Mineral Resource Availability and Consumption by Colobus in Kibale National Park,Uganda

Very little information exists on mineral nutrition of tropical, forest-dwelling species, yet minerals are critical to growth, reproduction, and survival. We examined the mineral resources available to and consumed by colobus in Kibale National Park, Uganda. We combined behavioral data on black-and-white (Colobus guereza) and red colobus (Piliocolobus tephrosceles) in a section of unlogged forest, a heavily logged area, and a forest fragment with mineral analysis of their foods to estimate the proportion of the diet containing specific minerals (mineral content). We compared mineral content of colobus foods (natural and crops) across plant parts and among plant species. Additionally, we estimated mineral intake of frugivorous primates in Kibale from published dietary data and our estimates of mineral content of foods. Dietary mineral content for all colobus groups and frugivorous species is similar despite significant differences in the mineral content of foods. Ripe and unripe fruits are lower in mineral content than most foods. Foods rarely consumed, such as bark, petioles, and caterpillars have high levels of some minerals. The mineral content of crops is low in comparison to that ofnatural foods. For all colobus groups of both species, sodium content of foods was extremely low and iron content was generally low, suggesting that intake isbelow suggested requirements, though current suggested iron requirements may overestimate physiological needs. Copper content was marginal and deficient seasonally for most colobus groups. Despite a sodium-limiting environment, only one of 8 colobus groups appeared to select sodium; however, this may be due to a lack of variation in sodium content among plant species and a positive correlation between high plant sodium content and secondary compounds. Despite the lack of selection for sodium by colobines, some behaviors point to a potential sodium deficiency, including urine drinking, consumption of high-sodium swamp plants, and use of mud-puddles.     

Galago locomotion in Kibale National Park, Uganda

Very few locomotor studies have been conducted on galagos. This is surprising given their interesting anatomy and ecology, as well as their increasing species diversity. In this study we investigated locomotion and postures in two sympatric galagos species (Galagoides thomasi and Galago matschiei) living in Kibale National Park, Uganda. G. thomasi uses arboreal quadrupedalism and leaping, while G. matschiei is more leaping-oriented. Both species utilize small oblique branches in the mid-canopy. These similarities in substrate use are most likely due to the similar body sizes and anatomies of the two species, as well as to the structure and availability of trees in Kibale National Park. Lastly, we compare the locomotor patterns of G. thomasi and G. matschiei with those observed in the few other quantitative locomotor studies available for galagos.     

Tool use in insect foraging by the chimpanzees of Ngogo, Kibale National Park, Uganda

Chimpanzee tool use for resource acquisition has been reported at numerous research sites. The chimpanzees of the Kibale Forest, western Uganda, have not previously been observed to use tools in foraging for insects. Here I report the first observation of tool use by the chimpanzees of the Ngogo community of Kibale National Park, Uganda, in insect foraging. Three adult females, one adolescent male, and one juvenile male were observed making and using tools to probe into a fallen dead tree to collect insect and wood pieces. I discuss the importance of this observation, and the behavioral similarities with chimpanzees from other sites.     

Long-term trends in fruit production in a tropical forest at Ngogo,Kibale National Park,Uganda

Fruit production in tropical forests varies considerably in space and time, with important implications for frugivorous consumers. Characterizing temporal variation in forest productivity is thus critical for understanding adaptations of tropical forest frugivores, yet long-term phenology data from the tropics, in particular from African forests, are still scarce. Similarly, as the abiotic factors driving phenology in the tropics are predicted to change with a warming climate, studies documenting the relationship between climatic variables and fruit production are increasingly important. Here, we present data from 19 years of monitoring the phenology of 20 tree species at Ngogo in Kibale National Park, Uganda. Our aims were to characterize short- and long-term trends in productivity and to understand the abiotic factors driving temporal variability in fruit production. Short-term (month-to-month) variability in fruiting was relatively low at Ngogo, and overall fruit production increased significantly through the first half of the study. Among the abiotic variables, we expected to influence phenology patterns (including rainfall, solar irradiance, and average temperature), only average temperature was a significant predictor of monthly fruit production. We discuss these findings as they relate to the resource base of the frugivorous vertebrate community inhabiting Ngogo.     

Lethal intergroup aggression by chimpanzees in Kibale National Park, Uganda

Chimpanzees (Pan troglodytes) have hostile intergroup relations throughout most or all of their geographic range. Hostilities include aggressive encounters between members of neighboring communities during foraging and during patrols in which members of one community search for neighbors near territory boundaries. Attacks on neighbors involve coalitions of adult males, and are sometimes fatal. Targets include members of all age/sex classes, but the risk of lethal intergroup coalitionary aggression is highest for adult males and infants, and lowest for sexually swollen females. The best-supported adaptive explanation for such behavior is that fission-fusion sociality allows opportunities for low-cost attacks that, when successful, enhance the food supply for members of the attackers' community, improve survivorship, and increase female fertility. We add to the database on intergroup coalitionary aggression in chimpanzees by describing three fatal attacks on adult males, plus a fourth attack on an adult male and an attack on a juvenile that were almost certainly fatal. Observers saw four of these attacks and inferred the fifth from forensic and behavioral evidence. The attackers were males in two habituated, unprovisioned communities (Ngogo and Kanyawara) in Kibale National Park, Uganda. We also summarize data on other intercommunity attacks at Ngogo. Our observations are consistent with the "imbalance of power" hypothesis [Manson & Wrangham, Current Anthropology 32:369-390, 1991] and support the argument that lethal coalitionary intergroup aggression by male chimpanzees is part of an evolved behavioral strategy.     

Skeletal pathology in Pan troglodytes schweinfurthii in Kibale National Park, Uganda

The ecological pressures shaping chimpanzee anatomy and behavior are the subject of much discussion in primatology and paleoanthropology, yet empirical data on fundamental parameters including body size, morbidity, and mortality are rare for wild chimpanzees. Here, we present skeletal pathology and body size data for 20 (19 crania, 12 postcrania) chimpanzees (Pan troglodytes schweinfurthii) from Kibale National Park, Uganda. We compare these data with other East African populations, especially Gombe National Park. Estimated body size for Kibale chimpanzees was similar to other East African populations and significantly larger than Gombe chimpanzees. The high rates of trauma and other skeletal pathology evident in the Kibale chimpanzee skeletons were similar to those in the Gombe skeletal sample. Much of the major skeletal trauma in the Kibale skeletons was attributable to falls, although other pathologies were noted as well, including apparent injuries from snares, degenerative arthritis, and minor congenital abnormalities.     

Hunting Behavior of Chimpanzees at Ngogo,Kibale National Park,Uganda

Chimpanzees (Pan troglodytes) prey on a variety of vertebrates, mostly on red colobus (Procolobus spp.) where the two species are sympatric. Variation across population occurs in hunting frequency and success, in whether hunting is cooperative, i.e., payoffs to individual hunters increase with group size, and in the extent to which hunters coordinate their actions in space and time, and in the impact of hunting on red colobus populations. Also, hunting frequency varies over time within populations, for reasons that are unclear. We present new data on hunting by chimpanzees at Ngogo, Kibale National Park, Uganda, and combine them with earlier data (Mitani and Watts, 1999, Am. J. Phys. Anthropol. 109: 439–454) to examine hunting frequency and success, seasonality, and cooperation. The Ngogo community is the largest and has the most males of any known community. Chimpanzees there mostly hunt red colobus and are much more successful and make many more kills per hunt than at other sites; they kill 6–12% of the red colobus population annually. The number of kills and the offtake of meat per hunt increase with the number of hunters, but per capita meat intake is independent of hunting party size; this suggests that cheating occurs in large parties. Some behavioral cooperation occurs. Hunting success and estimated meat intake vary greatly among males, partly due to dominance rank effects. The high overall success rate leads to relatively high average per capita meat intake despite the large number of consumers. The frequency of hunts and of hunting patrols varies positively with the availability of ripe fruit; this is the first quantitative demonstration of a relationship between hunting frequency and the availability of other food, and implies that the chimpanzees hunt most when they can easily meet energy needs from other sources. We provide the first quantitative support for the argument that variation in canopy structure influences decisions to hunt red colobus because hunts are easier where the canopy is broken.     

Breeding birds in the tropical rain forests of Kibale National Park, Uganda

A combination of trapping and foot surveys was used to assess the breeding status of birds in unlogged and logged sites in the tropical rain forest of Kibale National Park, Uganda. Breeding of forest edge and gap species was greatly enhanced by logging, whereas crevice and hole nesting specialist breeders were adversely affected. Egg‐laying periods corresponded to higher rainfall regimes at the nearby field station. The relevance of these findings to bird conservation is discussed.     

浙江天童常绿阔叶林中11种常绿乔灌木叶片虫食状分析

为探讨昆虫对植物叶片的取食行为和伤害方式,作者选择浙江天童常绿阔叶林内的11种常绿乔灌木为对象,对叶片虫食状类型和格局进行分析。结果如下:(1)共发现16种虫食状类型,每种植物叶片虫食状类型数在10–13种之间,每种虫食状出现频率在0.5–28.7%之间。缘食状出现频率最高(28.7%),虫瘿和泡状出现频率最低(0.5%)。(2)叶片虫食状分布格局可分为3种类型,即一种虫食状占绝对优势的单优格局,如马银花(Rhododendronovatum)和檵木(Loropetalumchinense);两种虫食状(缘食状和顶食状)共占优势的双优格局,仅有木荷(Schimasu-perba);3种及3种以上虫食状占优势的多优格局(其余8种植物)。(3)叶片虫食状多样性指数变化在1.57–2.23之间,最高为苦槠(Castanopsissclerophylla),最低为马银花;乔木的多样性指数(2.040)高于灌木(1.882),优势种多样性指数高于伴生种,但差异均不显著;多样性指数反映了虫食状类型和出现频率的综合差异。(4)16种虫食状类型中有8对显著正相关,4对显著负相关,可能反映出不同类型昆虫取食植物的趋同和差异。     

Diet of chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda, 2. Temporal variation and fallback foods

Highly frugivorous primates like chimpanzees (Pan trogolodytes) must contend with temporal variation in food abundance and quality by tracking fruit crops and relying more on alternative foods, some of them fallbacks, when fruit is scarce. We used behavioral data from 122 months between 1995 and 2009 plus 12 years of phenology records to investigate temporal dietary variation and use of fallback foods by chimpanzees at Ngogo, Kibale National Park, Uganda. Fruit, including figs, comprised most of the diet. Fruit and fig availability varied seasonally, but the exact timing of fruit production and the amount of fruit produced varied extensively from year to year, both overall and within and among species. Feeding time devoted to all major fruit and fig species was positively associated with availability, reinforcing the argument that chimpanzees are ripe fruit specialists. Feeding time devoted to figs-particularly Ficus mucuso (the top food)--varied inversely with the abundance of nonfig fruits and with foraging effort devoted to such fruit. However, figs contributed much of the diet for most of the year and are best seen as staples available most of the time and eaten in proportion to availability. Leaves also contributed much of the diet and served as fallbacks when nonfig fruits were scarce. In contrast to the nearby Kanywara study site in Kibale, pith and stems contributed little of the diet and were not fallbacks. Fruit seasons (periods of at least 2 months when nonfig fruits account for at least 40% of feeding time; Gilby & Wrangham., Behavioral Ecology and Sociobiology 61:1771-1779, 2007) were more common at Ngogo than Kanyawara, consistent with an earlier report that fruit availability varies less at Ngogo [Chapman et al., African Journal of Ecology 35:287-302, 1997]. F. mucuso is absent at Kanyawara; its high density at Ngogo, combined with lower variation in fruit availability, probably helps to explain why chimpanzee population density is much higher at Ngogo.     

Tool Use by Chimpanzees at Ngogo,Kibale National Park,Uganda

Chimpanzees make and use a wide variety of tools in the wild. The size and composition of their toolkits vary considerably among populations and at least to some extent within them. Chimpanzees at several well documented sites mostly use tools in extractive foraging, and extractive tool use can substantially increase their foraging efficiency. They also use tools for hygiene and for several other purposes, including attracting the attention of conspecifics, as in leaf-clipping. Some of the interpopulation variation in toolkits results from ecological variation, but differences in the efficiency of social transmission, perhaps related to differences in social tolerance, presumably also contribute. I describe tool use by chimpanzees in an unusually large community at Ngogo, in Kibale National Park, Uganda. Researchers have described some tool use for the community previously, but this is the most extensive report and is based on observations over 11 yr. The Ngogo chimpanzees have a small toolkit and use tools rarely except in leaf-clipping displays and to clean body surfaces; notably, males often use leaf napkins to wipe their penes after copulation. Extractive tool use is rare and is limited mostly to leaf-sponging and, less often, honey-fishing. Social tolerance is not low at Ngogo, but use of tools for extractive foraging, in ways documented at other field sites, may have little potential to increase foraging efficiency. Future research will undoubtedly show more tool use by females, which were underrepresented in my observations, but will probably not document much increase in the toolkit or in the use of extractive tools.     

New Intercommunity Infanticides by the Chimpanzees of Ngogo,Kibale National Park,Uganda

Hostile intercommunity relations, including attacking and killing extra-community infants of both sexes have occurred at most wild chimpanzee sites. We describe three recent cases of intercommunity attacks on infants committed by members of the Ngogo chimpanzee community in Kibale National Park, Uganda. Two of the attacks resulted in confirmed infanticides while a third attack probably resulted in the infant's death. In common with previous accounts of chimpanzee infanticides, the attacks described here occurred during boundary patrols outside the Ngogo community's usual range, adult and adolescent males were the main participants, one infant was cannibalized after being killed, and the victims’ mothers did not accompany the attacking party back to the Ngogo range. However, the patrol parties during each infanticide were larger than before and included females from the Ngogo community. Our observations indirectly support both the range expansion and imbalance of power hypotheses, which address why and under which conditions chimpanzee intercommunity encounters lead to aggression. These cases of intercommunity infanticide add to the growing database of the phenomenon in wild chimpanzees.     

Population structure and behaviour of crop-raiding elephants in Kibale National Park, Uganda

We examined patterns of crop raiding by elephants across gender and age classes in relation to elephant life history and sociobiology and estimated the quantitative contribution of crops to elephant diet in Kibale National Park (KNP). Elephant dung‐boli sizes were used to estimate age and sex, while the presence of crop remains in the dung of crop‐raiding elephants was used as evidence of repeated raiding. To estimate the expected proportion of elephants raiding per age class, the age distribution of raiders was compared with the age distribution of all KNP elephants. Elephants raiding crops were predominantly males. They began raiding in expected proportions at 10–14 years while a higher than expected proportion raided crops at 20–24 years. These results suggest that crop raiding is initiated at an age when male elephants leave their families and a large proportion of elephants raid when they are approaching reproductive competition. Evidence from dung of crop raiders, suggests that repeated raiding increases with age. Crop raiders derived 38% of their daily forage from the short time spent raiding, consistent with expectations of foraging theory. Males may be more likely to learn crop raiding because they are socially more independent and experience intense mating competition than females.     

Forest association and phenology of wild coffee in Kibale National Park, Uganda

A study on the forest association and phenology of wild coffee ( Coffea canephora Pierre) was conducted in Kibale forest, Uganda. Nested quadrats were used to enumerate tree species, including coffee and herbaceous plants associated with forest and coffee stands. A total of 150 coffee trees was marked along transects and monthly scans carried out to score for fruits, flowers, leaves and leaf insect damage. Pre- and post-dispersal predation levels and coffee yield estimates were made by examining fruits from trees, forest floor and seasonal fruit falls into demarcated plots. In the forest, wild coffee stands are associated with low-quality forest types in terms of timber species (about 10.5 canopy species/study site) and low stocking densities of trees ≥ 50 cm d.b.h. (average 38 trees ha⁻¹ for each site) and poor forest regeneration. In the forest, wild coffee reproductive phases overlap with ripening, coinciding with flower bud and flower production. The variable peak ripening season falls between November and April. The wild coffee yields are generally low (average of 3.5 intact fruits 16 m⁻² month⁻¹), with low insect fruit/seed damage (4–19%) but high levels of wastage due to monkeys, bats and birds.     

Forest Regeneration in Logged and Unlogged Forests of Kibale National Park, Uganda

Processes of forest regeneration in two unlogged areas and in three areas that were logged nearly 25 years ago were quantified in Kibale National Park, Uganda. For forests to recover from logging, one would predict recruitment and growth processes to be accelerated in logged areas relative to unlogged areas, facilitating increased recruitment of trees into the adult size classes. We examined this prediction first by determining the growth of 4733 trees over a 51 to 56 month period and found that growth rates in the most heavily logged area were consistently slower than in the two unlogged areas. In contrast, the lightly logged forest had similar growth rates to unlogged areas in the small size classes, but trees in the 30 to 50 cm DBH size cohort exhibited elevated growth rates relative to the unlogged areas. Mortality was highest in the heavily logged areas, with many deaths occurring when healthy trees were knocked over by neighboring treefalls. We found no difference in the density or species richness of seedlings in the logged and unlogged forests. The number of seedlings that emerged from the disturbed soil (seed bank+seed rain) and initially seed-free soil (seed rain) was greater in the logged forest than in the unlogged forest. However, sapling density was lower in the heavily logged areas, suggesting that there is a high level of seedling mortality in logged areas. We suggest that the level of canopy opening created during logging, the lack of aggressive colonizing tree species, elephant activity that is concentrated in logged areas, and an aggressive herb community, all combine to delay vegetation recovery in Kibale Forest.     

Tree population dynamics in Kibale National Park, Uganda 1975–1998

Changes in species composition, stem abundance, and basal area of trees taller than or equal to 10 m in a medium altitude tropical rain forest at the Ngogo study area, Kibale National Park, Uganda are described for the period between 1975 and 1998 ( n  = 23 years). Two enumeration episodes were conducted in 263 plots of 5 m by 50 m during 1975–80 and 1997–98. During this period, species richness decreased by 3% (from 92 to 89). Species diversity (H') also declined slightly from 2.97 to 2.86. The number of trees recorded in the sample plots decreased by 8% (from 2545 to 2329), while basal area decreased from 49.48 m² ha⁻¹ to 48.68 m² ha^−1. However, stem abundance and basal area increased for some species.