Intratree Variation in Fruit Production and Implications for Primate Foraging

We tested the hypothesis that fruit quantity and quality vary vertically within trees. We quantified intratree fruit production before exploitation by frugivores at different heights in 89 trees from 17 species fed on by primates in Kibale National Park, Uganda. We also conducted a pilot study to determine if the nutritional value of fruit varied within tree crowns. Depending on the species and crown size, we divided tree canopies into 2 or 3 vertical layers. In 2-layered trees, upper crowns produced fruits that were 9.6–30.1% bigger and 0.52–140 times the densities of those from lower crowns, with one exception. Among 2-layered trees, upper crowns produced a mean of 46.9 fruits/m³ (median 12.1), while lower crowns produced a mean of 14.1 fruits/m³ (median 2.5). Among 3-layered trees, upper crowns produced a mean density of 49.9 fruits/m³ (median 12.5), middle crowns a mean of 16.8 fruits/m³ (median 6.6), and lower crowns a mean of 12.8 fruits/m³ (median 1.8). Dry pulp and moisture were systematically greater per fruit in the highest compared to the lowest canopy layers (22.4% and 16.4% respectively in 2-layered trees, 49.7% and 21.8% respectively in 3-layered trees). In 1 tree of Diospyros abyssinica, a pilot nutritional study showed that upper crown ripe fruit contained 41.9% more sugar, 8.4% more crude proteins, and 1.8 times less of the potentially toxic saponin than lower crown ripe fruit, but the result needs to be verified with more individuals and species of trees. We discuss the consequences of intratree variations in fruit production with respect to competition among frugivorous primates.     

Thirty Years of Research in Kibale National Park,Uganda, Reveals a Complex Picture for Conservation

Kibale National Park, Uganda, has a rich and abundant primate community and a complicated history of anthropogenic disturbance. Moreover, it has been the focus of over 30 yr of research and has received considerable attention from nongovernmental and governmental conservation organizations. As a result, Kibale serves as a valuable case study with which to evaluate the factors that regulate primate population density and the challenges of deriving generalizations for conservation. We review the impact of logging and forest fragmentation on primate population density and trace the efficacy of various conservation strategies. A 28-yr comparison of primate abundance in logged and unlogged forests and a 10-yr study of forest dynamics showed that primate recovery in logged areas is generally slow or not occurring at all for some species, which is likely driven by the fact that the forest is not recovering as expected. No primate species characteristic predicted their ability to live in forest fragments around Kibale. While a nutritional model was useful to predict the abundance of colobus in forest fragments outside of Kibale, a 5-yr study revealed that human land-use practices are more fundamentally shaping population dynamics. We evaluate data on primate abundance in light of Milton’s protein/fiber model to predict colobine biomass. We demonstrate that while the model can predict colobus biomass in pristine habitats, the 2 colobus species respond differently to disturbance. We offer suggestions for future conservation research and consider strategies to conserve forested national parks based on experiences gained over 30 yr.     

Nutritional ecology of Preuss's red colobus monkey (Colobus badius preussi Rahm 1970) in Korup National Park, Cameroon

Studies on the nutritive ecology of red colobus monkeys (Colobus badius preussi) in Korup National Park, Cameroon, investigated food plants and the parts eaten as well as nutritive values of the various food items. Food plants were identified through direct observations and collected for laboratory determination of nutrient content. Food plants were analysed for crude protein (CP), crude fibre (CF), ether extract (EE), nitrogen‐free‐extractive (NFE) and total ash (A). A total of seventeen species belonging to nine different families were identified as food plants for red colobus. Food and feeding habits showed that the species were highly folivorous, tending to prefer the young leaves of food plants. Lecomtedoxa klaineana (27.0%) and Xylopia aethiopica (22.2%) of families Sapotaceae and Annonaceae, respectively, constituted about 50% of total food items consumed. Results of nutrient composition indicated individual nutrients varied significantly (P < 0.01) with food species.     

Led by the nose: Olfaction in primate feeding ecology

Olfaction, the sense of smell, was a latecomer to the systematic investigation of primate sensory ecology after long years in which it was considered to be of minor importance. 1 This view shifted with the growing understanding of its role in social behavior 2 and the accumulation of physiological studies demonstrating that the olfactory abilities of some primates are on a par with those of olfactory‐dependent mammals such as dogs and rodents. 3 , 4 Recent years have seen a proliferation of physiological, behavioral, anatomical, and genetic investigations of primate olfaction. These investigations have begun to shed light on the importance of olfaction in the process of food acquisition. However, integration of these works has been limited. It is therefore still difficult to pinpoint large‐scale evolutionary scenarios, namely the functions that the sense of smell fulfills in primates’ feeding ecology and the ecological niches that favor heavier reliance on olfaction. Here, we review available behavioral and physiological studies of primates in the field or captivity and try to elucidate how and when the sense of smell can help them acquire food.     

Primate Group Size and Abundance in the Caatinga Dry Forest, Northeastern Brazil

The Caatinga dry forest poses a series of ecological challenges for mammals in general and primates in particular. The erratic rainfall pattern impacts on plant diversity and phenological patterns; from year to year there is marked variability in fruit production and failure to fruit is common. The harshness apparently accounts for the impoverished mammalian fauna. However, data on primate abundance, distribution, and possible environmental effects on primate density are lacking in this type of dry forest. I censused the primate community in 3 habitats of the Serra da Capivara National Park, Piaui, NE Brazil, over a total distance of 318 km. Overall, the abundance of primates in the Caatinga dry forest is very low as a consequence of low abundance of food resources both in space and time. Alouatta caraya (predominantly folivorous) occurs at extremely low density, and during the dry season are apparently confined to canyon areas, where trees retain their leaves. Callithrix jacchus has morphological feeding specializations for gum-eating, and gum is an important resource during food bottleneck periods. Nonetheless, Callithrix jacchus occurs at comparatively low densities. Group sizes for howlers and marmosets in the Caatinga are significantly smaller than in other forest types. Contrarily, Cebus apella libidinosus had an average group size within the range reported for Amazonian and Atlantic forests. Researchers consider the generalized diet of capuchins as the explanation for their similar abundance in different habitats, indicating relative independence from ecological constraints. However, I suggest that capuchin foraging style and cognitive abilities are important factors accounting for their unreduced group size and density even under extreme conditions.     

Assessment of scale-dependent foraging behaviour in southern elephant seals incorporating the vertical dimension: a development of the First Passage Time method

1. Identifying the spatial scales at which top marine predators forage is important for understanding oceanic ecosystems. Several methods quantify how individuals concentrate their search effort along a given path. Among these, First-Passage Time (FPT) analysis is particularly useful to identify transitions in movement patterns (e.g. between searching and feeding). This method has mainly been applied to terrestrial animals or flying seabirds that have little or no vertical component to their foraging, so we examined the differences between classic FPT and a modification of this approach using the time spent at the bottom of a dive for characterizing the foraging activity of a diving predator: the southern elephant seal. 2. Satellite relayed data loggers were deployed on 20 individuals during three successive summers at the Kerguelen Islands, providing a total of 72 978 dives from eight juvenile males and nine adult females. 3. Spatial scales identified using the time spent at the bottom of a dive ( = 68.2 +/- 42.1 km) were smaller than those obtained by the classic FPT analysis ( = 104.7 +/- 67.3 km). Moreover, foraging areas identified using the new approach clearly overlapped areas where individuals increased their body condition, indicating that it accurately reflected the foraging activity of the seals. 4. These results suggest that incorporating the vertical dimension into FPT provides a different result to the surface path alone. Close to the Antarctic continent, within the pack-ice, sinuosity of the path could be explained by a high sea-ice concentration (restricting elephant seal movements), and was not necessarily related to foraging activity. 5. Our approach distinguished between actual foraging activity and changes in behaviour induced by the physical environment like sea ice, and could be applied to other diving predators. Inclusion of diving parameters appears to be essential to identify the spatial scale of foraging areas of diving animals.     

Biodiversity, ecosystem functioning and food webs in fresh waters: assembling the jigsaw puzzle

1. Dramatic advances have been made recently in the study of biodiversity–ecosystem functioning (B-EF) relations and food web ecology. These fields are now starting to converge, and this fusion has the potential to improve our understanding of how environmental stressors modulate ecosystem processes and the supply of 'goods and services'.
2. Food web structure and dynamics can exert particularly strong influences on B-EF relations in fresh waters, as consumer–resource interactions (e.g. trophic cascades) are often more important than horizontal interactions within trophic levels. For instance, many freshwater food webs are size structured, with large organisms tending to occupy the higher trophic levels and often exerting powerful effects on ecosystem processes. However, because they are also vulnerable to perturbations, non-random losses of these large taxa can alter both food web structure and ecosystem functioning profoundly.
3. Recently, the focus of food web research has shifted away from exploring patterns, towards developing an understanding of processes (e.g. quantifying fluxes of individuals, biomass, energy, nutrients) and how the two interact. Many of the best-characterized food webs are from fresh waters, and these ecosystems are now being used to address some of the shortcomings of earlier B-EF studies. I have identified several key gaps in our current knowledge and highlighted potentially fruitful avenues of future B-EF and food web research.
4. A major challenge for this newly emerging research is to place it within a unified theoretical framework. The application of metabolic theory and ecological stoichiometry may help to achieve this goal by considering biological systems within the constraints imposed upon them by physical and chemical laws.     

Variation in daily and seasonal foraging routines of non-breeding barnacle geese (Branta leucopsis): working harder does not overcome environmental constraints

This study describes the changes in stored body fat in Svalbard barnacle geese Branta leucopsis over the non-breeding period, and uses behavioural patterns to explain the variation in body fat stores. It was predicted that foraging effort would: (1) increase at low food densities; (2) increase when days were short; (3) decrease in smaller flocks; (4) be bimodally distributed throughout the day for long days, but constant for short days. Time constraints were found to be the major driving force behind foraging decisions during the shortest days of mid-winter, whereas food density was found to drive decisions during longer days. Field observations of fat stores showed that fat was rapidly accumulated at the start and end of the non-breeding period, but that fat stores remained constant during mid-winter. It is suggested that day length prevented a positive fat storage rate in mid-winter through the limitation of foraging time, even though foraging effort was high during this period. During a single day, evidence for a bimodal foraging routine was found, where feeding activity was concentrated in the early morning and late afternoon periods. This pattern was found in the full range of day lengths, which suggests that even for short days, feeding must be interrupted so that other essential activities can be conducted. It is concluded that the behavioural choices of barnacle geese were constrained by environmental conditions, and that these behavioural patterns allow the variation in fat stores to be explained.     

Variation in diet and ranging of black and white colobus monkeys in Kibale National Park,Uganda

Recently, considerable intraspecific variation in the diets and ranging behavior of colobine monkeys has been described, although in most cases this has involved documenting variation between, not within, sites. Some African colobines, such as guerezas (Colobus guereza), are relatively abundant in disturbed habitats that are very heterogeneous, raising the intriguing possibility that even groups with overlapping home ranges may exhibit large behavioral differences. If such differences occur, it will be important to understand what temporal and spatial scales adequately portray a species’ or population’s diet and ranging behavior. This study documents within-site variation in the diet and ranging behavior of guerezas in the habitat types in which they are described to be most successful—forest edge and regenerating forest. We collected data on eight groups of guerezas with overlapping home ranges for 3–5 months each in Kibale National Park, Uganda. The guerezas were highly folivorous, with leaves constituting 78.5–94.0% of the groups’ diets. The percentage of mature leaves and fruit in the diet varied widely among and within groups. We show that differences among groups in the intensity with which they fed on specific tree species were not just related to phenology, but also to differences in the forest compositions of groups’ core areas. Range size estimates varied more than fivefold among groups and the minimum distance from groups’ core areas to eucalyptus forest (which all groups regularly fed in) was a better predictor of range size than was group size. These results reveal considerable variation in the diet and ranging behavior among groups with overlapping ranges and have implications for comparative studies, investigations of within- and between-group feeding competition, and the potential for populations to adapt to anthropogenic or natural environmental change.     

Multi-scale Data Used to Analyze the Spatial Distribution of French Grunts, Haemulon Flavolineatum, Relative to Hard and Soft Bottom in a Benthic Landscape

We evaluated the day-time distribution of juvenile and adult French grunts, Haemulon flavolineatum, relative to the spatial configuration of hard and soft bottom areas in a benthic landscape. Probability of juvenile presence on hard bottom sites was inversely correlated with distance to soft bottom. Adults presence at hard bottom sites showed no significant relationship with distance to soft bottom. A significant and positive relationship was found between presence of juveniles on hard bottom sites and area of soft bottom within 100m, but no significant relationship was found for area of soft bottom within 500m. Adults exhibited no significant relationship with area of soft bottom for either distance tested. These distributions are suspected to be the result of the combined influence of larval settlement patterns and foraging behaviors associated with hard and soft bottom. This study indicates that data collected at very fine scales can be analyzed in the context of the broad-scale mosaic of habitats in the benthic landscape to predict patterns of fish distribution. Such spatially explicit conclusions are not possible through analysis of fine-scale or broad-scale data alone.     

Foraging in tropical rain forests: The case of the penan of Sarawak,East Malaysia (Borneo)

Bailey et al. (1989) and Headland (1987) have recently proposed hypotheses stating that human foragers are unable to live in undisturbed tropical rain forests without some reliance on cultivated foods. The present discussion considers these hypotheses, as well as some of the evidence by which they have been tested. Four conceptual problems in the way these hypotheses have been formulated are identified: (1) assumptions about the relationship between key features of tropical forest ecosystems and human subsistence potential, (2) in-consistencies in the definition of pure foraging, (3) adherence to a dichotomy between foraging and agriculture, the result being that conscious and unconscious effects of exploitation on the demographic parameters of key resources is ignored, and (4) problems in defining the significance of ecotones. I consider the case of Penan hunter-gatherers of Borneo, a population which, by virtue of their reliance on the sago palm Eugeissona utilis, contradicts the conclusions of Bailey et al. and Headland. I consider salient aspects of Penan reliance on Eugeissona, and describe how Penan exploitation of this resource may positively effect its availability. This case is seen to provide a challenge to the hypotheses of Bailey et al. and Headland, not only in the extent to which it contradicts their conclusions but, more significantly, in what it reveals about the assumptions upon which their hypotheses are based. This points to the need for greater precision in the definition of future hypotheses about foraging in tropical forests.     

Interspecific Variation in Primate Countershading: Effects of Activity Pattern,Body Mass,and Phylogeny

Countershading is defined as an animal displaying a dark dorsal and light ventral surface. This color pattern is found in numerous species, yet there are relatively few quantitative tests that examine its potential adaptive nature. The most frequently employed explanation for countershading is its ability to increase crypsis. I used a comparative approach to test 2 predictions of this hypothesis in primates. First, diurnal species should exhibit stronger countershading versus nocturnal species because light levels are low at night, reducing the benefit of countershading. Second, the degree of countershading should decrease as body mass increases because large animals should have a lower predation risk. I collected data from 171 museum specimens representing 63 primate species. Using digital photography, I quantified the average luminance values on the ventral and dorsal surfaces of each specimen and defined the degree of countershading as the ratio of these values. Nocturnal primates displayed similar levels of countershading compared to diurnal species. Previous studies have shown that some nocturnal primates are more active when moonlight levels are high, suggesting that countershaded pelages may act as an antipredator adaptation. I also found support for a negative relationship between body mass and the degree of countershading using both species values and phylogenetically independent contrasts. There were some exceptions to this pattern, including low levels of countershading for most callitrichine species, which may be due to the vertical positional behavior of many callitrichines, thereby reducing any benefit gained from countershading. In sum, variation in primate countershading is related to body mass and phylogeny, but not to activity pattern.     

Feeding ecology and foraging behaviour of impala Aepyceros melampus in Lake Mburo National Park, Uganda

Intersexual and seasonal variation in foraging behaviour of impala (Aepyceros melampus), was studied in the Lake Mburo National Park, Uganda. There was a moderate seasonal difference in foraging efficiency (as measured by ‘acceptable food abundance’), with a minimum in dry season and a maximum in Rainy season. The variation between sexes was more distinct with a pronounced minimum in time spent browsing of males in early wet season. By distinguishing between feeding time spent grazing and feeding time spent browsing the seasonal variation was confirmed. The proportion of foraging time spent feeding (expressed as ‘food ingestion rate’) showed an inverse pattern with a maximum in the late dry season (75.5%), decreasing values throughout the Rainy season and a minimum in early dry season (57.8%). Differences between sexes were explained in terms of reproductive demands and seasonal balance in terms of moderate climate throughout the year. Impala foraging patterns in the bimodal tropics (two Rainy seasons) is discussed and compared with unimodal tropics. The findings are matched against current ideas on optimal foraging.     

Habitat selection and seasonal patterns of activity and foraging of mantled howling monkeys(Alouatta Palliata) in Northeastern Costa Rica

I conducted a 15- month ecological study of habitat preferences and activity and foraging patterns of two troops of mantled howling monkeys, Alouatta palliata,in a lowland rain forest at La Selva Biological Reserve in northeastern Costa Rica. The two troops specialized on different habitats in spite of the fact that both of them had all habitats available and were not constrained by neighboring troops since the population density of howlers is low (7- 15 howlers/km ² ).Troop 1 spent the majority of time in primary forest (80%) followed by secondary forest (10%), while troop 2 spent the majority of time in undisturbed riparian habitat (60%) followed by primary forest (30%). Habitat sampling indicates that neither the total number of stems, species, or families nor the diversity (Shannon index) or evenness is a good indicator for howler habitat selection. Instead the density of trees from the 12 species most commonly consumed by each troop is the most important factor. Activity and foraging patterns were not dependent upon the season as has been described for howling monkeys in forests with a more pronounced dry season at Barro Colorado Island, Panama, and La Pacifica in northwestern Costa Rica. This is likely a result of the more constant food supply at La Selva, combined with less intraspecific competition due to the low howler density. The intraspecific variability of foraging patterns and troop- specific habitat specialization observed in Alouatta palliatashould be considered in the conservation biology of primates. Primate relocation programs should include not only an ecological assessment of the release site but also a comparison of the release site with the habitat that the groups currently occupy.     

Temporal Variation in the Relative Abundance of Fruit Bats (Megachiroptera: Pteropodidae) in Relation to the Availability of Food in a Lowland Malaysian Rain Forest

The aims of this study were to investigate the diet and relative abundance of fruit bats in a lowland Malaysian rain forest and to test the hypothesis that the local assemblage structure of fruit bats varies significantly over time in relation to the availability of food. In total, 352 fruit bats of eight species were captured during 72,306 m² mist‐net hours of sampling between February 1996 and September 1999. Three species of fruit bats (Balionycteris maculita, Chironax melanocephalus, and Cynopterus brachyotis) that fed on a wide range of “steady state” and “big bang” food resources were captured continuously throughout the study period, with no significant variation in capture rates over time. In contrast, five species that fed exclusively or almost exclusively on “big bang” food resources were sampled intermittently, with significant temporal variation in the capture rates of two species (Cynopterus horsfieldi and Megaerops ecaudatus). Significant variation in the capture rates of the remaining three species (Dyacopterus spadiceus, Eonycteris spelaea, and Rousettus amplexicaudatus) could not be detected due to small sample sizes. Since ephemeral “big bang” food resources were only sporadically available within the study area and were associated with large canopy trees and strangler figs, these results suggest that food abundance, or the availability of specific food items, may be important factors limiting local fruit bat species diversity in old‐growth Paleotropical rain forest. Thus, only three fruit bat species were locally resident within the forest throughout the study period. Therefore, further studies on the ranging behavior and habitat requirements of Malaysian fruit bats are required to assess the adequacy of existing reserves and protected areas.