at other sites had reported that heterosexual patas groups had only a single resident adult male and that mating was harem-polygynous. At the Kenya site, by contrast, as many as six males were simultaneously resident and mated in the group during the conception period. Males adopted a variety of tactics to gain access to receptive females, ranging from opportunistic mating to attempts at sequestration that resembled consort behavior in other cercopithecoids such as savanna baboons and rhesus macaques. Aggressive competition for access to females took place among the males, although the number of completed copulations per male did not bear a positive relation to agonistic dominance rank. For patas monkeys, harem polygyny is only one available option within an overall mating system that is best described as a form of promiscuous polygyny, especially during periods when conception is most likely.     

Interspecific and temporal variation of ant species within Acacia drepanolobium ant domatia, a staple food of patas monkeys (Erythrocebus patas) in Laikipia, Kenya

The ants that live in the swollen thorns (domatia) of Acacia drepanolobium are staple foods for patas monkeys (Erythrocebus patas). To obtain a better understanding of these insects as resources for patas monkeys, we sampled the contents of 1,051 swollen thorns (ant domatia) over a 22-month period from December 1999 to September 2001, in Laikipia, Kenya. First, we confirmed that of the four species of ants that live on A. drepanolobium, Crematogaster sjostedti, the competitively dominant ant in this system, does not rear significant brood in the swollen thorns and is therefore not a major food item of patas monkeys. Second, across the other three species that do use swollen thorns for rearing their brood, C. nigriceps, C. mimosae, and Tetraponera penzigi, the number of worker ants per swollen thorn increased with increasing competitive dominance. Third, although there was considerable month-to-month variation in the number of workers, immatures, and especially alates (winged reproductives) within species, there was less variation across species because ant production was asynchronous. Variation in domatia contents was poorly related to rainfall for each of the three species. Finally, distal thorns held more alates and fewer workers than interior thorns, and branches higher off the ground held more alates and more workers than lower branches. For the numerically dominant C. mimosae, higher branches held significantly more immature ants than did lower branches. Ants are reliable food resources for patas monkeys, and are probably more reliable than many plant resources in this highly seasonal environment. We estimate that patas monkeys may get as much as a third of their daily caloric needs from these ants year-round. As ants and other insects are widely consumed by primates, we suggest that greater consideration be given to species differences in animal food choices and that further studies be conducted to examine the degree to which ants influence energy intake and reproduction in other primates.     

Allomaternal behavior in a group of free-ranging patas monkeys

A free-ranging group of patas monkeys (Erythrocebus patas) containing 18 individually identifiable adult females was observed for approximately 400 hours, equally distributed over two two-month periods corresponding to the breeding and birth seasons, at the La Parguera facility of the Caribbean Primate Research Center. The large number of infants born in the spring and early summer (n = 14) allowed for detailed observations of alloparental behaviors, with a focus on allomothering by the adult females. Seven types of alloparental behaviors were recorded: contact, nuzzling, grooming, agonism, close visual inspection, attempted kidnapping, and kidnapping. Adult females emitted the vast majority of allomaternal behavior, the patterning and frequency of which closely resembled the patterning and frequency of inter-adult female social grooming. Relative dominance status of the participants did not consistently predict the directionality of allomothering. The most commonly observed allomaternal behaviors were contact and nuzzling, which are primarily affiliative behaviors; agonism was rare. Successful kidnapping occurred eight times. Immature monkeys (n = 22) emitted an additional 32 alloparental acts. A propensity towards allomothering by experienced females would be most beneficial to patas infants due to the patas' tendency towards dispersed foraging and rapid flight in the presence of danger. It is possible that direct competition with groups of rhesus macaques for available resources on the island served as a proximal cause for the allomothering observed in this patas group.     

Comparing measures of travel distances in primates: Methodological considerations and socioecological implications

Travel costs can influence numerous aspects of the lives of primates, including net energy balance (and therefore reproductive success of females) and maximum group size. Despite their potential impact, there has been no systematic comparison of different measures of travel distance. We compared three measures of travel distance in 30 min (actual distance of individuals, straight‐line distance of individuals, and straight‐line distance of groups) and their ratios in a small group and a large group of vervet monkeys (Cercopithecus aethiops) and between the large group of vervets and a group of patas monkeys (Erythrocebus patas) of roughly similar size. The large group of vervets traveled farther than the small group regardless of the measure used, but the ratios of the different measures were not significantly different between those groups. Patas monkeys traveled significantly farther than the large group of vervets regardless of the measure used. In both vervets and patas, straight‐line distances of individuals (ISLD) and groups (GSLD) underestimated actual distances traveled by individuals (IAD), but the degree to which they did so differed between species. IAD is more accurate than the other two measures and is preferred for studies of energetics and individual reproductive success, although ISLD or GSLD may be substituted when the ratios of IAD/ISLD or IAD/GSLD do not differ between groups or species. The ratio of IAD/ISLD was larger in vervets than in patas, suggesting that individual vervets meander more over short periods of time than patas. The ratio of ISLD/GSLD was larger in patas than in vervets, suggesting that patas move at angles or across the group's center‐of‐mass whereas vervets move more consistently along with others in their group. This has implications for the formation of spatial subgroups and alliances within groups. Am. J. Primatol. 48:87–98, 1999. © 1999 Wiley‐Liss, Inc.     

Passive joint mobility in patas monkeys (Erythrocebus patas): rehabilitation of caged animals after release into a free-ranging environment

A previous study of passive joint mobility in patas monkeys (Erythrocebus patas) showed that laboratory-caged animals had significantly greater mobility in most joints than age/sex matched free-ranging monkeys. Passive joint mobility on 27 of the same animals was measured 6 months after the caged animals were released onto a 40-hectare island. The results show that within 6 months of becoming free-ranging, typical passive joint mobility is restored. Thus, although caging directly affects measurements of morphologically-determined features in patas monkeys, confinement itself does not necessarily prevent rehabilitation if the immature monkeys are released into a free-ranging environment.     

Differential habitat utilization by patas monkeys (Erythrocebus patas) and tantalus monkeys (Cercopithecus aethiops tantalus) living sympatrically in northern Cameroon.

In order to obtain reliable evidence for differences in habitat preferences between two closely related savanna-dwelling primate species, namely, patas monkeys (Erythrocebus patas) and tantalus monkeys (Cercopithecus aethiops tantalus), I collected data on vegetation and patterns of range use concurrently at a single study site, Kala Maloue, Cameroon, in a similar manner for a group of each species. Kala Maloue consisted of 64% grassland mostly dominated by Gramineae spp. and the rest was woodland. Tantalus monkeys showed preference for woodland, especially gallery forest, much more than did the patas irrespective of the season. Moreover, patas preferentially established their home range in grassland in the wet season. Interspecific and seasonal differences in habitat preferences could be interpreted on the basis of interspecific and seasonal differences in preferences for main food. In dry season, tantalus utilized water-containing areas at a frequency closely in proportion to the availability of such areas while the patas utilized water-containing areas more frequently than expected. This is because tantalus established a smaller home range along the river where water was never completely depleted throughout the dry season. Both the patas and the tantalus preferred woodland to grassland as sleeping sites possibly owing to predation avoidance. Both the daily travel distance per group weight and the home range size per group weight were greater for patas than for tantalus partly because of higher preference for grassland with low habitat quality in the case of patas. It is suggested, however, that high locomotive ability enabled patas to effectively utilize small and widely dispersed items of food such as grasshoppers and to explore areas with high availability of food and water and with preferable sleeping sites.     

Social and environmental determinants of reproductive cycles in patas monkeys

The menstrual cycles of a captive group of patas monkeys were followed for 15 months by taking vaginal smears and lavages three times a week. Without an adult male in the group, menstrual cycles still showed the expected qualitative changes previously associated with the onset and with the end of a mating period. The addition of an adult male to the female group, once mating season cycles were evident, did not result in further changes in erythrocytes or sediment levels in vaginal samples or cycle regularity. Menstrual-cycle onsets for related females were significantly more synchronized than onsets for unrelated females. Preliminary observations on adult male patas housed separately from the females indicate that they too undergo seasonal changes in physiology and behavior.     

Male Competition,Mating Success and Female Choice in a Seasonally Breeding Primate (Erythrocebus patas)

Mammals with restricted breeding seasons often show brief but intense bouts of male competition for mates and male reproductive success has been attributed to male competitive abilities, with the most aggressively successful males able to control access to fertile females, or with females choosing to mate with such males. We studied male competition, mating behaviour and female mate selection in patas monkeys, a primate with a restricted breeding period. We observed two habituated patas groups in Laikipia District, Kenya, during Jun.-Aug. 1983. During the study, one group had a single resident male while the other group had multiple adult males. Within the multimale group, experienced adult males were no more successful than the subadult male. The sole resident male had a significantly higher rate of fights won (p < 0.02) although he did not differ from the multimale group males in rate of aggression or initiation of fights. We found no significant differences in either mating success or female preference based on males' experience or residency. The rates at which males copulated with and were solicited by females were not significantly correlated. We found no evidence of stable dominance ranks among males in the multimale group and aggressive success was not significantly correlated with copulation rate for males in the multimale group. Subadult males were responsible for the majority of copulations observed during the final third of the breeding season. Our observations of this patas population showed a fluid number of males in groups, with the same groups able to shift rapidly from single to multimale structure. This fluidity may result from the large fluctuations in numbers of breeding-age males and females observed over 4 yr of studying this population.     

Patas monkey copulations: One mount,repeat if necessary

Early, anecdotal reports of patas monkey (Erythrocebus patas) sexual behavior suggested that these primates were series-mount copulators. In 1975, limited evidence was presented in favor of single-mount copulations as the species-typical pattern for patas. The situation remained confused, however, as indicated by statements in a recent review article. This paper adds to our knowledge of patas sexuality by presenting quantitative data on the copulatory behavior of two additional adult males. Both males usually gave pelvic thrusts to intravaginal ejaculation once they had mounted a female and gained intromission. Mounts without ejaculation were clearly failed attempts at copulation, rather than segments of a stereotyped series-mount pattern. All of the available quantitative data indicate that the species-typical mating pattern for patas monkeys is copulation in a single mount.     

Longitudinal study of dominance relations among captive patas monkeys

Eight and a half years of dominance relations within a captive group of patas monkeys were analyzed. It was found that matrilineal kinship significantly influenced individuals' ranks. In contrast, with the exception of certain intramatriline changes, increasing age had no predictable effect on overall rank, at least for females (this was untestable for males). Offspring typically challenged maternal dominance and in eight of twelve dyads, offspring either rose fully over their mothers (three cases, all daughters) or at least achieved dominance ambiguity with them. Additionally, two of the four younger sisters with an opportunity to rise in rank over an older sister did so. The group dominance hierarchy was unstable for 75% of the study due to a combination of agonistically induced and demographically induced rank changes. Concentration of the highest ranks in a single matriline showed a stronger association with group hierarchy stability than did the presence of an adult, nonnatal male. Group hierarchy stability was associated with increased affiliation (sitting close and sitting touching), but otherwise there were no behavioral correlations. Individuals' ranks within the group hierarchy were unrelated to their chances of being wounded or having diarrhea. Adult females' ranks were over twice as stable as the group hierarchy (57.1% stability), but stability/instability was not correlated with any behavioral changes. Available evidence suggests that dominance relations play only a minor role in the organization of patas monkeys' intragroup behavior. Am. J. Primatol. 42:41–51, 1997. © 1997 Wiley-Liss, Inc.     

Diurnal births and perinatal behavior among wild patas monkeys: Evidence of an adaptive pattern

Data from a 2-year field study of patas monkeys (Erythrocebus patas) in Kenya support our earlier suggestion that diurnal births are a species-typical pattern of patas. In this respect patas are very unusual, as all existing information shows that nocturnal births are typical of both captive and freeranging monkeys. Patas do not give birth at night because to do so would render ineffective their night-resting strategy which reduces vulnerability to predation at night. Giving birth during the day, however, does not eliminate the risk of being preyed on; nor are all times of day equally favorable for giving birth. Our field data suggest that a patas female gives birth at those times of day when she is least likely to lose contact with her group or to encounter predators.