Locomotor activity differences between sympatric patas monkeys (Erythrocebus patas) and vervet monkeys (Cercopithecus aethiops): Implications for the evolution of long hindlimb length in Homo

Homo erectus is notable for its taller stature and longer lower limbs relative to earlier hominids, but the selective pressures favoring such long limbs are unclear. Among anthropoid primates, patas monkeys (Erythrocebus patas) and extant hominids share several extreme characteristics involved with foraging and movement, including the relatively longest lower limb proportions, longest daily travel distances and largest home ranges for their body or group size, occupancy of some of the driest habitats, and very efficient thermoregulatory systems. We suggest that patas monkeys are an appropriate behavioral model with which to speculate on the selective pressures that might have operated on H. erectus to increase lower limb length. Here, in a comparison of the locomotor activities of patas monkeys and sympatric, closely related vervet monkeys (Cercopithecus aethiops), we provide evidence for the hypothesis that patas use their long stride more to increase foraging efficiency while walking than to run, either from predators or otherwise. Am J Phys Anthropol 105:199–207, 1998. © 1998 Wiley-Liss, Inc.     

Rank Differences in Ecological Behavior: A Comparative Study of Patas Monkeys (Erythrocebus patas) and Vervets (Cercopithecus aethiops)

One of the central dichotomies in primate behavior is between species in which there are relationships among females that include stable dominance relationships, and those in which the relationships include weak or unstable dominance relationships. This dichotomy has been attributed to differences in food resources, with stable dominance hierarchies occurring in species that feed on usurpable foods. We compared rank-related differences in nonagonistic behaviors considered to be tightly linked to ecology in broadly sympatric vervets (Cercopithecus aethiops) and patas monkeys (Erythrocebus patas), two closely related cercopithecines that are exemplars of this dichotomy, with the expectation that vervets would exhibit stronger rank differences than patas monkeys in these behaviors. Overall, rank explained more than twice as much variation among vervets as among patas monkeys in ranging behavior, activity budgets, and diet. Vervets did not, however, exhibit stronger rank differences when they used Acacia xanthophloea habitat, in which foods are more usurpable, compared to Acacia drepanolobium habitat, in which foods are less usurpable. In Acacia drepanolobium habitat, to which patas are restricted, higher-ranking vervets converged in behavior with patas monkeys to a greater extent than lower-ranking vervets, suggesting that social constraints interfere with the foraging efficiency of lower-ranking vervets even in habitats in which there are fewer opportunities to usurp foods.     

Demography and Life Histories of Sympatric Patas Monkeys, <Emphasis Type="Italic">Erythrocebus patas</Emphasis>, and Vervets, <Emphasis Type="Italic">Cercopithecus aethiops</Emphasis>, in Laikipia,Kenya

Mortality patterns are thought to be strong selective forces on life history traits, with high adult mortality and low immature mortality favoring early and rapid reproduction. Patas monkeys (Erythrocebus patas) have the highest potential rates of population increase for their body size of any haplorhine primate because they reproduce both earlier and more often. We report here 10 yr of comparative demographic data on a population of patas monkeys and a sympatric population of vervet monkeys (Cercopithecus aethiops), a closely related species differing in aspects of social system, ecology, and life history. The data reveal that 1) adult female patas monkeys have significantly higher mortality than adult female vervets; 2) infant mortality in patas monkeys is relatively low compared to the norm for mammals because it is not significantly different from that of adult female patas monkeys; and 3) infant mortality is significantly higher than adult female mortality in vervets. For both species, much of the mortality could be attributed to predation. An epidemic illness was also a major contributor to the mortality of adult female patas monkeys whereas chronic exposure to pathogens in a cold and damp microenvironment may have contributed to the mortality of infant vervets. Both populations experienced large fluctuations during the study period. Our results support the prediction from demographic models of life history evolution that high adult mortality relative to immature mortality selects for early maturation.     

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.     

A case of patas-vervet hybrid in captivity

A female patas monkey (Erythrocebus patas) housed with two males of different species, Hamadryas baboon (Papio hamadryas), and vervet monkey (Cercopithecus aethiops pigerythrus), gave birth to a female offspring in a semi-open enclosure on June 11, 1975. Karyological analyses and electrophoretic examinations were carried out in order to determine the real father of the hybrid. From the results of these observations it was concluded that the father of the hybrid individual wasCercopithecus aethiops. Additionally, somatometric and hematological investigations were performed.     

An assessment of dominance and kinship among patas monkeys

Kaplan andZucker (1980) argued that dominance and kinship do not function as important organizing features for intragroup behavior and social structure among patas monkeys (Erythrocebus patas). This paper reviews the available data pertinent to this argument and concludes that dominance probably is not a reliable structural variable for captive patas, despite its clear development in most groups. In contrast, kinship is a major organizing feature that strongly affects allogrooming and other affiliative interactions, and socialization.     

Difference in food selection between patas monkeys (Erythrocebus patas) and tantalus monkeys (<Emphasis Type="Italic">Cercopithecus aethiops tantalus</Emphasis>) in Kala Maloue National Park,Cameroon, in relation to nutrient content

Phytochemical or nutrient analyses of primate diets have revealed clues to their food selection in a single species. On the other hand, few interspecific comparisons of phytochemical or nutrient composition of primate diets have been made, although diets are considered to differ in phytochemical or nutrient content from primate species to species, since different species have different body weights and different morphological and physiological characteristics. I compared the nutrient content of diet between patas monkeys (Erythrocebus patas) and tantalus monkeys (Cercopithecus aethiops tantalus) living sympatrically in Cameroon. Patas subsisted on a smaller number of food items, most of which were also tantalus food items. Then, I compared the protein–fiber ratio and the available energy content of the food items eaten by patas (patas foods) with those items eaten only by tantalus (tantalus foods). Both variables were higher in patas than tantalus foods, although there was no significant difference in available energy of plant foods. Next, when I performed discriminant analysis for patas foods and tantalus foods, employing the above two variables, a discriminant function with positive coefficients for both variables was obtained. The mean discriminant-function score of patas foods was higher than that of tantalus foods. Despite being somewhat larger in weight, patas selectively fed on a smaller number of foods of higher quality than did tantalus. I discuss why the results are inconsistent with a well known body weight–diet relationship (Jarman–Bell principle). Energy-efficient locomotion enables patas to exploit not only small dispersed food items of high quality but also areas where high-quality foods are distributed in clumps. Electronic Publication     

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.     

Reconciliation following aggression in patas monkeys,Erythrocebus patas

Following intra-group aggression, obvious conciliatory displays are absent from the behavioural repertoire of patas monkeys, Erythrocebus patas, while many other Old World primate species show special reconciliation gestures. When 10-min focal-animal samples that began after spontaneous aggression were compared with matched-control samples, captive adult female patas followed up on aggressive interactions, interacting sooner and more often with former opponents during post-conflict observations than during matched-control observations. Almost one-third of post-conflict observations included affiliative behaviour between former opponents, which is termed reconciliation. Matrilineally related opponents were more likely to reconcile with one another than were unrelated animals. No effect of the dominance hierarchy on tendency to reconcile was found. Thus, patas monkeys showed general patterns in post-conflict behaviour that were similar to those seen in other primates previously investigated.     

Observations of parturition among captive patas monkeys (Erythrocebus patas)

Six cases of labor and delivery by laboratory-housed patas monkeys (Erythrocebus patas) are described. Five of the births occurred during the daytime, and the sixth occurred just after the usual “lights out” time. Although it was impossible to determine the actual onset of labor, the final 1.5–0.5 hr before parturition were observed in all cases. Throughout labor, females tended to begin a contraction every 2–4 min and contractions usually lasted 35–60 sec. Females did not show common patterns of change in contraction frequency or duration as birth neared. It was not possible to describe patas monkey labor and delivery in terms of “stages.” Postpartum, females typically began cleaning their infants within 90 sec and picked them up within 5 min. Placental delivery times varied among females, as did the extent of placental consumption. One “mother-only” - reared female proved to be an inadequate mother.     

Nutritional benefits of crematogaster mimosae ants and acacia drepanolobium gum for patas monkeys and vervets in laikipia,Kenya

Patas monkeys (Erythrocebus patas) are midsized primates that feed extensively on the gum of Acacia drepanolobium and the ants are housed in swollen thorns of this Acacia. Their diet resembles that expected more of smaller bodied primates. Patas monkeys are also more like smaller bodied primates in reproducing at high rates. We sought to better understand the convergence of patas monkeys with smaller bodied primates by comparing their feeding behavior on ants and gum with that of closely related, sympatric vervets (Chlorocebus pygerythrus), and analyzing the nutrient content of the gum of A. drepanolobium and of Crematogaster mimosae, the most common ant species eaten by patas monkeys in Laikipia, Kenya. All occurrences of feeding and moving during focal animal sampling revealed that 1) patas monkeys seek A. drepanolobium gum but vervets avoid it; 2) both species open swollen thorns most often in the morning when antsare less active; 3) patas monkeys continually feed onswollen thorns and gum while moving quickly throughout the day, whereas vervets reduce their consumption of these items and their travel rate at mid‐day, and; 4) vervets eat young swollen thorns at a higher rate than patas monkeys. Patas monkeys are able to spend little time acquiring substantial amounts of energy, protein, and minerals from A. drepanolobium gum and C. mimosae ants each day. These findings, when coupled with evidence of causes of infant and adult female mortality, suggest that reproductive success of female patas monkeys is more immediately affected by illness, disease, interactions between adults and infants, and access to water than by food. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

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.     

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.     

Seasonal, sex, and interspecific differences in activity time budgets and diets of patas monkeys (Erythrocebus patas) and tantalus monkeys (Cercopithecus aethiops tantalus), living sympatrically in northern Cameroon

I examined seasonal, sex, and interspecific differences in activity time budgest and diets of patas (Erythrocebus patas) and sympatric tantalus monkeys (Cercopithecus aethiops tantalus) on the basis of 5-day data sets collected in three and two different seasons, respectively, by the method of focal animal sampling. The seasons included species-specific mating and birth seasons. As compared with not only the birth season but also conspecific females, both patas resident male and tantalus male spent less time feeding and more time resting, day and night, in their respective mating seasons. Given that day-resting time includes time for vigilance for non-resident males and receptive females, this may reflect that males should minimize time spent feeding to allow maximum participation in other fitness-increasing activities such as mating-relating activities asSchoener (1971) predicted. In both species, the males consumed fruits containing less protein but more calories and showed a high feeding rate to compensate for the shorter time spent feeding in the mating season. In contrast, females consumed protein-rich food types (i.e. animals, protein-rich seeds, leaves, and flowers) in the birth season to meet the high demand for protein due to pregnancy and lactation. Given that the season for males was considered to be not a calendar but a reproductive “season” (i.e. mating or birth season), both sexes of patas spent more time moving and less time day- and night-resting than did the tantalus counterparts irrespective of the “season”. Patas subsisted on fruits, gums, and supplementarily lipid-rich seeds as an energy source and animal matters and protein-rich seeds as a protein source. In contrast, tantalus subsisted on fruits and lipid-rich seeds as energy and flowers and leaves as protein.     

Schistosoma intercalatum Fisher, 1934 (Cameroon) infection in the patas monkey (Erythrocebus patas Schreber, 1775).

Kuntz R. E., McCullough B., Huang T. C. and Moore J. A. 1978. Schistosoma intercalatum Fisher, 1934 (Cameroon) infection in the patas monkey (Erythrocebus patas Schreber, 1775). International Journal for Parasitology8: 65–68. Patas monkeys (Erythrocebus patas) have been infected with Schistosoma intercalatum, a schistosome of increasing concern in Africa, to study definitive host-parasite relationships. A high compatibility for parasitism in the S. intercalatum-patas monkey system was demonstrated suggesting this combination for long-term investigations in which minimal parasite destruction by the host would be expected. Egg production by S. intercalatum was high. Most eggs are deposited in the large intestine, but they elicit only slight pathology. No pathologic involvement of the urinary bladder was observed, in contrast to previous investigations in which other species of nonhuman primates infected with S. intercalatum developed extensive pathology.     

Banding patterns of the chromosomes of Erythrocebus patas (Schreber 1774) compared to other primate species

The karyotype of Erythrocebus patas was studied using G, Q and C bands and NOR staining and compared to those of Miopithecus talapoin, Macaca mulatta and Cercopithecus aethiops. The comparison shows a large number of homoeologies, but several chromosome pairs cannot be matched. The mechanisms that may have originated the karyotypes of these species are discussed.     

Differences Between Vervets (Cercopithecus aethiops) and Patas Monkeys (Erythrocebus patas) in Agonistic Interactions Between Adult Females

We examined agonistic interactions between adult females in wild, unprovisioned patas monkeys (Erythrocebus patas) and vervets (Cercopithecus aethiops). The dominance hierarchy of patas is far less clear than that of vervets. Patas had fewer interactions per dyad, fewer dyads with interactions, and a high percentage (18%) of reversals in which lower-ranking females won in agonistic interactions with higher-ranking females. Although the rank ordering of the kinds of interactions patas and vervets displayed is similar, with avoidance being the most frequently observed agonistic response to approaches by other females, patas were chased and supplanted more often than vervets were. The resources over which females were supplanted also differ between species. Supplants over food comprise smaller proportion of total supplants patas than for vervets. Patas appear to feed on less usurpable foods than vervets. We conclude that (1) Erythrocebus and Cercopithecus spp., except C. aethiops, should not be categorized with other Cercopithecinae, and C. aethiops should not be categorized with other Cercopithecus spp. and Erythrocebus, in discussions and analyses of relationships between females within groups and (2) ecological conditions, i.e., usurpability of foods, can override phylogenetic history as the selective pressure determining the nature of female competitive relationships within groups.     

Embryonic development in Erythrocebus patas

The developmental stages of 12 Erythrocebus patas embryos, ranging in gestational age from 30 to 50 days, is described. The pattern of embryogenesis in E. patas closely parallels the anatomic characteristics of human and other nonhuman primate embryos between stages 12 and 23. However, there is a delay in development in E. patas similar to that observed in human embryos which differs from the macaques and baboons. This temporal difference in the embryonic period is an important factor in the design and analysis of early pregnancy studies 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.