Fruit selection by Andean forest birds: influence of fruit functional traits and their temporal variation

Fruit selection, i.e., the consumption of fruits disproportionately to their availability, results from the interaction between diet preferences and ecological factors that modify them. We assessed the importance of functional fruit traits to explain fruit selection by birds in Andean subtropical forests, taking into account temporal variation in trait distribution in the assembly of available fruits. During 2 yr, we measured the abundance of ripe fruits and their consumption by birds in a 6‐ha plot during 11 bimonthly samplings, and we used 17 phenological, morphological, and nutritional traits to characterize fruits selected by four bird species. Fruit selection was pervasive year‐round, highly variable over time and across bird species. Fruit species were selected over time periods shorter than their ripening phenology, and the selection of fruits with particular traits was specific to the fruit‐eating species. Maximization in pulp reward per consumed fruit seems to be the main driving force behind fruit selection, indicating that birds select fruits with traits that directly affect net energy gain. Our results can be interpreted in a framework of a hierarchy of foraging decisions, under which the spatiotemporal context of the fruiting environment modifies the relative intake rates of a particular fruit, while the ability to discriminate fruit contents becomes increasingly important on a smaller dimension. We show that fruit‐selection properties are contingent on specific fruit traits and particular spatiotemporal conditions, which modify the structure of mutualistic interactions.     

Diet and seasonal changes in sympatric gorillas and chimpanzees at Kahuzi–Biega National Park

Based on 8 years of observations of a group of western lowland gorillas (Gorilla beringei graueri) and a unit-group of chimpanzees (Pan troglodytes schweinfurthii) living sympatrically in the montane forest at Kahuzi–Biega National Park, we compared their diet and analyzed dietary overlap between them in relation to fruit phenology. Data on fruit consumption were collected mainly from fecal samples, and phenology of preferred ape fruits was estimated by monitoring. Totals of 231 plant foods (116 species) and 137 plant foods (104 species) were recorded for gorillas and chimpanzees, respectively. Among these, 38% of gorilla foods and 64% of chimpanzee foods were eaten by both apes. Fruits accounted for the largest overlap between them (77% for gorillas and 59% for chimpanzees). Gorillas consumed more species of vegetative foods (especially bark) exclusively whereas chimpanzees consumed more species of fruits and animal foods exclusively. Although the number of fruit species available in the montane forest of Kahuzi is much lower than that in lowland forest, the number of fruit species per chimpanzee fecal sample (average 2.7 species) was similar to that for chimpanzees in the lowland habitats. By contrast, the number of fruit species per gorilla fecal sample (average 0.8 species) was much lower than that for gorillas in the lowland habitats. Fruit consumption by both apes tended to increase during the dry season when ripe fruits were more abundant in their habitat. However, the number of fruit species consumed by chimpanzees did not change according to ripe fruit abundance. The species differences in fruit consumption may be attributed to the wide ranging of gorillas and repeated usage of a small range by chimpanzees and/or to avoidance of inter-specific contact by chimpanzees. The different staple foods (leaves and bark for gorillas and fig fruits for chimpanzees) characterize the dietary divergence between them in the montane forest of Kahuzi, where fruit is usually scarce. Gorillas rarely fed on insects, but chimpanzees occasionally fed on bees with honey, which possibly compensate for fruit scarcity. A comparison of dietary overlap between gorillas and chimpanzees across habitats suggests that sympatry may not influence dietary overlap in fruit consumed but may stimulate behavioral divergence to reduce feeding competition between them.     

Fruit Choice by Woolly Monkeys in Tinigua National Park,Colombia

I combined morphological fruit data, phenological, and demographic information for 128 plant species, with nutritional information for 78 species, to assess feeding preferences of woolly monkeys (Lagothrix lagothricha lugens) in Tinigua National Park, Colombia. I used multiple and simple regression analyses to predict fruit feeding time and one index of fruit preference from 20 independent variables presumed to affect fruit choice. The results of the independent regressions indicated a trend to spend more time consuming good tasting fruits from abundant plants that produce large quantities of fruits and produce large crops. Among these variables, abundant fruit production and the astringency index are the most important variables explaining feeding times when other variables are statistically controlled in a multiple regression analysis. Furthermore, the preference index showed that woolly monkeys prefer fruits from large trees that produce in periods of fruit scarcity, with high pulp/seed ratios, low lipid contents, and clumped spatial distributions. The results suggest that woolly monkeys are primarily generalized, opportunistic frugivores that avoid some fruits with secondary compounds. Their feeding behavior is determined by many factors, but the predictive power of each one is always low (<15%). No multiple regression model explained >69% of the variation in feeding time. I suggest that a detailed quantification of secondary compounds might increase the predictive power of the models to explain fruit choice by frugivorous primates.     

Advantages of granivory in seasonal environments: feeding ecology of an arboreal seed predator in Amazonian forests

Specialized seed predators are uncommon in arboreal vertebrate assemblages, and the hypothesis that consuming seeds of immature fruits – which may be available for relatively long periods compared to mature fruit – could reduce seasonal food scarcity experienced by generalist frugivores remains largely untested. To test this hypothesis, we examined the diet and feeding ecology of bald‐faced saki monkeys Pithecia irrorata in a largely intact forest mosaic of southeastern Peru based on systematic monitoring of five habituated groups over a three‐year period and compared the relative availability of ripe and unripe fruits in their diet. Plant phenology data from individual tree crowns showed that, compared to ripe fruits, immature fruits were available in more tree species, in greater quantities, and for longer periods. Despite pronounced community‐wide seasonal changes in fruit production at our study area, feeding patterns of bald‐faced saki remained largely invariant: fruits comprised approximately 95% of the species’ monthly diet, with seeds alone accounting for 75%, with no major monthly dietary shifts. The flexible exploitation by this species of a consistently available food supply for which it faces little competition likely reduces foraging effort and consumption of less desirable foods, even during prolonged periods of overall fruit scarcity. The relative rarity of immature fruit specialists in tropical forests may reflect the fact that processing the hard pericarps and neutralizing the toxicities of immature seeds present substantial evolutionary hurdles that few arboreal vertebrate species have overcome.     

Dietary Differences Among Sympatric Callitrichinae in Northern Bolivia: Callimico goeldii,Saguinus fuscicollis and S. labiatus

Studies of sympatric species can provide important data to define how dietary and habitat requirements differ among them. I collected dietary data during a first yearlong comparative study of wild groups of Callimico goeldii, Saguinus labiatus and S. fuscicollis. Dietary overlap was highest between Saguinus fuscicollis and Saguinus labiatus throughout the year, and lowest between Saguinus labiatus and Callimico goeldii. All three species had high dietary overlap in February and March when a few abundant fruit species dominated their diets. Although all three species rely heavily on many of the same fruits and arthropods, there are several important distinctions among their diets. Surprisingly, Callimico goeldii consume large quantities of fungus throughout the year: 29% of annual feeding records. Mycophagy is more frequent in the dry season when fruits are scarce. In contrast, Saguinus labiatus rarely eat fungus during the period of fruit scarcity, and instead rely on nectar, a resource never exploited by Callimico goeldii. Saguinus fuscicollis also rely on nectar during periods of low fruit availability and increase their intake of arthropods and exudates. During April, a period of fruit scarcity, exudates comprise >50% of the feeding records of Saguinus fuscicollis. The use of different food resources during fruit scarcity, and differences in the heights at which each species feeds and forages appear to define a distinct ecological niche for each of them and allow them to maintain long-term associations throughout the year. Furthermore, I hypothesize that the limited distribution of Callimico goeldii may result from their restriction to forests that have high disturbance rates, where microhabitats appropriate for fungal growth are abundant, but which also contain abundant fruit and insects.     

The Influence of Temporal Changes in Fruit Availability on Diet Composition and Seed Handling in Blue Monkeys (Cercopithecus mitis doggetti)1

We investigated the relation between temporally varying resources, diet composition, and seed-handling behaviors in a group of blue monkeys (Cercopithecus mitis doggetti) in a tropical montane forest of Rwanda. Changes in diet composition were related to concurrent phenological studies of fruit-producing trees, and density and abundance of tree resources within the monkey's home range. Fruit composed nearly 50 percent of the diet. Over 50 percent of the fruits eaten had juicy fleshy pulp. Observations of seed handling behavior provided insights into the role of these animals as potential seed dispersal agents. The monkeys moved the seeds of 29 species out of parent canopies by defecating seeds intact and by potentially carrying seeds in cheek pouches and dropping them later. Seeds of 18 species were found intact in fecal piles. Our study showed community-level phenology patterns did not indicate a decrease in fruit availability during the study period, but an analysis of the preferred fruits consumed by the monkeys showed distinct periods of low fruit availability. The study period included two dry seasons; only one of these produced a period of fruit scarcity for the animals. The animals employed different strategies during times of preferred fruit scarcity. They increased consumption of leaves and other fleshy fruits, and diet diversity increased, or became mainly seed predators and diet diversity decreased. The variable responses of these monkeys to changes in food availability highlights their dietary plasticity and imposes significant variations in their role as potential seed dispersers.     

Fallback Foods of Red Leaf Monkeys (Presbytis rubicunda) in Danum Valley, Borneo

Animals in Southeast Asia must cope with long periods of fruit scarcity of unpredictable duration between irregular mast fruiting events. Long-term data are necessary to examine the effect of mast fruiting on diet, and particularly on the selection of fallback foods during periods of fruit scarcity. No such data is available for colobine monkeys, which may consume substantial amounts of fruits and seeds when available. We studied the diet of red leaf monkeys (Presbytis rubicunda, Colobinae) in Danum Valley, Sabah, northern Borneo, using 25 mo of behavioral observation, phenology and vegetation surveys, and chemical analysis to compare leaves eaten with nonfood leaves. The monkeys spent 46% of their feeding time on young leaves, 38% on seeds, 12% on whole fruits, 2.0% on flowers, 1.0% on bark, and 1.2% on pith. They spent more time feeding on seeds and whole fruit when fruit availability was high and fed on young leaves of Spatholobus macropterus (liana, Leguminosae) as fallback foods. This species was by far the most important food, constituting 27.9% of the total feeding time, and the feeding time on this species negatively correlated with fruit availability. Consumed leaves contained more protein than nonconsumed leaves, and variation in time spent feeding on different leaves was explained by their abundance. These results suggest that red leaf monkeys show essentially the same response to the supra-annual increase in fruit availability as sympatric monogastric primates, increasing their seed and whole-fruit consumption. However, they depended more on young leaves, in particular Spatholobus macropterus, as fallback foods during fruit-scarce periods than did gibbons or orangutans. Their selection of fallback food appeared to be due to both nutrition and abundance.     

The response of the frugivorous lion-tailed macaque (Macaca silenus) to a period of fruit scarcity

Tropical rainforests show seasonal fluctuations in the abundance of fruits resulting in periods of resource scarcity for frugivores. We examined the response of an obligate frugivore, the lion‐tailed macaque (LTM) (Macaca silenus), to a period of fruit scarcity in a rainforest in the Western Ghats, India. We estimated the abundance and distribution of fruit resources from food tree densities obtained from 348 point centered quadrats, and fruit availability from phenological monitoring of 195 trees of 15 reported major food species. Macronutrient content was estimated for fruits of 15 major food species. We estimated time spent feeding on different food items from 1,853 individual scans spanning 120 hr of observation of one habituated study group. There was a distinct period of fruit scarcity during the drier months of February to mid‐March (Period 1) compared with late March and April (Period 2), separated by summer showers. Fruits available in Period 1 had lower soluble carbohydrate and lipid content and overall caloric value compared with Period 2. During the lean period, the LTM fed more on fruits of Drypetes wightii, which had the highest carbohydrate content, than on nectar of Palaquium ellipticum or Ficus spp., which had low carbohydrate content. During this period, the resource availability in a location significantly influenced the occurrence of feeding there. In Period 2, the group fed most on the seeds of Cullenia exarillata, the most abundant tree in the home‐range and with the highest content of soluble carbohydrates. During this period, the abundance of food trees in fruit in a location did not seem to influence the occurrence of feeding. Low abundance, stochastic fruiting and, low quality might make Ficus spp. a poor fallback option for the LTM. Am. J. Primatol. 73:1250–1260, 2011. © 2011 Wiley Periodicals, Inc.     

Masting in a temperate tree: Evidence for environmental prediction?

Many plant species produce large fruit crops in some years and then produce few or no fruits in others. Synchronous, inter‐annual variation in plant reproduction is known as ‘masting’ and its adaptive significance has yet to be fully resolved. For 8 consecutive years, I quantified every fruit produced by 22 females of a New Zealand tree species (Dysoxylum spectabile), which has an unusual habit of taking a full calendar year to mature fruits after flowering. Fruit production varied strongly among years and was tightly synchronized among trees. Annual variability in fruit production declined with total reproductive output, indicating trees with lower fecundity exhibited a stronger tendency to mast. Although unrelated to temperature, annual fruit production was positively related to precipitation during annual periods of fruit development, and negatively related to fruit production in the previous year. Seedlings had higher rates of survivorship in a wet, high‐seed year than in a dry, low‐seed year, suggesting that seedlings might be drought sensitive. Therefore, D. spectabile produced large fruit crops during periods of high rainfall prior to fruit maturation, which may enhance survivorship of drought‐intolerant seeds. Results were inconsistent with several hypotheses that are widely believed to be the most likely explanations for masting. Instead, results were consistent with the environmental prediction hypothesis, suggesting that this hypothesis may be more important than previously appreciated.     

The fruits of selectivity: how birds forage on Goupia glabra fruits of different ripeness

Although many studies have been published on avian fruit selection, few have addressed the effects of fruit scarcity on the patterns of fruit choice. Here, we compared the consumption of seven bird species for six simultaneously present maturation stages of Goupia glabra fruits. Ripe G. glabra fruits contain more lipids, carbohydrates and energy, and fewer phenols, than unripe fruits. All bird species selected from among ripening stages and removed a higher proportion of ripe fruits than of intermediate or unripe fruits. Importantly, however, fruit choice was flexible in all species. Whether birds preferred or avoided fruits of intermediate ripeness depended on the overall fruit supply. When ripe fruits were scarce, birds showed a higher acceptance of fruits of intermediate ripeness, but still rejected the least ripe fruit stages. In a foraging bout, most birds fed on fruits of the same ripeness. By doing so, birds maximised instantaneous energy gain per time, because search time was longer for riper fruits while energy intake was lower for less ripe fruits. The results suggest that birds select fruits based on fine-scale differences in profitability, but accept less profitable fruits during low fruit abundance. If environmental factors such as overall fruit availability influence avian fruit choice, we suggest that the potential for directional selective pressures on fruit compounds is restricted.     

Preferences of cedar waxwings in the laboratory for fruit species,colour and size: A comparison with field observations

Fruit preferences of cedar waxwings, Bombycilla cedrorum, in the laboratory were compared with preferences in the field to identify fruit characteristics that influence choice by avian dispersers. Waxwings ate 12 of 13 species of fruits offered individually in laboratory tests, but in the field ate only eight of the same 13 species. When given a choice of 10 fruit species offered simultaneously, waxwings showed a strong preference for fruits that were small or red. Preferences for fruit species in the laboratory were not correlated with morphological or nutritional characteristics of the fruits. When offered food that differed only in colour, waxwings initially preferred red over blue, yellow and green. In later tests, preference for red fruit was less marked. In tests for size preference, waxwings preferred small pieces of fruit (6 mm) over medium pieces (9 mm), and medium pieces over those that were large (12 mm). Sizes of fruits preferred in the field and in the laboratory tests were approximately 6·0–7·5 mm. Thus, waxwings can discern differences in food items and they have definite preferences. The lack of complete agreement between preferences for fruits in the field and in the laboratory suggests that factors important in the field but controlled in the laboratory (e.g. abundance, location) override preferences for certain fruits. However, some fruit characteristics, particularly size, were consistently influential in the laboratory and in the field.     

Fruit removal of Coprosma quadrifida (Rubiaceae) by birds in south-eastern Australia

Abstract Removal of the fleshy fruits of Coprosma quadrifida by birds was examined in relation to fruit crop size, neighbouring fruit crop size and time in the season in montane wet sclerophyll forest in south-eastern Australia. On average, 84% of individual fruit crops were removed. Frugivores removed fruit in proportion to its abundance, indicating that they continually assess availability rather than respond to a critical abundance. The size of neighbouring fruit crops did not influence fruit removal rates, suggesting that there is little competition among neighbours for dispersers. The ripe fruits of C. quadrifida were eaten by 14 bird species, comprising 50% of the bird species trapped. Silvereyes (Zosterops lateralis) were the most abundant consumers of fruits. With the exception of the parrot, Platycercus elegans, all birds defecated seeds intact and had no influence on seed viability.     

Spatial and seasonal variation in niche partitioning between blue manakin (Chiroxiphia caudata) and greenish schiffornis (Schiffornis virescens) in southeastern Brazil

The diet of Chiroxiphia caudata and Schiffornis virescens were studied over three years in four forest sites in the southeastern Atlantic Forest of Brazil. Theory suggests that coexistence of similar species is possible by resource-use differences. To test this, patterns of niche overlap and niche breadth in diet were measured in different seasons and forest sites, based on fruit morphological traits and plant life forms. The results suggest seasonal and spatial variation in resource partitioning. During periods of high fruit availability, there was no evidence of fruit trait selection. In contrast, during periods of fruit scarcity in primary forests, there was evidence of niche partitioning, where C. caudata foraged for larger fruits in canopy and S. virescens searched for understory plants. The niche overlap was higher in secondary forest during periods of scarcity, when both species used small understory fruits.

Foi estudada a dieta de Chiroxiphia caudata e Schiffornis virescens durante três anos, em quatro tipos de fisionomias florestais no sudeste da Mata Atlântica, Brasil. A teoria sugere que a coexistência de espécies semelhantes é possível pelas diferenças no uso dos recursos. Assim, nós mensuramos a amplitude e sobreposição de nicho considerando a morfologia dos frutos consumidos e a forma de vida das espécies de plantas utilizadas em quatro tipos florestais e em diferentes estações. Os resultados sugerem mudanças sazonais e espaciais na partilha de recursos. Durante períodos de alta disponibilidade de frutos, não há evidências de partilha de recursos. Já em períodos de escassez de frutos nas florestas primárias, C. caudata consumiu frutos grandes presentes no dossel enquanto S. virescens utilizou os frutos pequenos do sub-bosque. Na floresta secundária, o período de escassez foi marcado por sobreposição intensa de nicho onde ambas as espécies consumiram frutos pequenos de sub-bosque.     

Crop size,plant aggregation,and microhabitat type affect fruit removal by birds from individual melastome plants in the Upper Amazon

We studied the efficiency (proportion of the crop removed) and quantitative effectiveness (number of fruits removed) of dispersal of Miconia fosteri and M. serrulata (Melastomataceae) seeds by birds in lowland tropical wet forest of Ecuador. Specifically, we examined variation in fruit removal in order to reveal the spatial scale at which crop size influences seed dispersal outcome of individual plants, and to evaluate how the effect of crop size on plant dispersal success may be affected by conspecific fruit abundance and by the spatial distribution of frugivore abundance. We established two 9-ha plots in undisturbed terra-firme understory, where six manakin species (Pipridae) disperse most seeds of these two plant species. Mean levels of fruit removal were low for both species, with high variability among plants. In general, plants with larger crop sizes experienced greater efficiency and effectiveness of fruit removal than plants with smaller crops. Fruit removal, however, was also influenced by microhabitat, such as local topography and local neighborhood. Fruit-rich and disperser-rich patches overlapped spatially for M. fosteri but not M. serrulata, nonetheless fruit removal of M. serrulata was still much greater in fruit-rich patches. Fruit removal from individual plants did not decrease in patches with many fruiting conspecifics and, in fact, removal effectiveness was enhanced for M. fosteri with small crop sizes when such plants were in patches with more conspecifics. These results suggest that benefits of attracting dispersers to a patch balanced or outweighed the costs of competition for dispersers. Spatial pattern of fruit removal, a measure of plant fitness, depended on a complex interaction among plant traits, spatial patterns of plant distribution, and disperser behavior.     

Dietary Response of Chimpanzees and Cercopithecines to Seasonal Variation in Fruit Abundance. I. Antifeedants

In order to understand dietary differentiation among frugivorous primates with simple stomachs, we present the first comparison of plant diets between chimpanzees and cercopithecine monkeys that controls for food abundance. Our aim was to test the hypothesis that monkeys have a more diverse diet as a result of their dietary tolerance for chemical antifeedants. Our study species are chimpanzees, blue monkeys, redtail monkeys, and gray-cheeked mangabeys living in overlapping ranges in Kibale National Park, Uganda. We indexed food abundance by the percentage of trees having ripe fruit within the range of each group; it varied widely during the year. Chimpanzees spent almost 3 times as much of their feeding time eating ripe fruits as the monkeys did and confined their diets almost exclusively to ripe fruits when they were abundant. Monkeys maintained a diverse diet at all times. When ripe fruit was scarce chimpanzee and monkey diets diverged. Chimpanzees relied on piths as their main fallback food, whereas monkeys turned to unripe fruits and seeds. For each primate group we calculated the total weighted mean intake of 5 antifeedants; condensed tannins (CT), total tannins assayed by radial diffusion (RD), monoterpenoids (MT), triterpenoids (TT), and neutral-detergent fiber (NDF). Monkeys had absolutely higher intakes of CT, RD, MT, and TT than those of chimpanzees, and their intake of NDF did not differ from that of chimpanzees, appearing relatively high given their lower body weights. However contrary to expectation, dietary divergence during fruit scarcity was not associated with any change in absolute or relative intake of antifeedants. For example, fruit scarcity did not affect the relative intake of antifeedants by cercopithecines compared to chimpanzees. Our results establish chimpanzees as ripe-fruit specialists, whereas cercopithecines are generalists with a higher intake of antifeedants. The low representation of ripe fruits in the diets of cercopithecines has not been explained. An important next step is to test the hypothesis that the difference between Kibale chimpanzees and cercopithecines represents a more general difference between apes and monkeys.     

Compensatory response of two dune annuals to simulated browsing and fruit predation

Abstract. Greenhouse and field studies were conducted to determine the effects of artificial fruit removal and clipping (simulated deer browsing) on components of reproduction of two indeterminate dune annuals: Cakile edentula var. lacustris (Bigel.) Hook and Corispermum hyssopifolium L. The results showed that both species were able to partially or completely compensate or even overcompensate for low to moderate levels of clipping or fruit removal only if the damage occurred in the early stages of fruit development. The indeterminate habit of growth of both species facilitated their compensatory ability. The removal of fruits and clipping of plants which mimick browsing by white tailed deer along Lake Huron resulted in a significant increase in their period of flowering and fruiting, especially in the early stages of fruit development. The plants compensated for fruit loss by increasing the number of fruits per plant. A delay in fruit removal or clipping to later stages of fruit maturation of both species drastically reduced reproduction. Similar experiments in the field revealed that, although the critical periods during which the plants were most susceptible were the same, the plants had lower compensatory ability. Fruit damage and clipping generally did not affect the seed mass of both species.     

The effect of accessibility on rates of fruit removal from tropical shrubs: An experimental study

Summary In a series of field experiments using Costa Rican rain forest plants, we examined the effect of accessibility on fruit removal rates. We compared the effects of fruit placement in terminal and axillary infructescences on diurnal and nocturnal removal rates, visitation rates, and incidence of fruit damage. We used three different species of berries (Phytolacca rivinoides, Psychotria brachiata, and Psychotria pitteri) and worked in three different habitats (fallow fields, treefall gaps, and forest understory) and in two different seasons (July–September, a season of fruit abundance and December–January, a season of fruit scarcity.)We found that in oldfields especially, diurnal removal rates by birds were significantly greater from axillary than from terminal infructescences. Nocturnal removal from axillary infructescences-presumably by rodents-is also occassionally significant. From these data, from observations on climbing ability and fruit use in captive rodents, and from reports in the literature, we suggest that rodents are significant sources of fruit and seed loss in tropical shrubs. We hypothesize that placement of the infructescence on the plant affects fruit removal by both seed-dispersing birds and by the less agile, often seed-destroying rodents. The balance between the two rates is an important component of a plant's dispersal success.Diurnal fruit removal rates were higher during the season of fruit scarcity than during the season of fruit abundance and higher in old fields than in forest gaps or understory. Fruit damage rates-probably due to orthopterans-were slightly greater in gaps and understory than in old fields.     

Vertical Stratification and Caloric Content of the Standing Fruit Crop in a Tropical Lowland Forest1

Fruit abundance in tropical forests thus far has been studied in relation to consumer populations. Area–based surveys that focus on the quantity and quality of the standing fruit crop of an entire plant community, however, are lacking. This paper presents the results of a four–month study on the seasonality and vertical distribution of the standing fruit crop within a tropical forest during the dry season in southern Venezuela. Fruit numbers ranged from 262,000 to 424,000 fruits/ha. The standing crop was between 39 and 92 kg/ha, yielding 67,000 to 126,000 kj. Fruits were not evenly distributed in different forest layers. They were most abundant in the canopy above 16 m and scarce between 4 and 12 m above the ground. Despite a scarcity of fruits in the mid–strata, the caloric value of the total fruit supply peaked within a narrow layer of subcanopy at 12 to 16 m above ground due to a high amount of energy per fruit. Palms fruited mostly in this layer, their fruits contributing 59 percent of the energy supplied by all fruits. Above 12 m, we found a broader range of fruit sizes than in the forest below 12m. Small–sized fruits were distributed in the understory and in the canopy but were nearly absent from die mid–story. Corresponding to the abundance of small–sized fruits in the canopy, small frugivorous species, such as members of the Thraupidae, limited their foraging to the canopy strata whereas larger avian frugivores foraged mainly in die mid–story.     

Ecological correlates to social structure in two lemur species in Madagascar

In this study, I tested two hypotheses regarding the relationship of ecological variables (size, density, and distribution of patches) and infant developmental patterns to lemur social structure using two prosimian primates in Ranomafana, Madagascar: the rufous lemur (Eulemur fulvus rufus) and the red-bellied lemur (Eulemur rubriventer). Three predictions regarding the general effects of patch size and subgroup size on lemur feeding rates were supported: (1) Rufous lemurs used large patches; red-bellied lemurs used smaller patches; (2) larger subgroups of rufous lemurs used larger patches; and (3) rufous lemur feeding rates decreased significantly with increases in subgroup size and patch size, whereas size and patch size had no significant effect on red-bellied lemur feeding rates. However, food item size (fruit) had a more significant effect on rufous and red-bellied lemur feeding rates than either patch size or subgroup size. When similar-sized fruits were compared, rufous lemur feeding rates on small fruit were most affected by patch size, yet feeding rates on medium-sized fruit were most affected by subgroup size. Neither lemur species used patches in consistent ways seasonally. During periods of food abundance, rufous lemurs used many small, common, and clumped patches. In food scarcity periods, they used fewer, larger, rarer, and less clumped patches; groups migrated when food became most scarce. Red-bellied lemurs also used patches in variable ways, but these patterns were not linked with food availability. Finally, infant development patterns differed between lemur species; red-bellied lemur males cared for offspring and infants reached developmental landmarks faster than rufous lemur infants. Therefore, red-bellied lemur group size may be constrained by the need for additional infant care by other group members. In contrast, rufous lemur group size may be constrained by patch availability during the most critical period of food scarcity. © 1996 Wiley-Liss, Inc.