Fruit Availability and Habitat Use by Chimpanzees in the Kalinzu Forest,Uganda: Examination of Fallback Foods

We studied seasonal change in habitat use by chimpanzees in the Kalinzu Forest, Uganda. The forest comprises various types of vegetation. For each vegetation type, we compared number of chimpanzees (per km²) that used the vegetation with fruit availability in different census periods. We estimated the number of chimpanzees by nest count and fruit availability via density of fallen fruit. The mixed mature forest contained a large amount of fruit during the high-fruiting season, but it decreased rapidly in the low-fruiting season. The number of chimpanzees also decreased in mixed mature forest in approximate proportion with fruit availability. In the Parinari-dominated mature and secondary forests, both fruit availability and number of chimpanzees were very low throughout the study. In the Musanga-dominated secondary forest, the number of chimpanzees increased toward the low-fruiting season, though the fruit availability decreased slightly. A multiple regression analysis showed that various fruits had significant effects on the number of chimpanzees during the high-fruiting season, while only Musanga leo-errerae had a significant effect during the low-fruiting season. The results suggest that the fruit of Musanga leo-errerae functions as a fallback food, and a combination of different vegetation types supports the chimpanzees in the Kalinzu Forest.     

The fruit phenology of Musanga leo-errerae and its importance for chimpanzee diet in Kalinzu Forest Uganda

This study reports the rate of fruit phenological pattern of Musanga leo-errerae and how it sustains the chimpanzee population better than other fruits in Kalinzu Forest Reserve. We analysed 2635 faecal samples to determine the proportion of M. leo-errerae by composition of fruit diet compared with other fruits eaten by chimpanzees. Musanga leo-errerae trees were monitored for fruit production between November 2002 and December 2004. Musanga leo-errerae fruit production did not vary significantly between months (ANOVA, F  = 2.0, d.f. = 11, P  = 0.13). The size of fruit and rate maturation varied with seasons, although fruit production was synchronous and available all year round. From the 2635 faecal samples analysed, 79.2% contained M. leo-errerae fruit seed. Chimpanzee diet in Kalinzu is 75% frugivorous, 37.2% of which is solely contributed by M. leo-errerae fruit. The continuous availability of M. leo-errerae fruit makes it the most important food for chimpanzees in this forest, especially during general fruit scarcity there by joining figs in importance for chimpanzee survival in tropical Africa.     

Tree fruiting phenology in Kalinzu Forest, Uganda

The spatial and temporal variations in species composition and abundance of trees that fruited in mechanically logged, intensively pit‐sawn and essentially undisturbed forest areas in Kalinzu Forest (0°17′S, 0°30′S and 30°00′, 30°07′E) were assessed. The duration and frequency of fruiting episodes of selected pioneer, understorey and canopy tree species were also determined. These phenology aspects were monitored for 18 months (between February 1997 and July 1998) in 99 plots, each 20 m × 20 m in size. Eighty species consisting of 1489 trees fruited during the study. Most of the species were recorded in the undisturbed forest area and least in the mechanically logged areas. Monthly number of individuals and species that fruited in each forest condition were significantly related to monthly rainfall. Two pioneer species (Musanga leo‐errerae and Trema orientalis), one sub‐canopy (Funtumia africana) and canopy (Parinari excelsa) species had extended fruiting phases (>5 months). Two understorey species (Oxyancius speciosus and Tabernaemontana spp.) had an extended phase while the third (Teclea nobilis) had a short (≤5 months) fruiting phase. Of the selected species, only Strombosia scheffleri (a sub‐canopy species) and P. excelsa had more than one fruiting episode.     

Fruiting phenology of fig trees in Kalinzu Forest,Uganda

The study determined the abundance and species composition of fig trees that fruited in the different forest types of Kalinzu Forest Reserve (KFR), Uganda. It also assessed the seasonal variations in abundance and species composition of the fig trees, the relationships between the fruiting patterns and rainfall and the figs’ inter‐ and intraspecific patterns of fruiting episodes. Sixteen fig species represented by 515 individuals were monitored monthly from December 2007 to January 2010. Most individuals and species that fruited were in the secondary forest types (the Musanga‐ and Parinari‐dominated secondary forests) and abundances of individuals of the different species were significantly associated with particular forest types. One colonizing species (Ficus sur) was the most abundant species that fruited and was mostly recorded in the secondary forests. Species composition and abundances of trees that fruited varied seasonally, and only the abundances of two canopy species (Ficus lingua and Ficus sansibarica) were significantly related with monthly rainfall. Most species experienced at least four fruiting phases, and F. sur displayed the longest episode covering 22 months. The results suggest that the past intensive logging in KFR promoted the regeneration of a diversity of fig species, and most species generally experience community‐wide asynchronous fruiting.     

Ranging Patterns of Chimpanzees in a Montane Forest of Kahuzi,Democratic Republic of Congo

I studied ranging patterns of a semihabituated unit-group of chimpanzees for 60 mo at Kahuzi. They had a total home range of 12.81 km² and a mean annual home range of 7.55 km². Considering the low density of chimpanzees in the area vis-à-vis chimpanzees in arid areas, their home range is very small. Kahuzi chimpanzees used the home range in a clumped pattern, frequently visiting the core area and only rarely entering peripheral areas. The monthly range changes with fruit availability, increasing during periods of fruit scarcity. There was no consistent seasonal difference in the size of the home range. However, use of different habitat types may vary seasonally. While there was no seasonal effect in the use of primary forest, the chimpanzees showed a statistically consistent seasonal difference in their use of secondary forest, visiting it mainly during the dry season when fig trees were in fruit. Since the primary forest provides them with more food fruits, chimpanzees tended to use more frequently the small patches of primary forest in their home range. Thus, the size and distribution of small fragmented primary forests may be an important factor influencing the ranging pattern of chimpanzees at Kahuzi.     

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.     

Censusing chimpanzees in the Budongo Forest,Uganda

It is difficult to make valid comparisons of chimpanzee densities among sites because observers calculate them using different methods. We argue that nest count estimates of density are preferable to densities from home range estimates because of the problems of defining home range. There are many problems associated with nest count methods, some of which have not been addressed in previous studies. In 1992, we censused chimpanzees in the Budongo Forest using three methods;the standing crop nest count (SCNC), the marked nest count (MNC), and visual sightings of the animals (VS). Each method is based on standard line transect techniques. Of 96 nests monitored for decay rate,those constructed in the dry seasons decayed faster than those in the wet seasons. All- day follows of individual chimpanzees and observations of nesting chimpanzees at dusk showed that about 15.8% of night nests were reused,17.5% of the population did not build nests, and 18.8% of nests were first constructed as day nests. Given the variability in nest decay rates, we argue that MNC is a better method than SCNC because it avoids having to calculate decay rates.     

The effect of human disturbance on tree species composition and demographic structure in Kalinzu Forest Reserve,Uganda

Tree abundance and species composition in the mechanically logged, intensively pitsawn and minimally disturbed areas of Kalinzu Forest Reserve were determined. The spatial and diameter size‐class distribution of ten selected tree species representing pioneer, secondary colonizer, understorey, canopy, dominant and endangered species were also assessed. The species were: Musanga leo‐errerae, Trema orientalis, Funtumia africana, Strombosia scheffleri, Oxyansus speciosus, Parinari excelsa, Tabernaemontana holstii, Newtonia buchananii, Lovoa swynnertonii and Entandrophragma angolense. The mean stem density of all trees ≥0.5 m in height was 2809.1 per hectare and 150 species were enumerated in the three forest areas. Most individuals (47.73%) and species (75.0%) were recorded in the minimally disturbed and least in the mechanically logged areas. Of the selected species, F. africana was the most abundant (n = 789) and widely distributed in each of the areas. Entandrophragma angolense was the least abundant (n = 63) and most of its individuals (74.6%) were recorded in the minimally disturbed area. Funtumia africana and S. scheffleri (subcanopy), O. speciosus and T. holstii (understorey), N. buchananii (canopy) and P. excelsa (an upper canopy and a climax species of this forest reserve) had an inverse J‐shaped diameter size‐class structure while pioneer species (M. leo‐errerae and T. orientalis) had a bell‐shaped size‐class structure.     

Dietary selection of L'Hoest monkeys in Kalinzu forest reserve, southwestern Uganda

Reports on patterns of dietary selection of forest guenons tend to be general and often classify Cercopithecus l’hoesti as a frugivores–folivores species just like the other guenons, and to date there has been no systematic investigation into the inventory of the species’ food items in the remaining forest fragments to guide major conservation and management decisions. Here we report on the L’Hoest monkeys’ main food species and how the selection of these foods varies among different age groups of the troop in Kalinzu Forest, Uganda for a period of 12 months. Behavioural observations were determined using scan samples [Behaviour, 49 (1974) 227]. Two hypotheses were evaluated in this study. Results indicate that individuals of different age groups have the ability to select a variety of both plant and invertebrate food items in the vicinity to supplement their diets. There was no significant difference in dietary selection between different age groups of L’Hoest monkey for a particular food type (ANOVA: Column analysis, F_3,84 = 1.541337, P = 0.209827). However, significant difference was realized in dietary selection of the different food types by a particular age group (ANOVA: F_22,46 = 40.86429, P = 3.69 × 10^?23). Selective predation pressure against the infants and juveniles by Stephanoaetus coronatus was believed to be one of the likely reasons why infants and juveniles least exposed themselves in the trees feeding on fruits of Musanga leo‐errerae and Ficus spp. Invertebrate feeding was found to constitute high percentage (47.2%) of the species diet and plant materials comprised (52.8%) than previously reported in other study sites. These data provide potentially useful insight on the feeding ecology of the little studied C. l’hoesti and can provide baseline information on conservation of its food items in the remaining forests.     

A quantitative comparison of terrestrial herbaceous food consumption by Pan paniscus in the Lomako Forest,Zaire, and Pan troglodytes in the Kibale Forest,Uganda

Differences in the social organization and dental morphology of Pan paniscus (bonobos) and Pan troglodytes (chimpanzees) have been related to differences in the spatiotemporal availability of food and its exploitation. The presence of abundant terrestrial herbaceous vegetation (THV) in the bonobo's habitat and the apparent greater reliance on herbs for food has been used to explain differences in party size and, by extension, social organization. Using fecal analysis, we assess quantitatively the amount of herbaceous foods consumed by Pan paniscus in the Lomako Forest, Zaire, compared to similar data for Pan troglodytes in the Kibale Forest, Uganda. We examine this data in the context of spatiotemporal patterns of availability of herbaceous foods and fruit, as well as their nutritional content. The results support the suggestion that bonobos consume more herbaceous food than do the Kibale chimpanzees and that these foods are more prevalent in the bonobo's habitat than in the Kibale Forest. However, temporal changes in fruit availability and herb consumption, along with nutritional analyses, suggest that chimpanzees consume herbs as a fallback source of carbohydrates, whereas bonobos consume herbs as a source of protein regardless of season or fruit abundance. Available data suggest that party size while feeding on terrestrial herbs is restricted at both sites, but a determination of the relative strength of this constraint is not possible at this time. Difficulties in methods used for data collection are discussed and areas where more information is needed are highlighted. © 1994 Wiley-Liss, Inc.     

Honey‐Making Bee Colony Abundance and Predation by Apes and Humans in a Uganda Forest Reserve1

Honey‐making bee colonies in Bwindi Impenetrable National Park were investigated with Batwa Pygmies locating 228 nests of Apis and five stingless bees (Meliponini). The relative importance of predation, food supply, nesting site, and elevation affecting abundance were studied for meliponines in particular. Nest predation and overall nest abundance had no correlation with elevation along a 1400 m gradient, nor did flowering phenology or pollen collection. Many suitable, large trees were unoccupied by bee nests. In 174 ha of forest plots, 2 Meliponula lendliana, 13 M. nebulata, 16 M. ferruginea, 16 M. bocandei, and 20 Apis mellifera adansonii nests occurred, suggesting a habitat‐wide density of 39 nests/km². Compared to other studies, Ugandan Meliponini were uncommon (0.27 colonies/ha, tropical mean = 1.9/ha), while Apis mellifera was numerous (0.12 nests/ha, tropical mean = 0.06/ha), despite park policy allowing humans to exploit Apis. Meliponine colony mortality from predators averaged 12 percent/yr and those near ground were most affected. Tool‐using humans and chimpanzees caused 82 percent of stingless bee nest predation. Selective factors affecting nest heights and habit may include auditory hunting by predators for buzzing bees, and indirect mutualists such as termites that leave potential nesting cavities. Mobility and free‐nesting by honey bee colonies should enable rapid community recovery after mortality, especially in parks where human honey hunting is frequent, compared to sedentary and nest‐site‐bound Meliponini.     

Frequent insectivory by two guenons (Cercopithecus lhoesti and Cercopithecus mitis) in the Kalinzu Forest, Uganda

I investigated the feeding ecology of lHoests monkeys (Cercopithecus lhoesti) and blue monkeys (Cercopithecus mitis) in the Kalinzu Forest, Uganda. Although forest guenons are generally thought to be frugivores or folivores, these two guenons spent a large proportion of their time feeding on invertebrates. The lHoests monkey and the blue monkey spent as much as 66 and 50% of their time on insectivory, respectively. These proportions of time spent on invertebrate feeding are higher than those reported elsewhere for forest guenons. LHoests monkeys mainly utilized the area near the ground for invertebrate feeding, while blue monkeys utilized the area around 20 m above the ground. It seemed that guenons have an ability to change their diet according to the environment.     

Soils Consumed by Chimpanzees of the Kanyawara Community in the Kibale Forest,Uganda

We previously reported on a study of 4 soils that chimpanzees of the Kanyawara community in the Kibale National Park, Uganda consumed on a near-daily basis. We suggested that iron was a possible chemical stimulus in association with high quantities of Si:Al = 1:1-dominated clay minerals in the consumed material. To test our initial findings, we analyzed 18 samples from the same general area including 7 samples that the chimpanzees did not eat. Among the chemical elements, As, Au, Br, Ca, Cl, Dy, Mg, Ni, Sb, Sr, and I are below detection limits. Only Fe stands out as a potentially important nutritional element present in sufficient quantity to provide a physiological stimulus for chimpanzees living at high elevations near the flanks of the Ruwenzori Mountains. Along with Fe, metahalloysite is present in high amounts in these soils. In its pure crystalline form as a pharmaceutical grade clay mineral metahalloysite may well counteract the debilitating effects of diarrhea, with an effect similar to what is achieved with kaolinite (cf. Kaopectate^TM). An unexpected result, the relatively high nitrogen and carbon in the eaten samples relative to the uneaten group, indicates the chimpanzees may have a higher threshold for organic-rich material than previously believed. Contrarily, the color of the ingested material, depicts a material with less humus than in the uneaten group, a finding that is compatible with previous work reported at other geophagy sites in Africa. Of all the choices of soil available to them, the chimpanzees appear to be selecting highly homogeneous chemical natural earths with well-leached and uniform mineralogical material similar to the uneaten group, but with higher relative amounts of clay size material.     

Ficus sp. and Musanga leo‐errerae: coexistent keystone fruits for chimpanzees in Kalinzu forest

The composition of Ficus sp. and Musanga leo‐errerae in Chimpanzees’ diet was investigated by faecal analysis and direct observation in the medium altitude forest of Kalinzu, along the albertine rift, south‐western Uganda. The fruit availability of Ficus species showed significant variations while that of Musanga leo‐errerae was consistent and significantly higher than that of Ficus (P = 0.053; t = ?2.034) all year round. Their consumption was not opportunistic as no correlation existed between their fruit abundance and their occurrence in chimpanzee faecal samples/diet (Musanga leo‐errerae: r = 0.153, P = 0.456; Ficus sp.: r = 0.039, P = 0.848). Results showed that Musanga leo‐errerae and Ficus species seeds occurred in 80.2% and 67.2% respectively of the total 2635 chimpanzee faecal samples analyzed. Although there was no significant difference between chimpanzees party size that fed on Musanga leo‐errerae and Ficus tree species, the rate of consumption was significantly different in the low (t = 3.835; P = 0.031) than the high fruiting season (t = 2.379; P = 0.063). Ficus sp. and Musanga leo‐errerae genera function as coexistent keystone fruits for chimpanzees because they perfectly complement each other in terms of chimpanzees’ sustenance. This information has significant implications in the management of tropical forests like Kibale, Budongo, Bwindi Impenetrable, Gombe and Mahale inhabited by primate populations especially the endangered ones like the chimpanzee.     

Factors affecting Florida scrub-jay nest survival on Ocala National Forest,Florida

One of the main populations of the Florida scrub-jay (Aphelocoma coerulescens), a federally threatened species, occurs on Ocala National Forest, Florida. We determined the nest daily survival rate (DSR) of 474 nests of Florida scrub-jays in stands subject to sand pine reforestation management after timber harvesting or wildfire on Ocala National Forest. We used the information-theoretic approach with logistic-exposure modeling to determine the most likely models to account for DSR for the incubation and nestling stages separately. The models consisted of 4 components (temporal, management, habitat, and helpers) with each consisting of one to several specific variables. In the incubation stage all the best models included the temporal component alone or in concert with the habitat, helpers, or habitat and helpers components. Model averaging in the incubation stage indicated support for the year and day × year in the temporal component and stand age in the habitat component. In the nestling stage, top models all included the temporal component alone or with helpers or habitat, helpers, and the interaction of habitat and helpers. Model averaging in the nestling stage showed support for the year, nest age, and to a lesser extent day in the temporal component and helpers. The management component, which consisted of no site preparation, wildfire burn, post-harvest burn, chop and seed, or seed only, had little influence on nest survival for the incubation or nestling stages. However, we identified several other management factors that may increase Florida scrub-jay populations despite having no effect on DSR. First, the proportion of Florida scrub-jay nests was significantly higher than expected in burned habitat based on habitat availability, indicating a potential preferred nesting habitat conducive to population growth. Second, incubation stage DSR with respect to stand age (habitat component) declined to stand age 10 yr and then began to increase, which may be attributed to the higher bird population in the prime habitat in the middle stand ages. The denser population may result in more competition for resources, and possibly may attract more predators, resulting in a lower DSR. Thus, although DSR may be lower at the mid-stand ages, the overall population may actually be optimal. Therefore, to increase the Florida scrub-jay population on Ocala National Forest, we recommend maintaining a mosaic of stands ≤20 yr of age and emphasizing natural reseeding and site preparation via burning. © 2011 The Wildlife Society.     

Probable Community Transfer of Parous Adult Female Chimpanzees in the Budongo Forest, Uganda

Female chimpanzees with dependent offspring generally avoid border areas of their community's home range because they risk aggression and infanticide from extracommunity males. Typically, only nulliparous females risk crossing the boundary areas to transfer between communities; while immigration of parous females occurs, it is extremely rare and dangerous for the females and their offspring. In the Budongo Forest, Uganda, where researchers have continuously studied the Sonso community since 1990, our field data provide strong indications that ≥5 adult females with offspring have immigrated into the community. If the interpretation is correct, then it has fundamental implications for our understanding of female chimpanzee social strategies and dispersal patterns. Further, the identification of such a large number of new individuals within a short time frame is remarkable and suggests a major event, such as the breakup of a neighboring community or major habitat disturbance. We explore the evidence that points to the events as immigrations and discuss the implications for understanding the chimpanzee social system.     

Phenology of figs in Budongo Forest Uganda and its importance for the chimpanzee diet

This paper reports on the phenological patterns of figs in Budongo Forest, Uganda, and how it relates to chimpanzee food availability in different seasons. In addition, we analysed the dung of chimpanzees to understand the composition of fruits in their diet. The aim of our study was to assess Ficus phenology and how it affects chimpanzee diet. Fifteen species of figs were monitored for fruit (syconium) and leaf phenology between June 2000 and 2001. Ficus fruit production varied significantly between and within species, and also with tree trunk and crown diameters. Fig fruit production was asynchronous and individual fig trees produced crops from one to five times in a year. In addition to fruits, chimpanzees fed on young leaves of some Ficus species. Shedding of old Ficus leaves coincided with the dry season, followed by appearance of young leaves. The dry season in Budongo is a period of general fruit scarcity. The combination of fig fruits and young leaves make up the most important food in the diet of chimpanzees. From the chimpanzee dung, more than 78% of seeds comprised fig ‘seeds’ (nutlets) and the rest of the diaspores were from other tree species. Our findings suggest that chimpanzees disperse large number of diaspores in their dung, thereby serving as important agents of natural forest regeneration.