High interannual variation in the diet of a tropical forest frugivore (Hylobates lar)

Frugivores must deal with seasonal changes in fruit availability and changes from year to year, as most species of tropical forest fruiting trees have considerable interannual variation in phenology and many are mast fruiters. We quantified seasonal and interannual changes in the fruit diet in a frugivore and important seed disperser, the white‐handed gibbon, Hylobates lar, in Thailand. We used 40‐d following data during April and May replicated in six consecutive years to study interannual variability in the diet and compared it with seasonal changes measured in monthly samples of the same size collected in three successive years. The 40‐d periods of following also allowed us to measure the decline in dietary similarity with time over a finer scale. We measured fruit diet similarity between replicated 5‐d periods using the percentage overlap (Renkonen's) index and Jaccard's similarity index. Seasonally, average dietary overlap between adjacent months was low, and similarity approached zero after four months. Average rate of decline in similarity exceeded 20 percent per 5‐d period. Variation in fruit species in the diet between years was high and was correlated with interannual variation in fruiting phenology. The strongest correlation occurred in the case of Nephelium melliferum, a highly preferred species that dominated the diet in good fruiting years. It is difficult to separate changes in food species preference from changes in availability from year to year. We devised a relative measure of preference that depends on the degree to which the gibbons rely on prior knowledge to find sources.     

Frugivory in sun bears (Helarctos malayanus) is linked to El Niño-related fluctuations in fruiting phenology, East Kalimantan, Indonesia

Sun bear ( Helarctos malayanus ) frugivory and fruiting phenology was investigated in a lowland dipterocarp forest in East Kalimantan, Indonesia. Two mast fruiting events, both coinciding with El Niño/Southern Oscillation events, occurred 4 years apart, resulting in large fluctuations in fruit availability. Sun bear fruit availability decreased from 13 trees ha⁻¹ fruiting month⁻¹ during the mast fruiting to 1.6 trees ha⁻¹ fruiting month⁻¹ during the intermast period. Almost 100% of sun bear diet consisted of fruit during mast fruiting period, whereas sun bear diet was predominantly insectivorous during intermast periods. The majority of sun bear fruit trees displayed 'mast-fruiting' and 'supra-annual' fruiting patterns, indicating sporadic productivity. Sun bears fed on 115 fruit species covering 54 genera and 30 families, with Ficus (Moraceae) being the main fallback fruit. The families Moraceae, Burseraceae, and Myrtaceae contributed more than 50% to the sun bear fruit diet. Sun bear fruit feeding observations were unevenly distributed over forest types with more observations in high-dry forest type despite fewer fruiting events, possibly due to a side-effect of high insect abundance that causes bears to use these areas more intensively. The possible evolutionary pathways of sun bears in relation to the Sundaic environment are discussed.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 489–508.     

Inter‐annual variability of fruit timing and quantity at Nouragues (French Guiana): insights from hierarchical Bayesian analyses

The timing and quantity of fruit production are major determinants of the functioning of a forest community, but simultaneous analyses of both are rare. We analyzed a ten‐year dataset (2001–2011) of fruit production for 45 tree and liana species from the Nouragues rain forest, French Guiana. We developed a hierarchical Bayesian approach to determine variation in the timing and quantity of fruit production. Our analysis accommodates missing censuses and quantifies variation at seasonal and inter‐annual scales. The fruiting peak of 22 of 45 species occurred during the peak of the rainy season, which is typical for central and eastern Amazon. The timing and quantity of fruit production varied substantially across years in most species, with greater variation in quantity than in timing. The timing of fruit production varied from continuously fruiting species to mast fruiting species that had two or more consecutive years without fruit production. Fully 40% of species were mast fruiting species. The seasonal timing and inter‐annual variation in fruiting were unrelated to seed dispersal mode across species. We saw no evidence for directional change in the level of fruit production, the timing of fruit production, or their variances; however, 10 yr is a short record for such analyses.     

Individual–resource networks reveal distinct fruit preferences of selective individuals from a generalist population of the Helmeted Manakin

Ecological studies traditionally assume that generalist populations are homogeneous in the use of food resources, but empirical evidence supports that intraspecific differences in morphology, physiology and behaviour affect foraging decisions and promote diet variation among individuals. Furthermore, the temporal availability of resources may shape the dynamics of population trophic niche, which ultimately depends on individual niches. In this study, we investigated the seasonal changes in individual-based networks between the Helmeted Manakin Antilophia galeata, a generalist frugivorous bird, and fruiting plants, following theoretical models of interindividual diet variation based on the Optimal Diet Theory. Selective individuals were the majority of the generalist population of the Helmeted Manakin. Our results suggest that the structure of the individual-resource networks varied seasonally. We found that modularity was higher than expected by chance in the wet season, when fruit availability was also higher. In the dry season, modules disappeared and the network became more nested. These findings are consistent with the Distinct Preference Model of diet variation. We suggest that downscaling ecological networks to the individual level may reveal emergent properties that, albeit existent, are not evident in species–resources networks.     

Floristic heterogeneity at Ngogo,Kibale National Park,Uganda and possible implications for habitat use by chimpanzees (Pan troglodytes)

Tropical forest tree communities exhibit heterogeneity at multiple spatial scales, with important implications for animals relying on these resources. However, different organisms may perceive heterogeneity in the floristic community in very different ways. Here, we characterize the overall extent of heterogeneity in the floristic community at Ngogo, in Kibale National Park, Uganda. Also, using information from studies on the diet of chimpanzees (Pan troglodytes) inhabiting Ngogo, we quantify the extent to which the habitat is likely perceived as heterogeneous by this species. The forest as a whole is slightly more diverse than comparable sites in the Congo Basin, but this diversity is driven by a relatively high proportion of rare species represented by few individuals. From the perspective of chimpanzees, the forest is, unsurprisingly, even more heterogeneous. Species that provide fruit for chimpanzees during times of low overall fruit abundance and that display interindividual synchrony in fruiting were the most common chimpanzee resource in our sample, whereas species that provide fruit during times of low overall fruit abundance and that display asynchronous fruiting were the least common. We discuss the implications of the differences in density and distribution of various classes of resources for chimpanzee habitat use and foraging efficiency.     

Feeding ecology of western lowland gorillas in the Nouabalé-Ndoki National Park,Congo

The feeding ecology of western lowland gorillas (Gorilla gorilla gorilla) living in the Nouabalé-Ndoki National Park, northern Congo, was surveyed for one full year. This is the first record to make clear the seasonal changes in the feeding habits of gorillas in a whole year, living in the primary lowland forest almost completely undisturbed. Fecal contents, feeding traces, and direct observation were analyzed with reference to a fruit availability survey. Although the gorillas fed largely on fruits in the forest, their basic diet was fibrous parts of plants, including shoots, young leaves, and bark. Terrestrial herbaceous vegetation, such as monocotyledons of the Marantaceae and aquatic herbs having much protein content and minerals, were frequently eaten even in the fruiting season. As these highly nutritious fibrous foods were superabundant all year, the major foods of the Ndoki gorillas seemed to be those plants. However, they selected fruits as their alternative food resources in the fruiting season. Gorillas foraged on many fruit species, while showing strong preferences for some particular species. The swamp forest, including marshy grasslands, was an important and regular habitat for the Ndoki gorillas.     

Comparison of frugivory by howling monkeys (Alouatta palliata) and bats (Artibeus jamaicensis) in the tropical rain forest of Los Tuxtlas,Mexico

Information on the fruit diets of howling monkeys and fruit-eating bats in the tropical rain forest of Los Tuxtlas, Mexico was collected for a year to compare the plant species used. Howling monkeys used 19 plant species whereas bats used 32 plant species as fruit sources. Eleven species were common in the diet of both mammals. A rank analysis at the plant species level showed that the fruit diets of Alouatta and Artibeus were very different. In contrast to bats, howling monkeys displayed a seasonal pattern in fruit consumption. Diet overlap between the two mammals was highest during the monkeys' fruit-eating season. Measures of fruit production in eight trees (four species) indicated marked variations in fruit biomass produced and in length of fruiting from tree to tree and species to species. Peaks in fruit production were typical both at the species and the individual tree levels, demonstrating the very patchy nature of the fruit available to the monkeys and the bats.     

Seasonal shifts in large mammal activity relative to fruit availability and hunting risk along a protected area boundary in Borneo

Understanding wildlife spatiotemporal dynamics at protected area boundaries is critical to conservation. In SE Asia, protected areas are often bordered by indigenous communities whose traditional practices result in increased landscape heterogeneity within their community managed forests (CF). Because SE Asian forests exhibit supra-annual mast fruiting (3–7 yrs) and sustained fruit scarcity, wildlife using CF may benefit from greater availability of fruits and seeds encouraged by traditional management, but incur greater risk of being hunted. We examined shifts in wildlife activity in forests within and adjacent to Lanjak Entimau Wildlife Sanctuary (LEWS), in Sarawak, Malaysia. Using camera traps and surveys of fruiting phenology, we related presence of fruits and seeds to shifts in large mammal activity over 3 years (October 2016–2019). We first compared relative frequency of five focal species (sun bear, bearded pig, muntjac, mouse deer, and sambar) by forest type for a given survey year and season (rainy vs. dry). We then modeled activity to assess how fruit availability, landscape features, and proxies for hunting risk, influence seasonal shifts in habitat use. Our results suggested seasonal variation in concert with increased landscape heterogeneity influenced patterns of activity. We found no evidence that CF excluded any detected mammal species, suggesting CF contribute important habitat to wildlife, particularly during periods of fruit scarcity in primary forest. However, our findings also imply increased hunting risk for wildlife attracted to resources near human settlements, emphasizing the need to consider the influence of both annual and supra-annual seasonality when developing best practices for the sustainable management.     

Fruit availability, chimpanzee diet, and grouping patterns on Rubondo Island, Tanzania

We examined seasonal patterns of fruit availability, dietary quality, and group size in the descendants of an introduced chimpanzee population on Rubondo Island, Tanzania. The site has supported a free-ranging population without provisioning for 40 years. Our goals were to determine whether Rubondo chimpanzees experience periods of fruit shortage, and whether they respond to changes in fruit availability similarly to chimpanzees at endemic sites. We indexed the fruit availability of tree and liana species on transects stratified across three chimpanzee ranging areas. We used fecal analyses to evaluate seasonal changes in diet, and used data on party size and nesting group size to examine seasonal patterns of grouping. Tree fruit availability was positively correlated with rainfall, with a period of relative tree fruit scarcity corresponding with the long dry season. Liana fruit availability was not related to rainfall, and lianas exhibited less variable fruiting patterns across seasons. Fruits made up the majority of the chimpanzee diet, with lianas accounting for 35% of dietary fruit species. Fruits of the liana Saba comorensis were available during all months of phenological monitoring, but they were consumed more when tree fruit was scarce, suggesting that Saba comorensis fruits may be a fallback food for Rubondo chimpanzees. There were no increases in consumption of lower-quality plant parts between seasons, and there were no changes in group size between seasons. These results contrast with evidence from several endemic chimpanzee study sites, and indicate that Rubondo chimpanzees may have access to abundant and high-quality foods year round.     

Dietary variability in Cebus apella in extreme habitats: evidence for adaptability

The dietary composition of Cebus apella in two subtropical forest environments of Argentina reflects the seasonal availability of potential food resources. In the marginal areas of the geographical distribution of this species, different resources are used during periods of scarcity. A greater tendency to use leaves in the diet is observed in the northwest (El Rey National Park), where there is greater seasonal variation of fruit availability. In El Rey, where there are few potential resources, the diet is dominated by a few resources, notably bromeliads. Where the resources are more abundant and the availability is more constant throughout the years (Baritú and Iguazú National Parks), the monkeys exhibit a feeding behavior similar to that seen in tropical areas. This ability of the genus Cebus to exploit resources not accessible to other primate species is one of the reasons for its wide geographical distribution and its widespread existence in ecosystems marginally used by primates.     

Resource tracking and its conservation implications for an obligate frugivore (Procnias tricarunculatus,the three‐wattled bellbird)

In Monteverde, Costa Rica, the vulnerable Three‐wattled Bellbird (Procnias tricarunculatus) feeds primarily upon the fruit of Lauraceae species during its reproductive and post‐reproductive seasons. To understand and advance appropriate conservation measures, this study identified the bellbird's foraging challenges in its search for a temporally and spatially fluctuating resource. Although there are at least 96 species of Lauraceae found in the five life zones of Monteverde, the distinct distributions of tree species both among and within life zones require the bellbirds to track seasonal fruiting across the various zones. In this 6‐year study, we monitored the fruiting of tree species and bellbird abundance in 24 study plots within its post‐reproductive life zone, the Premontane Wet forest, to identify preferred bellbird food resources and how the fruiting of these species drives the spatial distribution of the bellbird. Our research revealed phenological patterns of annual, biennial, and triennial fruiting with high levels of fruiting synchrony within several identified key fruit species. Of critical conservation importance is that no single species of Lauraceae produced a consistent food supply for bellbirds each year. Therefore, even within life zones, the bellbird's survival depends on its mobility to search for and obtain fruit, as well as the availability of fruits of multiple tree species. The conservation implications include focused attention on multiple core areas within given life zones, protection of existing forest and remnant trees, and forest restoration with plantings of multiple tree species. We suspect that other tropical frugivorous species face similar conservation challenges.     

Diversity of climbing seed plants and their reproductive habit in a karst seasonal rain forest in Nonggang,Guangxi, China

Aims Diversity of climbing seed plants and their reproductive habits and characteristics are central for the understanding of community structure and dynamics of forests and hence are important for forest protection. However, little is known about the climbing seed plants in northern tropical karst seasonal rain forests. Here, using the data of the species diversity and reproductive habits of climbing seed plants in Nonggang, Guangxi, China, we aim to 1) explore the species diversity and distribution of climbing seed plants in northern tropical karst seasonal rain forests, 2) study the flowering and fruiting phenology and 3) the associations of reproductive characteristics to the environment. Methods Species composition, preferred habitat, flowering time, fruiting time and fruit types of climbing seed plants were surveyed. The seasonality of flowering and fruiting were analyzed by concentration ratio and circular distribution. Climbing seed plants were divided into three groups according to their growth forms and places in spatial forest structure: bush ropes, herbaceous vines and lianas. Monthly flowering ratios, fruiting ratios, fruit types and their ratios in different groups were determined. These relationships of flowering ratio, fruiting ratio, fruit type and its ratio to meteorological factors were investigated using Pearson correlation analysis. Important findings There were a total of 333 species of climbing seed plants in Nonggang karst seasonal rain forest, belonging to 145 genera and 56 families. Bush ropes, herbaceous vines and lianas contained 119, 88 and 126 species, respectively. At species level, herbaceous vines were more abundance in valleys, while bush ropes and lianas were more abundance on slopes. Flowering and fruiting of climbing seed plants occurred seasonally, with flowering peaking in April to September, while fruiting peaking in July to December. The seasonality of flowering and fruiting in bush ropes was weaker than in herbaceous vines and lianas. Flowering ratio was significantly positively correlated with rainfall and air temperature, which suggest that flowering peaks in monsoon season. Peak time for fruiting was about three months later than the peak time of flowering, around the end of monsoon season. The ratio of samara species to all fruiting species in lianas was significantly positively correlated with wind speed, but negatively correlated with rainfall and air temperature. It showed that samara in lianas tended to occur in dry season with high wind speed. In conclusion, species diversity and the seasonal features of reproduction of climbing seed plants in Nonggang karst seasonal rain forest were closely related to the spatial and temporal variations of habitat resources.     

Quetzal Abundance in Relation to Fruit Availability in a Cloud Forest in Southeastern Mexico1

The Resplendent Quetzal (Pharomachrus mocinno) is an altitudinal migrant that nests in high elevation cloud forests and migrates toward lower areas during the summer rainy season. It has been suggested that its migratory movements are related to the abundance of ripe Lauraceae fruits. We studied the quetzal diet during two consecutive years, as well as changes in fruit abundance of the plant species on which the bird feeds at El Triunfo Biosphere Reserve, southeastern Mexico. The quetzal was observed feeding on 32 plant species; of these, 24 are new records in its diet. We chose 20 of these 32 species and studied their fruit phenology for two years in order to describe the relationship between fruit and quetzal abundance. Our results showed that quetzal abundance in the breeding area was correlated with the total number of fruiting species, whereas the correlation between quetzal abundance and the number of fruiting Lauraceae species was only marginal. Additionally, a correlation test showed that quetzal abundance was marginally correlated with total fruit availability (total no. of fruits per month); however, the correlation between quetzal abundance and the number of fruits in the Lauraceae was not significant. Our results suggest that the dynamics of food resources may be playing a major role in the quetzal's migratory behavior. Knowing the bird's diet may aid in characterizing the type of habitat adequate for its conservation. Our observations in this respect suggest that conservation efforts to preserve this bird species should concentrate on the protection of its habitat, including both breeding and nonbreeding (migration) locations.     

Global patterns in fruiting seasons

Aim   To identify geographical and climatic correlates of the timing of fruit production in fleshy fruited plant communities.
Location   Global.
Methods   We searched the literature for studies documenting monthly variation in the number of fleshy fruited species bearing ripe fruits in plant communities (i.e. fruit phenologies). From these data, we used circular vector algebra to characterize seasonal peaks in fruit production (mean date, as an angle) and the length of fruiting seasons (as a circular standard deviation). Generalized linear models and circular correlations were used to assess whether latitudinal patterns in fruit phenologies could be explained by variation in temperature, precipitation and actual evapotranspiration (AET).
Results   Dates of peak fruit production and the length of fruiting seasons showed consistent differences with latitude. Annual peaks in fruit production occurred 1 to 3 months after the summer solstice at high-latitude sites in both hemispheres. Fruiting seasonality increased with latitude, indicating that fruiting seasons were longer in the tropics and shorter toward the poles. AET was the best climatic predictor of fruit phenologies. Annual peaks in fruit production were positively associated with annual peaks in AET and temperature, while fruiting seasons were shorter in areas with pronounced annual variation in AET.
Main conclusions   Global patterns in fruiting seasons are associated with global variation in climate. Across the globe, fleshy fruits are produced after annual periods of elevated water–energy availability. Fruiting seasonality is also more pronounced in areas with strongly seasonal water–energy inputs. Therefore, the timing of reproduction in fleshy fruited plant communities appears to be determined, at least in part, by spatial and temporal variation in energy supplies needed to subsidise plant reproduction.     

Fruiting phenology of palms and trees in an Atlantic rainforest land-bridge island

Tropical forests show periods of scarcity and high fruit production in the same year and/or between years. Palms are an important component of Neotropical rainforests and a significant food resource for several frugivores. Therefore, their role as keystone resource may be exacerbated in highly impoverished areas. In Anchieta Island, São Paulo/Brazil, human settlements have modified and impoverished the forest, mainly through overharvesting and the introduction of exotic plants and several mammal species. We assessed the offer of fruits consumed by vertebrate frugivores at this island, the vegetation of which is belonging to the Brazilian Atlantic rainforest. We compared whether the fruiting patterns and fruit fall differ between palms and trees, and discuss the importance of palms as a food resource for frugivores and the implications for Anchieta Island conservation. Phenological patterns were seasonal for both trees and palms; however, the times of fruiting occurrence differed. Fruit fall biomass was at least twice lower than reported for other Atlantic rain forests and was also different between trees and palms. Palms contributed more than 80% of the overall fruit fall biomass. Palms may constitute an alternative food resource in periods of low fruit availability, although they do not provide resources for the entire assemblage of vertebrate frugivores. Energy-rich fruits, such as those produced by palms, may play an important role in the maintenance of frugivore populations in isolated, disturbed environments with a high density of vertebrate frugivores, low diversity of fruiting species and fruit biomass such as those found on Anchieta Island.     

Diet and Feeding Ecology of Chimpanzees (Pan troglodytes) in Bulindi, Uganda: Foraging Strategies at the Forest–Farm Interface

Wild animals increasingly inhabit human-influenced environments such as forest fragments amid agricultural systems. Dietary studies provide a means of assessing wildlife responses to anthropogenic habitat changes. Chimpanzees are specialist frugivores that consume other plant parts, e.g., fibrous pith and leaves, in greater amounts during fruit shortages. I examined the plant diet and seasonal foraging strategies of chimpanzees inhabiting small forest fragments within a cultivated landscape in Uganda. I determined diet over 13 mo via systematic fecal analysis, supplemented by direct observation and feeding trace evidence. I identified important foods and examined their role as seasonal fallbacks. Diet composition and breadth were overall species typical. Chimpanzees were highly frugivorous and the fruit component of fecal samples exceeded that of nonfruit fiber in all months. Forest fruit availability fluctuated seasonally, including a 3-mo low fruiting season, when overall fruit intake declined. During this time chimpanzees pursued a mixed strategy of increasing fiber consumption and feeding more heavily on energy-rich cultivars, including those obtained through crop raiding. The data suggest that exploiting agricultural fruits helped chimpanzees maintain a fruit-dominated diet when forest fruit was scarce. No evidence suggested this disturbed forest–farm mosaic is a food-impoverished habitat for chimpanzees overall. Nevertheless, cultivar feeding creates conflict with people and the high nutritional quality of crops is likely offset by the inherent risk associated with obtaining them. This study adds to growing evidence of ecological and behavioral adaptability of Pan troglodytes in response to anthropogenic habitat alteration. Targeted conservation of key natural foods for wildlife —particularly fallbacks— would help reduce conflicts and improve the survival prospects of threatened species sharing environments with people.     

Photosynthesis and carbohydrate partitioning in sweet cherry: Fruiting effects

The effect of fruiting on carbon fixation and retention in leaves was monitored by measuring net photosynthesis (Pn) and total non-structural carbohydrates (TNC) on a seasonal basis on mature fruiting and non-fruiting sweet cherry trees ( Prunus avium L. cv. Bing). Pn was also measured diurnally during stages II and III of fruit development. Pn rates increased to between 18 and 20 mg CO₂ dm^-2 h^-1 during stage II of fruit development and were maintained until harvest. Diurnally, Pn increased in the morning to 20 mg CO₂ dm^-2 h^-1 and this rate continued until sunset. Leaf carbohydrate levels decreased in both fruiting and non-fruiting trees beginning at the equivalent of stage II of fruit growth. Carbohydrates were lower in leaves and woody portions of current, 1- and 2-year-old shoots of fruiting trees. Although differences were found in levels of non-structural carbohydrates, no differences in Pn were found in fruiting vs non-fruiting plants on either a seasonal or a diurnal basis. Pn rates in swet cherry in the field were primarily affected by ontogeny and environment and not by sink strength.     

Seasonality,diaspore traits and the structure of plant-frugivore networks in Neotropical savanna forest

Complex frugivory networks are common in heterogeneous environments, but how the structure of those networks varies due to seasonality and other environmental factors remains unclear. For example, seasonal variation in rainfall can influence fruit production and diaspore characteristics, which could alter the quantity and quality of resources available to different animals in the network and, hence, network structure. We investigated how a frugivory network varied seasonally in Brazilian savanna (Cerrado), where there are well-defined dry and wet seasons and fructification mainly during the rainy season for most tree species. We recorded fruit consumption by animals during the dry and wet seasons in two different gallery forests and used these data to test the hypotheses that connectance, links per species and nestedness would be higher in the dry season than rainy season due to low available food in the former that would be consumed by various species of frugivores. Concomitantly, we also measured seed width and lipid content from diaspores of the fruiting trees to determine if these characteristics influenced interaction properties between fruiting trees and frugivores. Among the measured network parameters, connectance, links per species and specialization varied between seasons in one site but not in the other, indicating that seasonal variation in networks is not necessarily consistent over time or space. The number of tree species with small diaspores with high lipid content differed between seasons, and those characteristics were key factors increasing the interaction parameter of fruiting trees. We suggest that network stability between seasons may be related to local frugivore diversity, resource availability, and fruit quality.     

Flowering and Fruiting Patterns in a Subtropical Rain Forest,Taiwan

Reproductive patterns of tropical and temperate plants are usually associated with climatic seasonality, such as rainfall or temperature, and with their phylogeny. It is still unclear, however, whether plant reproductive phenology is influenced by climatic factors and/or phylogeny in aseasonal subtropical regions. The plant reproductive phenology of a subtropical rain forest in northern Taiwan (24°45′ N, 121°35′ E) was monitored at weekly intervals during a 7‐yr period (2002–2009). The flowering patterns of 46 taxa and fruiting patterns of 26 taxa were examined and evaluated in relation to climatic seasonality, phylogenetic constraints, and different phenophases. Our results indicated that most of the studied species reproduced annually. Additionally, both community‐wide flowering and fruiting patterns exhibited distinct annual rhythms and varied little among years. The community flowering peak matched seasonal changes in day length, temperature, and irradiance; while the community fruiting peak coincided with an increase in bird richness and the diet‐switching of resident omnivorous birds. In addition, phylogenetically closely related species tended to reproduce during the same periods of a year. Neither the mean flowering dates nor seasonal variation in solar radiation, however, was related to the fruit development times. Our results indicate the importance of abiotic, biotic, and evolutionary factors in determining the reproductive phenology in this subtropical forest.     

The Influence of Seasonal Variation on Chimpanzee (Pan troglodytes schweinfurthii) Fallback Food Consumption, Nest Group Size, and Habitat Use in Gishwati, a Montane Rain Forest Fragment in Rwanda

The increased number of primates living in fragmented habitats necessitates greater knowledge of how they cope with large-scale changes to their environment. Chimpanzees (Pan troglodytes) are exceptionally vulnerable to forest fragmentation; however, little is known about chimpanzee feeding ecology in fragments. Although chimpanzees have been shown to prefer fruit when it is available and fall back on more abundant lower quality foods during periods of fruit scarcity, our understanding of how chimpanzees use fallback foods in forest fragments is poor. We examined how chimpanzees cope with periods of fruit scarcity in Gishwati Forest Reserve, a disturbed montane rain forest fragment in Rwanda. We assessed seasonal changes in chimpanzee diet and the use of preferred and fallback foods through fecal and food site analysis. We also examined seasonal variation in nest group size and habitat use through marked nest censuses. We found that chimpanzees experienced a seasonal reduction in preferred fruit availability, which led to a seasonal diet shift to more fibrous foods, including several that functioned as fallback foods. Our results suggest that during periods of fruit scarcity the chimpanzees also reduced nest group size. However, we found that the chimpanzees did not alter their habitat use between high- and low-fruit seasons, which suggests that the small size of the forest limits their ability to change their seasonal habitat use. Consequently, fallback foods appear to be particularly important in small food-impoverished habitats with limited ranging options.