Showing papers in "International Journal of Primatology in 1998"

Dietary Response of Chimpanzees and Cercopithecines to Seasonal Variation in Fruit Abundance. II. Macronutrients

Abstract: In a continuation of our study of dietary differentiation among frugivorous primates with simple stomachs, we present the first comparison of differences in dietary macronutrient content between chimpanzees and cercopithecine monkeys Previously we have shown that chimpanzee and monkey diets differ markedly in plant part and species content We now examine whether this diet diversity is reflected in markedly different dietary macronutrient levels or the different feeding strategies yield the same macronutrient levels in their diets For each primate group we calculated the total weighted mean dietary content of 4 macronutrients: crude lipid (lipid), crude protein (CP), water-soluble carbohydrates (WSC), and total nonstructural carbohydrates (TNC) We also calculated 4 fiber fractions: neutral-detergent fiber (NDF), which includes the subfractions hemicellulose (HC), cellulose (Cs), and sulfuric acid lignin (Ls) The HC and Cs are potentially fermentable fibers and would contribute to the energy provided by plant food, depending on the hind gut fermenting capacity of the individual primate species The chimpanzee diet contained higher levels of WSC and TNC because during times of fruit abundance the chimpanzees took special advantage of ripe fruit, while the monkeys did not The monkey diets contained higher levels of CP because the monkeys consumed a constant amount of leaf throughout the year All four primate species consumed diets with similar NDF levels However, the chimpanzees also took advantage of periods of ripe fruit abundance to decrease their Ls levels and to increase their HC levels Conversely, the monkey diets maintained constant levels of the different fiber fractions thoughout the year Nevertheless, despite these differences, the diets of the 4 frugivores were surprisingly similar, considering the substantial differences in body size We conclude that the chimpanzee diet is of higher quality, particularly of lower fiber content, than expected on the basis of their body size

Changes in Orangutan Caloric Intake, Energy Balance, and Ketones in Response to Fluctuating Fruit Availability

Abstract: I used novel quantitative measures to assess the impact of fruit seasonality on changes in diet and energy balance during 4918 hours of observation on <60 orangutans in Gunung Palung National Park, West Kalimantan, Indonesia, from September 1994 through September 1995. During this period a mast fruiting occurred, resulting in large fluctuations in fruit availability, measured by monitoring of 558 orangutan fruit trees. Orangutan diet varied tremendously in accordance with these fluctuations. During the month of highest fruit production, 100% of the orangutan diet was composed of fruit. In contrast, during the fruit-poor period as little as 21% of the diet was fruit, while 37% was bark. Nutritional analyses of 78 of the foods most commonly eaten during this study show that mast foods were significantly higher in caloric content than were nonmast foods, which translates into substantial changes in caloric intake. During the month of highest fruit consumption (January), males consumed an estimated 8422 kcal/day and females consumed 7404 kcal/day. During the month of lowest fruit consumption (May), males consumed 3824 kcal/day and females consumed 1793 kcal/day. These differences were significantly different between the 2 months for both males and females. In May, males ate significantly more kcal/day than females did. Time spent feeding did not differ between the two periods for either males or females. I assessed the impact of these changes in diet and caloric intake on orangutan physiological functioning by measuring ketones in 257 urine samples collected from adult males and females. Ketones—products of fat metabolism—were present in urine only during the fruit-poor period. These data suggest that orangutans are highly efficient at storing fat during fruit-rich periods and utilizing the reserves during times of fruit shortfall. This response may have important implications for orangutan survivorship, reproduction, and behavior.

Conservation Biology of the Genus Alouatta

Abstract: As assessed by the IUCN Mace–Lande system, seven (35%) of the 20 Alouatta species and subspecies with adequate data are classified as “threatened,” i.e., critically endangered, endangered, or vulnerable (Rylands et al., 1995). This percentage is much lower than the 75 to 100% threatened taxa for the other large-bodied genera: Ateles, Lagothrix, and Brachyteles. Only 5 of the 16 Neotropical genera have lower percentages of threatened taxa than that of Alouatta: Cebuella, Pithecia, Saguinus, Saimiri, and Cebus. The threatened howler taxa occupy small distributions in areas of forest fragmentation. In general, populations are most affected by major habitat disturbance, such as total deforestation and flooding from dam construction, and by human hunters. Facilitated by their ability to exploit folivorous diets in small home ranges, howlers can tolerate considerable habitat fragmentation but not the increased exposure to hunting that may accompany it. Howlers seem particularly vulnerable to yellow fever and bot fly parasitism. Although the former threat may decrease by increasing fragmentation of the habitat, other sorts of parasitism may increase in disturbed habitats. The low genetic variability of the Central American howlers suggests a resistance to inbreeding depression potentially experienced during population bottlenecks. Greater between-population variability may still exist. Although howlers are not readily bred in captivity, they respond well to translocation. Translocation has been successfully achieved for ≥4 howler species and is a viable option for introducing new genetic variability into population fragments and repopulating areas from which howlers are extinct. Their pattern of bisexual dispersal facilitates colonization of regenerating habitats, and in suitable, protected habitats they have shown the capacity for strong population recovery.

Physiological ecology of howlers (Alouatta) : Energetic and digestive considerations and comparison with the Colobinae

Abstract: Remarkably little attention has been focused on the physiological ecology of free-ranging primates Yet without such information, it may prove difficult to advance our understanding of factors influencing the dietary behavior of wild primates much beyond its present state Mantled howlers (Alouatta palliata) have been studied in terms of some features of physiological ecology Results of this work have helped to clarify some factors influencing howler and other primate food choices in the natural environment and have called into question various assumptions about leaf-eating primates For example, though howlers eat considerable foliage, they do not exhibit a lower than predicted basal metabolic rate, nor do available data suggest that secondary compounds strongly influence howler food selection Comparison of howlers with members of the Colobinae reveals some differences in features of their respective energetic and digestive physiology and raises timely issues for future research

Social Organization and Range Use in the Yunnan Snub-Nosed Monkey Rhinopithecus bieti

Abstract: We studied social organization, behavior, and range use of the Yunnan snub-nosed monkey (Rhinopithecus bieti) at Wuyapiya (99°12′E, 28°30′N, the People's Republic of China) over 12 months between May 1992 and June 1994. The Wuyapiya band contained ≥175 members and had two levels of social organization. At one level, the monkeys formed multifemale, one-male units (OMUs) similar to those of many other colobines. At another level, 15 to 18 OMUs traveled together in a cohesive band. Unlike the bands of other species of Rhinopithecus, the Wuyapiya band of R. bieti did not show seasonal fission–fusion, although some social behavior, such as male–male aggression, was seasonal. With regard to range use, the Wuyapiya band had a large home range and long daily travel distances compared with other colobines. Minimum range size in 1 year at Wuyapiya is 16.25 km 2 , although there is no asymptote for range size as a function of observation time. Range size for the Wuyapiya band is 25.25 km 2 over the 2-year study and appeared to cover 100 km 2 between 1985 and 1994. The primary food of R. bieti at Wuyapiya is lichens, which are ubiquitous in fir trees. The multitiered social organization of R. bieti appears to result from the interaction of food resource characters with the forces of mate competition, with band sizes based on female responses to the spatial and temporal characteristics of lichens and subdivisions within bands based on male competition for mates.

Long-Term Habitat Use by Mountain Gorillas (Gorilla gorilla beringei). 1. Consistency, Variation, and Home Range Size and Stability

Abstract: Mountain gorillas are highly folivorous. Food is abundant and perennially available in much of their habitat. Still, limited research has shown that single gorilla groups heavily used areas where food biomass and quality were relatively high and where they met daily nutritional needs with relatively low foraging effort. Also, ecological factors influenced solitary males less than groups with females. Long-term data on habitat use by multiple mountain gorilla social units and more extensive data on variation in food distribution, presented here, confirm that food distribution influences areal occupation densities across groups and over time. These data also confirm the group/solitary male distinction and show that food distribution became more important for one male once he acquired females. Groups used ≤25 km2, and inter-annual home range and core area overlap was often low. Annual home range and core area size varied considerably within groups and across years. It bore no simple relationship to group size and estimated group biomass. Core areas were biased samples of total home ranges and were relatively good foraging areas. One group abruptly shifted its home range in response to male mating competition. Home ranges of two others expanded from 1981 to 1987, though at a decreasing rate. Data on one such group, which varied considerably in size, are consistent with arguments that costs of scramble competition are low except in unusually large groups. Low site fidelity, low scramble costs, and high home range overlap should decrease the ecological costs of female transfer.

Population Characteristics of Howlers: Ecological Conditions or Group History

Abstract: We examined the relative importance of ecological parameters—habitat productivity and seasonality—and group history—episodic predation, disease, and sudden habitat deterioration—to explain variation in the density and group structure of howlers (Alouatta spp.). We use data from a census of Guanacaste National Park, Costa Rica, and a literature review characterizing 80 howler populations. In Guanacaste National Park both habitat type and degree of protection affect howler density and group structure. Howlers were found at the highest density and in the largest groups in areas of semievergreen forest, which ecological sampling indicates have the most consistent level of food production. Differences in density between the sector of the park that first received protected status and more recently protected areas may be due partially to the degree of protection the areas received. We test the prediction that howler density and group structure would be influenced by habitat productivity as indexed by rainfall. Average group size and sex ratios differ among species, but female-to-immature ratios do not. Considering all censuses at one site to be independent, there are significant interspecific differences in density, with Alouatta pigra occurring at lower densities than the other species. In spite of such variability, there is no relationship between annual rainfall and howler density, and rainfall had a variable effect on group size depending on the level of independence that was considered. While such ecological comparisons are unrefined, e.g., rainfall must be used as a surrogate for habitat production, the fact that so few relationships were documented suggests that factors other than the ecological factors considered here are responsible for the observed differences in population characteristics. We suggest that much of the variability in howler population characteristics is related to events occurring in the recent history of the groups, such as habitat alteration, hunting, food tree crop failure, and disease.

Effects of Fruit Patch Availability on Feeding Subgroup Size and Spacing Patterns in Four Primate Species at Tinigua National Park, Colombia

Abstract: We examined the effects of fruit patch size, density, and distribution on feeding subgroup size and feeding bout duration in four Neotropical primates—Lagothrix lagotricha, Ateles belzebuth, Cebus apella, and Alouatta seniculus—based on 2494 hr of focal animal sampling in Tinigua National Park, Colombia. All four species show positive correlations between patch size (tree diameter at breast height; DBH) and subgroup size (maximum number of individuals from the focal group feeding at the same time), but the determination coefficients are very low and this relationship does not seem to influence strongly the actual size of the groups. Ateles showed the weakest correlation between patch size and feeding party size, and their fission–fusion sociality is best explained as a mechanism to reduce intragroup competition rather than to adjust the subgroup size to the availability of patches. Feeding associations as well as proximal spacing patterns, in general, are more alike in species with similar ecological requirements.

Long-Term Habitat Use by Mountain Gorillas ( Gorilla gorilla beringei ). 2. Reuse of Foraging Areas in Relation to Resource Abundance, Quality, and Depletion

Abstract: Resource depression caused by current feeding and the rate of resource renewal should influence foragers' decisions about when to revisit foraging areas. Adjustment of foraging paths and revisit rates should be particularly important when resources renew slowly. Foragers can also benefit by returning more often to highly profitable than to less profitable foraging areas. Many highly frugivorous primates seem to time revisits to fruit sources so as to harvest fruit efficiently and, also, use efficient search paths. Fewer data on non-frugivores exist. Mountain gorillas are folivores that eat mostly perennially available, continuously growing herbs and vines. Vegetation regenerates slowly from the effects of gorilla trampling, though trampling can also facilitate food species productivity, at least in the short term. Adjustment of intervals between visits to foraging areas to the extent of previous use and to resource renewal rates should increase gorilla foraging efficiency. Long-term data on habitat use by 6 mountain gorilla social units show that revisit intervals vary in association with variation in the extent of previous use and in plant productivity. However, they also revisit areas more often, the higher the biomass and nutritional quality of food there. These data are generally consistent with the hypothesis that the gorillas crop resources on a sustained-yield basis, though more precise data on areal revisits and complementary long-term data on vegetation composition would be needed to test the hypothesis.

Factors Affecting Party Size in Chimpanzees of the Mahale Mountains

Abstract: We studied factors affecting party size and composition of wild chimpanzees at Mahale (M group) over an 11-month period. Parties with 1–5 individuals were most frequent (37.8%; 153/405 parties); they included 94.7% of all male parties (n = 76) and 81.3% of all female parties (n = 75). The median of monthly values was the standard for analysis. We divided the year into four periods based on the median size of monthly bisexual parties (30.9 individuals; includes both males and females): monthly bisexual party sizes were larger in May–June (period II) and October–January (period IV) and smaller in February–April (period I) and July–September (period III). Only bisexual parties changed in size with period. The number of fruit items (=species) eaten was fewer in periods II and IV when abundance per item appeared to be great. The sizes of bisexual parties, which included cycling females with maximal anogenital swelling, were larger, and their representation (%) in all bisexual parties was greater in periods III and IV. The numbers of both cycling females and cycling females with maximal anogenital swelling were also larger in periods III and IV. The percentage of cycling females with maximal anogenital swelling was greater in periods II and III. The results of this study and those of Nishida (1979) suggest that seasonal variation in party size of Mahale chimpanzees maintains a relatively consistent annual cycle. The factors assumed to affect party sizes are fruit availability and the presence of cycling females with maximal anogenital swelling.

Habitat Use and Ranging of Wild Bonobos (Pan paniscus) at Wamba

Abstract: The relationship between vegetation and ranging patterns of wild bonobos at Wamba, Democratic Republic of the Congo, was examined. Via Landsat data, we distinguished three types of vegetation—dry forest, swamp forest, and disturbed forest—at Wamba. The home ranges of the study groups changed considerably from year to year, due mainly to intergroup relationships. The population density of each group varied between 1.4 and 2.5 individuals per km2and was lowest during a period of population increase. Home ranges consisted mainly of dry forest. The bonobos used dry forest more frequently than the other forest types, though they also used swamp and disturbed forest almost every day. The latter types of forest seemed to be important resources for the bonobos, owing to the abundant herbaceous plants that are rich in protein and constantly available. The bonobos tended to use dry forest more frequently in the rainy season than in the relatively dry season, probably because the favored fruits in the dry forest were mostly available in the rainy season. There was no seasonal difference in the size of the daily ranging area.

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

Abstract: 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.

Parasites of Wild Howlers (Alouatta spp.)

Abstract: A literature review of howler parasites provides the basis for an overview of the ecological significance of parasite surveys in primates. Within this framework, we have added insights into the interactions between primate hosts and their parasites from a long-term study in Costa Rica. We collected fecal samples from mantled howlers (Alouatta palliata) over a 9-year period (1986–1994 inclusive) and analyzed them for parasite eggs, larvae, cysts, and oocysts. We found many misperceptions inherent in the typical methodology of primate parasite surveys and in the reporting of the findings. Our work in Costa Rica suggests that a snapshot effect occurs with most surveys. A static view does not reflect the dynamic and changing ecological interaction between host and parasite. We describe some problems with parasite data analyses that emphasize the need for long-term longitudinal surveys in wild primate groups.

Population Dynamics of Wild Bonobos ( Pan paniscus ) at Wamba

Abstract: We analyzed population dynamics and birth seasonality of wild bonobos at Wamba, Democratic Republic of the Congo, based on 20 years of observations (1976–1996). Wamba Bonobo infant mortality is much lower than that reported for chimpanzees. This seemes to be related to several socioecological characteristics of bonobos: the use of abundant fruit and herbaceous foods, larger food patch size, female feeding priority, and the absence of infanticide. The mean interval between live births of 4.8 years is shorter than those reported for chimpanzees, and some females simultaneously carried and nursed two successive offspring. Mother–offspring conflicts, such as refusal of suckling attempts and interference with mothers' copulation, which are common in chimpanzees, are rare in Wamba bonobos. A birth peak seems to occur during the light rainy season from March to May, just after the season with the least rainfall. This timing of births is similar to those reported for chimpanzee populations, and might benefit both mother and offspring by maximizing the amount of time before the next dry season.

Tree Holes Used for Resting by Gray Mouse Lemurs (Microcebus murinus) in Madagascar: Insulation Capacities and Energetic Consequences

Abstract: I studied the insulation capacity of tree holes used by gray mouse lemurs (Microcebus murinus) in a primary dry deciduous forest in western Madagascar during the cool dry season. Tree holes had an insulating effect, and fluctuations of air temperatures were less extreme inside the holes than outside them. The insulation capacity of the tree holes peaked between 0800 and 1100 hr, when ambient temperatures ranged between 25 and 30°C. To compare tree holes, I calculated the mean difference between the internal temperature )(Ti ) and the external temperature (Te ) for each tree hole. Thus large differences indicate good insulation capacities. The mean difference of tree holes in living trees was significantly larger than that of tree holes in dead trees, which shows that insulation in living trees is more effective. During the dry season, the insulation capacity of tree holes in living trees decreased and was lowest in July, whereas the insulation capacity of holes in dead trees remained approximately constant. Physiological studies under natural temperature and light condition in Microcebus murinus reveal that daily torpor saves around 40% of the daily energy expenditure compared to normothermia. However, torpor can be maintained only up to the threshold body and ambient temperature of 28°C, whereat Microcebus murinus have to terminate torpor actively. By occupying insulating tree holes, mouse lemurs may stay longer in torpor, which increases their daily energy savings by an extra 5%.

Survey of Grauer's Gorillas (Gorilla gorilla graueri) and Eastern Chimpanzees (Pan troglodytes schweinfurthi) in the Kahuzi-Biega National Park Lowland Sector and Adjacent Forest in Eastern Democratic Republic of Congo

Abstract: We describe the distribution and estimate densities of Grauer's gorillas (Gorilla gorilla graueri) and eastern chimpanzees (Pan troglodytes schweinfurthi) in a 12,770-km2area of lowland forest between the Lowa, Luka, Lugulu, and Oku rivers in eastern Democratic Republic of Congo, the site of the largest continuous population of Grauer's gorillas. The survey included a total of 480 km of transects completed within seven sampling zones in the Kahuzi-Biega National Park lowland sector and adjacent Kasese region and approximately 1100 km of footpath and forest reconnaissance. We estimate total populations of 7670 (4180–10,830) weaned gorillas within the Kahuzi-Biega lowland sector and 3350 (1420–5950) individuals in the Kasese survey areas. Within the same area, we estimate a population of 2600 (1620–4500) chimpanzees. Ape nest site densities are significantly higher within the Kahuzi-Biega lowland sector than in the more remote Kasese survey area in spite of a significantly higher encounter rate of human sign within the lowland sector of the park. Comparison of our data with information obtained by Emlen and Schaller during the first rangewide survey of Grauer's gorillas in 1959 suggests that gorilla populations have remained stable in protected areas but declined in adjacent forest. These findings underscore the key role played by national parks in protecting biological resources in spite of the recent political and economic turmoil in the region. We also show that forest reconnaissance is a reliable and cost-effective method to assess gorilla densities in remote forested areas.

Postconflict Affiliation and Stress-Related Behavior of Long-Tailed Macaque Aggressors

Abstract: Previous studies on macaques and baboons showed that after agonistic conflicts aggressees as well as aggressors show an increase in stress-related behavior such as scratching. Reconciliation reduces stress-related behavior of the aggressee. We investigated the influence of various affiliative postconflict behaviors of the aggressor on the aggressor's scratching rates in captive long-tailed macaques: reconciliation, contacts with the aggressee's kin (or substitute reconciliation), and contact with other group members (or triadic affiliation). After a conflict, the aggressor showed an increase in rates of scratching. Scratching rates were reduced after reconciled conflicts compared to nonreconciled conflicts. Substitute reconciliation did not reduce scratching when we controlled for the influence of reconciliation, i.e., the aggressor might not interpret it as a substitute for reconciliation. Triadic affiliation did not reduce scratching rates, hence, triadic affiliation probably does not console the aggressor. Scratching rates after reconciliation are significantly lower than scratching rates after triadic affiliation. This proves that the stress-reducing effect of reconciliation is not due to the calming effect of general body contact but that the stress reduction is specifically associated with contacts with the former opponent. The contestants are anxious about their relationship, and only reconciliation takes away this anxiety. Reconciliation is thus an important social repair strategy.

Seasonality and Socioecology: The Importance of Variation in Fruit Abundance to Bonobo Sociality

Abstract: The assumption that nonseasonal, evergreen, rain forests contain more continuously available food resources than seasonal rain forests is fundamental to comparisons made between the socioecology of the male-bonded Pan troglodytes and the female-based social system of the Pan paniscus. Chimpanzee females may be less social due to the high costs of feeding competition, whereas in the more food-rich central African rain forests such as the Lomako forest, female bonobos can associate and socially bond. The Lomako Forest experiences two wet and two dry seasons a year. Data on fruit abundance and sociality show that despite monthly variation in fruit availability, there was no consistent seasonal variation in fruit abundance or dietary breadth. Bonobo use of nonfig fruits, figs, THV, and leaves did not follow seasonal patterns. Leaves and THV may act as complementary sources of plant protein and their use was inversely correlated. Monthly variation in fruit abundance was associated with a significant decrease in the number of males in a party but not in the number of females. Focal males were frequently solitary during 1 of the 3 months with the smallest party sizes. In contrast, females remained social with each other throughout the year. Therefore, seaonality at Lomako appeared to be less marked than at comparable chimpanzee sites, such that the variation in fruit abundance did not fall below a level that prohibits female sociality.

Selectivity and Variability in the Diet of Milne-Edwards' Sifakas ( Propithecus diadema edwardsi ): Implications for Folivory and Seed-Eating

Abstract: Propithecus diadema edwardsi combines a strategy of high selectivity for many food items with the use of more readily available and abundant foods. Selectivity is seen in relation to taxonomic affiliations of plants eaten, forest composition, and phenological patterns. Feeding time is concentrated on relatively uncommon species as indicated by high selection ratios. Annual and monthly dietary diversity is low, with food choices restricted to a relatively narrow set of species: a total of 82 and 73 plant species were eaten by the two groups of which, only 22 and 16% contributed >1% feeding time, however, these values fall with commonly reported ranges. Plant parts eaten varied widely from month to month, with overall variance in seed-eating higher than variance in young or mature leaf feeding. No correlation between percentage feeding time and food availability is significant; instead, monthly feeding patterns are better explained by the availability of particular plant species, which also supports a view of high selectivity in food choice. In contrast to selectivity for many seed, whole fruit, and new leaf sources, reliably abundant food sources appear to be leaves from lianas, epiphytes, and parasites, which were eaten in all focal months. While dramatic interannual differences in dietary composition in response to the absence of a preferred seed source highlight the interpretation of folivores as surviving food crunches by turning to readily available, poor quality resources, this strategy is not unique to folivores.

Different Population Dynamics of Microcebus murinus in Primary and Secondary Deciduous Dry Forests of Madagascar

Abstract: The goal of this study was to identify causes for lower population densities of mouse lemurs (Microcebus murinus) in secondary than in primary dry deciduous forests of western Madagascar. Variations in the abundance of Microcebus murinus are linked to the capacity to enter energy-saving torpor during the dry season. Under natural conditions in primary forest, Microcebus murinus can maintain daily torpor (and possibly hibernation) as long as body temperatures remain below 28°C. Females are more likely to hibernate than males, resulting in skewed sex ratios of captured Microcebus murinus in the primary forest. In the secondary forest the sex ratio of subjects captured during the dry season is more balanced than in primary forest, indicating that fewer females go into torpor in secondary than in primary forest. Secondary forests have fewer large standing or fallen trees that might provide holes as shelter for Microcebus murinus. Ambient temperatures are higher in secondary than in primary forests and higher outside than inside tree holes. These high ambient temperatures might hinder the ability of Microcebus murinus to maintain torpor for prolonged periods in secondary forests. Mouse lemurs from secondary forest have lower body mass than in primary forest. The year-to-year recapture rate is zero in secondary forest and thus significantly lower than in primary forest. This indicates that survival rates are lower in secondary than in primary forests. Thus, secondary forests may be of limited value as buffer zones or even corridors for mouse lemurs.

Seasonality in the Ecology and Life Histories of Mountain Gorillas (Gorilla gorilla beringei)

Abstract: The abundance of food, especially that of fruit and often that of young leaves, varies considerably over time for most primates. This variation can depend on or be independent of seasonality in rainfall. Mountain gorillas (Gorilla gorilla beringei) in the Virungas are exceptional: their habitat contains almost no edible fruit, and they mostly eat perennially available herbs and vines that are densely and evenly distributed in much of their habitat. Earlier studies documented little consistent temporal variation in mountain gorilla diets and habitat use, except for seasonal use of bamboo by some groups, and documented no birth seasonality. Long-term data (≤ 7 years) on 6 mountain gorilla groups confirm these results for habitat use, except that they show some seasonality in use of the upper altitudinal extremes of the gorillas' home ranges for unclear reasons. Relatively low and inconsistent variation in habitat quality over time should lower the costs of grouping for gorillas compared to other apes. Long-term data also confirm the absence of seasonality in births and conceptions. However, they show that mortality rates and risk of respiratory infections vary directly with rainfall. These relationships are probably causal and may be mediated through thermoregulatory stress.

Growth of Mantled Howler Groups in a Regenerating Costa Rican Dry Forest

Abstract: We examined population dynamics in mantled howlers (Alouatta palliata palliata) in a regenerating tropical dry forest in Santa Rosa National Park (SRNP), Costa Rica. The population has grown at a rate of about 7% per annum during the past decade. The growth in numbers from 342 in 1984 to 554 in 1992 reflects an increase in the number of groups (from 25 to 34) and a slight increase in their average size (from 13.6 to 16.3). Population density has increased from 4.9 to 7.9 individuals per km2. Santa Rosa's population density and group compositions are similar to those at several other mantled howler sites, but densities of mantled howlers are much higher at two other well-studied sites: La Pacifica and Barro Colorado Island (BCI). We relate the low density of howlers at Santa Rosa to local historical and ecological factors. Howler populations at high and low densities differ in average group size and sex ratio. At high population densities, groups are larger and include more adult females. The number of male howlers per group appears to be more strictly limited and less variable than the number of females is. However, there is greater variation in male group membership at Santa Rosa than at La Pacifica or BCI, and at Santa Rosa there are more generating forests available into which males and females can disperse and form new groups. We present case studies describing two ways in which new howler groups are formed, and we suggest that, compared to females and compared to males at high density sites, males are relatively advantaged in the uncrowded habitats at Santa Rosa and other low density sites.

How Precisely Do Bonobos (Pan paniscus) Grasp Small Objects

Abstract: The general objective of this study was to compare the precise grasping behavior and intermanual differences in performance between three Pan paniscus and five Homo sapiens in grasping small objects. We compared the temporal pattern of two submovements of consecutive grasping cycles, the (visuomotor) reaching and the (sensorimotor) grasping. Both species were similarly successful in this task, they showed a behavioral right-hand preference and preferred specific types of grips. Bonobos required less time for reaching an object but a much longer time to grasp it than humans did. Thus, the species pursued different strategies. We assumed that this might be due to the different grip techniques. However, grip preferences did not serve a quicker intramanual performance but they pronounced differences between hands. Intermanual differences in timing were restricted to the reaching part and more strongly in bonobos than in humans. However, the right hand need not necessarily perform quicker. As in the case of humans, we assume that attentional cues were focused more on preparing a proper grip with the right hand than on a quick performance. However, strong intermanual differences in bonobos may indicate an overall stronger neuronal asymmetry in the motor organization of the finger musculature that prepare a proper grip than is true of humans.

Comparative Perspectives on Bimaturism, Ontogeny, and Dimorphism in Lemurid Primates

Abstract: We address questions regarding the general absence of dimorphism in lemurid primates (Hapalemur, Eulemur, and Varecia) through comparative analyses of ontogeny. We described and analyzed body mass growth data for 9 lemurid taxa and compared them to similar data for anthropoid primates. Lemurids tend to grow rapidly over a short period of time when compared to anthropoid primates of similar body sizes. Size variation among lemurid taxa arises primarily as a consequence of differences in rates of growth. Comparative analyses of body mass growth data suggest that natural selection has produced ontogenetic adaptations in lemurids that center on relatively short periods of growth. Reduced growth periods preclude the evolution of sexual dimorphism through bimaturism—a sex difference in the length of the growth period—despite high levels of intermale competition. Selective factors related to seasonal variability of lemurid habitats play important roles in limiting the potential for the evolution of bimaturism. Other selective factors that limit bimaturism are related to female reproductive synchrony. In combination, they favor relatively early male maturation, precluding sexual selection that would otherwise promote the evolution of dimorphism through bimaturism. Natural selection on growth rates may preclude somatic responses to sexual selection that involve elevated male growth rates. In general, existing ontogenetic or life history adaptations appear to restrict responses to sexual selection in male lemurids.

Effective solutions for howler conservation

Abstract: All species of howlers are at risk due to habitat destruction. I review the three most serious threats to howler survival—agricultural disturbance, logging disturbance, and hunting—both in a general context and in species accounts. Withal, the adaptability of howlers has allowed them to be a widespread genus throughout Central and South America. Alouatta is especially adaptable due to its generalized folivorous diet. I discuss how this adaptability relates to survival in situations of habitat destruction. In addition, I examine howler preference for riverine habitats and their ability to capitalize on secondary forests and secondary growth. I take a practical approach, suggesting some solutions for species survival, which include habitat management and reintroductions. Successful conservation models highlighted are the Community Baboon Sanctuary in Belize, in which villagers manage their lands for the benefit of the black howler and for their own economic benefit from ecotourism, and a village-sustainable logging system in Quintana Roo, Mexico, which has reduced levels of deforestation while benefiting both villagers and howlers, as well as other wildlife. A successful translocation of black howlers in Belize in order to reestablish a viable population is also described. The paper concludes with further suggestions to encourage howler conservation.

Intertroop Transfer and Dominance Rank Structure of Nonnatal Male Japanese Macaques in Yakushima, Japan

Abstract: We examined the interaction between intertroop transfer and male dominance ranks in a wild population of Japanese macaques (Macaca fuscata yakui) in Yakushima using data collected over 15 years. Intertroop transfer tended to maintain a linear, stable, and age-graded dominance rank order among nonnatal males irrespective of variation in troop size or composition. All males that joined a troop at the top of the rank order were prime adults. Among males joining at lower ranks, entry at the most subordinate position in the hierarchy was common. Males joining at lower ranks tended to join troops in which all other resident males were the same age or older. Adult males tended to join troops with few or no males. Young males tended to join troops with many resident males, and in which a relatively large proportion of males was other young ones. Intertroop transfer was responsible for most rank changes of resident males. The most common cause of males rising in rank was the emigration or death of a higher-ranking male. Males fell in rank most frequently as a result of a new male joining the troop at the top of the hierarchy. Rank reversals among resident males were rare. The cumulative effects of male transfers produce sociodemographic variation within a troop over time and sociodemographic diversity among troops in a local population. A key feature of intertroop diversity is that larger troops have a significantly greater proportion of young males than smaller troops. This diversity also creates the potential for intertroop variation in the severity of male competition and provides a range of options for transferring males.

Primate Faunivores: Physical Properties of Prey Items

Abstract: Despite increasing interest in food texture and how it affects primate masticatory systems, little quantitative work has been done in this area. We discuss a method to quantify physical properties of primate food items. To demonstrate its utility, we performed fracture experiments on a sample of adult coleopterans (to approximate a “hard-object” insectivorous diet), larval lepidoterans (to approximate a “soft-object” insectivorous diet), and vertebrate muscle tissue (to represent a more carnivorous feeding regime).

Group Size, Home Range Use, and Seasonal Variation in the Ecology of Eulemur mongoz

Abstract: We collected data during a 10-month study carried out on the mongoose lemur, Eulemur mongoz, at Anjamena in northwestern Madagascar, which provide baseline information on seasonal variation in the ecology, home range use and some aspects of the behavior of two neighboring groups. We monitored group size of nine groups in the study area and assessed them for seasonal variation. We present additional information collected during short-term surveys in other areas before and during the study for comparison. The study groups were small family units, and changes in group size were limited to births and emigrations of sexually mature progeny. In spite of clear seasonal changes in climate and vegetation, there is no variation in grouping patterns, so it is not possible to correlate variation in group size with seasonal variation of ecological variables. Comparison with ecological data from other field studies on lemurids reveals differences in food resource distribution in western forests versus other types of Malagasy forest. This distribution of food resources may predict home range size in mixed frugivorous–folivorous lemurs. Small home ranges, mainly in the West, could be correlated with a uniform distribution of food resources. Finally, we suggest that the dry season in the West may not present frugivorous–folivorous lemurs with major problems in finding an adequate food supply. This is supported by the lack of seasonal differences in ranging behavior of mongoose lemurs.

Infant-nonmother interactions of free-ranging mantled howlers (Alouatta palliata) in Costa Rica

Abstract: Proximate and ultimate explanations of interactions between infants and nonmothers vary depending upon the relatedness of the interactors. We investigated interactions of infants and nonmothers from a 22-month continuous study and from the long-term monitoring of the mantled howler population of La Pacifica, Guanacaste Province, Costa Rica. Relatedness is low or absent in these mantled howler groups. Juvenile females appeared to practice care skills with older infants, but as most first infants died, they failed to benefit. Infant positive interactions with adults occurred with the mother and probable father. Other adult females behaved aggressively toward the youngest infants. Mothers were retentive of infants and responded negatively to these interactions, suggesting that they perceived them as threatening. Interactions with infants appear to reflect competition in groups of unrelated adults. A review of other populations of Alouatta palliata and other species of howlers indicate variability in social group size and suggest variability in intragroup relatedness. We suggest that further study will confirm that social behavior (including interactions with infants) will vary by resource availability (group size) and associated demographic patterns (male and female migration) that affect relatedness in howler social groups.

Skeletal Pathologies in a Population of Alouatta palliata: Behavioral, Ecological, and Evolutionary Implications

Abstract: We examined the pathological conditions in a sample of 200 skulls of Alouatta palliata recovered from Barro Colorado Island (BCI), Panama, to investigate the prevalence of disease and injury in a free-ranging monkey population. Forty-five individuals (22.5%) have some type of nondental pathological condition, indicating that illness and injury are relatively common in this population. Ten individuals, all older adult males, exhibit clear signs of antemortem trauma in the form of healed fractures. We attribute the trauma primarily to fighting, and its frequency (16.4% of adult males) contradicts previous assertions that BCI howlers are nonaggressive. Nine of the 37 scorable immature individuals (24%) have signs of a larval infestation, supporting the suggestion that ectoparasites play an important role in BCI howler mortality. Other pathological conditions in the sample include shifted muscle attachments, periostitis, arthritis, neoplasm, periapical abscesses, and facial deformity, as well as various dental abnormalities.