Lauren J. Chapman

Ecological constraints on group size: an analysis of spider monkey and chimpanzee subgroups

The social organization of spider monkeys (Ateles geoffroyi) and chimpanzees (Pan troglodytes) appear remarkably similar. In this paper, field studies of these two species were used to (1) test a model of ecological constraints on animal group size which suggests that group size is a function of travel costs and (2) assess ecological and social factors underlying the social organization of these two species. Spider monkeys were studied over a 6-year period in Santa Rosa National Park, Costa Rica, and chimpanzees were studied for 6 years in Kibale National Park, Uganda. Adults of both species spent their time in small subgroups that frequently changed size and composition. Thus, unlike most primate species, spider monkeys and chimpanzees were not always in a spatially cohesive social group; each individual had the option of associating in subgroups of a different size or composition. Both species relied on ripe fruit from trees that could be depleted through their feeding activity. However, spider monkey food resources tended to occur at higher densities, were more common, less temporally variable, and did not reach the low levels experienced by chimpanzees. Analyses of the relationship between subgroup size and the density and distribution of their food resources suggested that travel costs limit subgroup size. However, these ecological factors did not influence all age/sex classes equally. For example, the number of adult males in a subgroup was a function of food density and travel costs. However, this was not the case for female chimpanzees, suggesting that the benefits of being in a subgroup for females did not exceed the costs, even when ecological conditions appeared to minimize subgroup foraging costs. Therefore, it seems likely that social strategies influenced the relationship between food resource variables and subgroup size.

Estimators of Fruit Abundance of Tropical Trees1

Many types of biological studies require the estimation of food abundance in tropical forests, and a variety of methods have been used to estimate this parameter. Here we compare the accuracy and precision of three methods for estimating the fruit abundance (biomass and number) of tropical tree species: tree diameter, crown volume, and visual estimation. Diameter at breast height (DBH) was the most consistently accurate method and exhibited low levels of interobserver variability. Generally, crown volume was neither precise nor accurate. The visual estimation method was accurate for trees with very large fruit, but exhibited high interobserver variability.

Biodiversity and Fishery Sustainability in the Lake Victoria Basin: An Unexpected Marriage?

Abstract Lake Victoria is Africas single most important source of inland fishery production. After it was initially fished down in the first half of the 20th century, Lake Victoria became home to a series of introduced food fishes, culminating in the eventual demographic dominance of the Nile perch, Lates niloticus. Simultaneously with the changes in fish stocks, Lake Victoria experienced dramatic changes in its ecology. The lake fishery during most of the 20th century was a multispecies fishery resting on a diverse lake ecosystem, in which native food fishes were targeted. The lake ended the century with a much more productive fishery, but one in which three species—two of them introduced—made up the majority of the catch. Although many fish stocks in Lake Victoria had declined before the expansion of the Nile perch population, a dramatic increase in the population size of Nile perch in the 1980s roughly coincided with the drastic decline or disappearance of many indigenous species. Now, two decades after the rise of Nile perch in Lake Victoria, this species has shown signs of being overfished, and some of the native species that were in retreat—or even thought extinct—are now reemerging. Data on the resurgence of the indigenous species suggest that heavy fishing of Nile perch may enhance biodiversity; this has spawned renewed interest in management options that promote both fishery sustainability and biodiversity conservation.

A long-term evaluation of fruiting phenology: importance of climate change

Within the last decade the study of phenology has taken on new importance because of its contribution to climate-change research. However, phenology data sets spanning many years are rare in the tropics, making it difficult to evaluate possible responses of tropical communities to climate change. Here we use two data sets (1970-1983 and 1990-2002) to describe the fruiting patterns of the tropical tree community in Kibale National Park, Uganda. To address variation in spatial patterns, we describe fruiting over 2-3 y among four sites each separated by 12-15 km. Presently, the Kibale region is receiving c. 300 mm more rain than it did at the start of the century, droughts are less frequent, the onset of the rainy season is earlier, and the average maximum monthly temperature is 3.5 ◦ C hotter than it was 25 y ago. The 1990-2002 phenology data illustrated high temporal variability in the proportion of the populations fruiting. Interannual variation in community-wide fruit availability was also high; however, the proportion of trees that fruited has increased over the past 12+ y. At the species level a variety of patterns were exhibited; however, a number of the most common species currently rarely fruit, and when they do, typically < 4% of the individuals take part in fruiting events. Combining the data set from 1990 to 2002 with that from 1970 to 1983 for specific species again reveals an increase in the proportion of trees fruiting between 1990 and 2002; however, the proportion of the populations fruiting decreased during the earlier period. When one examines particular species over this whole period a variety of patterns are evident. For example, Pouteria altissima exhibited a relatively regular pattern of fruiting during the 1970s; however, it rarely fruited in the 1990s. Contrasting phenological patterns at four sites revealed that at the community level the fruiting patterns of only one of the six pair-wise site combinations were correlated. Relationships between rainfall and fruiting were variable among sites. Contrasting changes in fruiting patterns over the 30 y with differences among the four sites varying in rainfall, suggests that the changes observed in fruiting may be due to climate change. Responses to this climate change are likely complex and will vary among species. However, for some species, current conditions appear unsuitable for fruiting.

Fruit and flower phenology at two sites in Kibale National Park, Uganda

Examination of phenological patterns of tropical trees at different temporal and spatial scales can elucidate biotic and abiotic factors that correlate with fruiting, flowering and/or leaf set patterns. In this study, 3793 trees from 104 species in Kibale National Park, Uganda were monitored. The trees were selected from two sites (Kanyawara and Ngogo) separated by 10 km. Trees were monitored monthly to document community-wide and population-level fruiting and flowering patterns for a maximum of 76 mo. Analysis of two sites over a number of years permitted examination of generalities of patterns found on smaller spatial and temporal scales. Spectral analysis indicated that community-level flowering and fruiting at Kanyawara exhibited regular annual peaks, although the flowering peaks were of shorter duration. At Ngogo, community-level flowering also dis- played regular annual peaks, but fruiting had an irregular pattern with no distinct peaks. The abundance of fruiting trees at Kanyawara was negatively related to the minimum temperature in the previous season (3-7 mo prior). Since fruiting tended to peak when the first wet season of the year was ending and the dry season was beginning, this suggests that the minimum temperature in the previous dry season is important in determining how many individuals fruit. Flowering at Kan- yawara peaked immediately after the maximum annual period of high irradiance. Within-species synchronization was evident in the flowering for all species exam- ined at Ngogo and for 64% of those at Kanyawara. Fruiting was synchronous within species for 64% of the species at both sites. Despite this general community-level synchronization, the months of peak fruiting and flowering for some species varied markedly among years. Furthermore, for a number of species the timing of fruiting or flowering events differed between Kanyawara and Ngogo. For some species, trends that were suggested from one year of data were not supported when addi- tional years were considered. Although these two sites are close together, share many of the same species, and experience similar climatic regimes, many phenolo- gical patterns were site-dependent.

Physiological refugia: swamps, hypoxia tolerance and maintenance of fish diversity in the Lake Victoria region

In Lake Nabugabo, Uganda, a satellite of Lake Victoria, approximately 50% of the indigenous fishes disappeared from the open waters subsequent to the establishment of the introduced predatory Nile perch, Lates niloticus. This pattern is similar to the faunal loss experienced in the much larger Lake Victoria. Several of these species persisted in wetland refugia (e.g. ecotonal wetlands, swamp lagoons); however, deep swamp refugia (habitats lying well within the dense interior of fringing wetlands), are available only to a subset of the basin fauna with extreme tolerance to hypoxia. Although air-breathers are common in deep swamp refugia; we also documented a surprisingly high richness and abundance of non-air-breathing fishes. We describe several mechanisms that may facilitate survival in deep swamp refugia including high hemoglobin concentration, high hematocrit, large gill surface area and a low critical oxygen tension (P(c)). In addition, swamp-dwelling fishes showed lower PO(2) thresholds for onset of aquatic surface respiration than the lake-dwelling fishes. This suggests higher tolerance to hypoxia in the swamp fishes because they are able to withstand a lower oxygen tension before approaching the surface. We suggest that physiological refugia may be important in modulating the impact of Nile perch and indigenous fishes in the Lake Nabugabo region; this highlights the need to evaluate relative tolerance of introduced predators and indigenous prey to environmental stressors.

Indices of Habitat-wide Fruit Abundance in Tropical Forests1

The assessment of fruit abundance is critical for studies of frugivore ecology. A variety of methods have been used to estimate habitat-wide fruit abundance. However, since the methods have not been calibrated with each other, it is difficult to compare results of different studies. Here we compare three methods used simultaneously to collect fruit abundance data in the Kibale Forest, Uganda. Estimates of fruit abundance derived from fruit traps were not correlated with estimates derived from either systematic transect sampling or estimates obtained from observing fruiting phenology of key species on a fruit trail. However, estimates based on fruit trail data and transect data were correlated. We review the advantages and disadvantages of methods that have been used to assess habitat-wide fruit abundance.

Nuclear markers reveal unexpected genetic variation and a Congolese-Nilotic origin of the Lake Victoria cichlid species flock

Phylogenetic analyses based on mitochondrial (mt) DNA have indicated that the cichlid species flock of the Lake Victoria region is derived from a single ancestral species found in East African rivers, closely related to the ancestor of the Lake Malawi cichlid species flock. The Lake Victoria flock contains ten times less mtDNA variation than the Lake Malawi radiation, consistent with current estimates of the ages of the lakes. We present results of a phylogenetic investigation using nuclear (amplified fragment length polymorphism) markers and a wider coverage of riverine haplochromines. We demonstrate that the Lake Victoria–Edward flock is derived from the morphologically and ecologically diverse cichlid genus Thoracochromis from the Congo and Nile, rather than from the phenotypically conservative East African Astatotilapia. This implies that the ability to express much of the morphological diversity found in the species flock may by far pre–date the origin of the flock. Our data indicate that the nuclear diversity of the Lake Victoria–Edward species flock is similar to that of the Lake Malawi flock, indicating that the genetic diversity is considerably older than the 15 000 years that have passed since the lake began to refill. Most of this variation is manifested in trans–species polymorphisms, indicating very recent cladogenesis from a genetically very diverse founder stock. Our data do not confirm strict monophyly of either of the species flocks, but raise the possibility that these flocks have arisen from hybrid swarms.

Application of protein-to-fiber ratios to predict colobine abundance on different spatial scales

The biomass of arboreal folivorous primates in Africa and Asia is related to an index of mature leaf quality: the ratio of protein-to-fiber concentration. Investigations have considered variation in folivore biomass and forest composition among sites separated by hundreds or thousands of km. However, large variation in folivore abundance has been documented over much smaller spatial scales. We quantify the degree to which the average protein-to-fiber ratio of mature leaves of the 20 most abundant tree species predicts the biomass of western red colobus (Piliocolobus trephosceles) and black-and-white colobus (Colobus guereza) over very small spatial scales. Four sites within Kibale National Park, Uganda, varied markedly in forest structure. Colobine biomass varied among sites from 191 to 2675 kg/km2 and was related to the average protein-to-fiber ratio of mature leaves of the 20 most abundant tree species at each site. We examined the generality of the relationship between protein-to-fiber ratios and colobine abundance by adding our biomass and leaf chemistry values to previously published values to produce 9 comparable sites. At these locations, colobine biomass varied from 84 to 2675 kg/km2 (mean biomass among sites = 910 kg/km2), and mean protein/fiber ratios varied from 0.167 to 0.577. Colobine biomass was related to the protein-to-fiber ratios of mature leaves (R2 = 0.616, P = 0.012).

Foraging challenges of red colobus monkeys: influence of nutrients and secondary compounds

The diet selection of two groups of red colobus monkeys (Procolobus badius) in Kibale National Park, Uganda are considered with respect to protein, fiber, digestibility, alkaloids, total phenolics, tannins, saponins, and cyanogenic glycosides. Both groups selected young leaves over mature leaves and young leaves had more protein, were more digestible, and had a higher protein to fiber ratio than mature leaves. Young and mature leaves did not differ with respect to secondary compounds. There were no differences in the phytochemical factors examined between frequently eaten foods and leaves that red colobus were never known to eat, but were relatively common in the environment. Regression analyses predicting foraging effort from the phytochemical components of the large groups diet revealed selection for only one factor, foods that are high in protein and low in fiber, when differences in food tree availability were taken into consideration. A similar analysis with the small group did not suggest selection or avoidance of foods with respect to any of the factors considered. Previous studies have found the biomass of folivorous primates to be related to the ratio of protein to fiber concentration of mature leaves in the environment. These investigations have considered variation in folivore biomass and forest composition among sites separated by hundreds of kilometers; however, large variation in folivore abundance occurs over much smaller spatial scales. In Kibale National Park the average protein to fiber ratio of the mature leaves of the 20 most abundant tree species predicted the biomass of red colobus among four neighboring sites. We examined the generality of this relationship by adding our biomass and leaf chemistry values to previously published values; 62% of the variance in colobine biomass was explained by variation in the protein to fiber ratios of mature leaves at the sites. There was no evidence that red colobus avoided plants with high levels of secondary compounds. In fact, one of the most preferred trees (Prunus africana) was the species with the highest levels of cyanogenic glycosides, and the highest saponin levels were found in the young leaves of Albizia grandibracteata, the sixth and fourth most preferred plant species for the large and small groups, respectively.