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.     

Feeding habits of chimpanzees (Pan troglodytes), red-tail monkeys (Cercopithecus ascanius schmidti) and blue monkeys (Cercopithecus mitis stuhlmanii) on figs in Budongo Forest Reserve, Uganda

Feeding habits of chimpanzees, red‐tail and blue monkeys on figs (Ficus) were studied in compartment N3 of Budongo Forest Reserve, western Uganda, from September 1997 to March 1998. The aim was to examine the spatial and temporal foraging habits of chimpanzees, red‐tail monkeys and blue monkeys on figs in the forest reserve. Both scan and focal sampling methods were used to assess the foraging habits of the primates. It was found that the primates fed on emerging leaves of Ficus mucuso Ficalho, F. varifolia Warb. and F. exasperata Vahl. They also preferred ripe fruits to emerging, young and unripe fruits. The primates spent 78% of the morning eating fruits and leaves and inhabited fig trees with fruits for about 4 h. Fig trees with ripe fruits attracted larger numbers of primate groups. It is concluded that information on the feeding habits of chimpanzees and monkeys is required in order to have a clear understanding of the social behaviour and pattern of movement of the primates and to assist in predicting the likely impacts of poor forest management, forest degradation and loss of food resources on their populations.     

Diet composition of chimpanzees inhabiting the montane forest of Kahuzi,Democratic Republic of Congo

The diet of chimpanzees was investigated by direct observations, feeding remains, and fecal analysis from January 1994 to December 2000 in the montane forest of Kahuzi-Biega National Park. A total of 171 food items were identified, among which 156 items were plant materials belonging to 114 species from 57 taxonomic families. Chimpanzees consumed 66 species of fruits (62 species of pulps and four species of seeds). Results of fecal analysis showed that fig fruits were the most frequently eaten. Their seeds occurred in 92% of a total of 7212 chimpanzee fecal samples. The chimpanzees changed their diet according to seasonal and annual variations in both abundance and diversity of fruit species. However, they are very selective frugivores. Only a few pulp-fruit species are regularly identified in their fecal samples. During the rainy season, when ripe fruit was scarce, chimpanzees relied heavily on piths and leaves. They swallowed leaves of two species of Commelinaceae without chewing, probably for medical purposes. Animal foods were eaten infrequently. The montane forest of Kahuzi, where chimpanzees range up to 2600 m above sea level, may be the highest altitudinal limit ever recorded for their distribution. Compared to other chimpanzee habitats, Kahuzi has a low diversity of fruit species and the availability of a few pulp-fruit species may be critical to the survival of Kahuzi chimpanzees.     

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.     

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

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

Foraging Patterns of Sympatric Hornbills during the Nonbreeding Season in Arunachal Pradesh, Northeast India

The diet and food resource partitioning of three sympatric hornbills (Great Hornbill Buceros bicornis, Wreathed Hornbill Aceros undulatus, and Oriental Pied Hornbill Anthracoceros albirostris) during the nonbreeding season were studied relative to fruit availability in a foothill semievergreen forest of Arunachal Pradesh, northeast India. Hornbills fed on fruits of 49 plant species that comprised over 95 percent of their diet. Hornbill species partitioned food resources by varying the relative contribution of figs and non‐fig fruits in the diet. Similarity in non‐fig fruit diet was low. Ten species contributed to over 90 percent of the non‐fig diet. The availability of non‐fig fruits was much lower in the nonbreeding season than in the breeding season; however, despite lower fruit availability during the nonbreeding season, hornbills had a wide diet breadth and indirect evidence suggests that Wreathed Hornbills foraged widely in this season for fruit. Great Hornbills relied more on figs that were available year‐round. The diverse diet breadth that included rare and patchy fruit resources underscores the importance of conserving large forest tracts for hornbills.     

Chimpanzee seed dispersal quantity in a tropical montane forest of Rwanda

We describe chimpanzee seed dispersal in the tropical montane forest of Nyungwe National Park (NNP), Rwanda, for a total of three years from January 1998 through May 2000 and May 2006 through March 2007. Relatively few studies have examined chimpanzee seed dispersal in montane communities where there are generally fewer fruiting tree species than in lowland forests. Such studies may reveal new insights into chimpanzee seed dispersal behaviors and the role that they play in forest regeneration processes. Chimpanzees are large‐bodied, highly frugivorous, and tend to deposit the seeds of both large‐ and small‐seeded fruits they consume in a viable state. We found that chimpanzees dispersed a total of 37 fruiting species (20 families) in their feces, 35% of which were large‐seeded trees (≥0.5 cm). A single large‐seeded tree, Syzygium guineense, was the only species to be dispersed in both wadges and feces. Based on phenological patterns of the top five large‐seeded tree species found in chimpanzee feces, our results indicate that chimpanzees do not choose fruits based on their availability. There was, however, a positive relationship between the presence of Ekebergia capensis seeds in chimpanzee feces and S. guineense seeds in chimpanzee wadges and their respective fruit availabilities. Our data reveal that proportionately fewer chimpanzee fecal samples at NNP contained seeds than that reported in two other communities in the Albertine Rift including one at mid‐elevation and one in montane forest. As in other chimpanzee communities, seeds of Ficus spp. were the most common genus in NNP chimpanzee feces. Our data do not support previous studies that describe Ficus spp. as a fallback food for chimpanzees and highlights an intriguing relationship between chimpanzees and the large‐seeded tree species, S. guineense. Am. J. Primatol. 71:901–911, 2009. © 2009 Wiley‐Liss, Inc.     

Few figs for frugivores: Riparian fig trees in Zimbabwe may not be a dry season keystone resource

Most plants flower and fruit at times of year when probabilities of pollination and seedling establishment are high. Fig trees (Ficus spp.) are often considered as keystone resources for vertebrate frugivores, in part because of year-round fig production. This unusual fruiting phenology results in the maintenance of fig wasp populations, but in seasonal environments this means fruiting occurs during periods when the chances of seedling establishment are low. Under these circumstances, selection is expected to favour any individuals that reduce or eliminate fruiting at these times. Here, we describe a large-scale survey of the extent of dry season fruiting by three riparian Ficus species in Gonarezhou National Park, Zimbabwe. Few trees of two monoecious species, F. sycomorus and F. abutilifolia, had figs, and most crops of F. sycomorus were far smaller than the trees were capable of producing. Large stands of the dioecious F. capreifolia were present, but fig densities were low and no mature female (seed containing) figs were recorded. Even though fig trees may have been the only species bearing fruit, the consequences of the low investment in reproduction by the three Ficus species were clear—there were too few figs for a landscape-scale keystone role.     

Chimpanzee feeding ecology and fallback food use in the montane forest of Nyungwe National Park,Rwanda

Almost all primates experience seasonal fluctuations in the availability of key food sources. However, the degree to which this fluctuation impacts foraging behavior varies considerably. Eastern chimpanzees (Pan troglodytes schweinfurthii) in Nyungwe National Park, Rwanda, live in a montane forest environment characterized by lower primary productivity and resource diversity than low‐elevation forests. Little is known about chimpanzee feeding ecology in montane forests, and research to date predominantly relies on indirect methods such as fecal analyses. This study is the first to use mostly observational data to examine how seasonal food availability impacts the feeding ecology of montane forest chimpanzees. We examine seasonal changes in chimpanzee diet and fallback foods (FBFs) using instantaneous scan samples and fecal analyses, supported by inspection of feeding remains. Chimpanzee fruit abundance peaked during the major dry season, with a consequent change in chimpanzee diet reflecting the abundance and diversity of key fruit species. Terrestrial herbaceous vegetation was consumed throughout the year and is defined as a “filler” FBF. In contrast to studies conducted in lower‐elevation chimpanzee sites, figs (especially Ficus lutea) were preferred resources, flowers were consumed at seasonally high rates and the proportion of non‐fig fruits in the diet were relatively low in the current study. These divergences likely result from the comparatively low environmental diversity and productivity in higher‐elevation environments.     

Behavioral Ecology of Sympatric Chimpanzees and Gorillas in Bwindi Impenetrable National Park,Uganda: Diet

Via a field study of chimpanzees (Pan troglodytes schweinfurthii) and gorillas (Gorilla gorilla beringei) in Bwindi Impenetrable National Park, Uganda, we found that their diets are seasonally similar, but diverge during lean seasons. Bwindi chimpanzees fed heavily on fruits of Ficus sp., which were largely ignored by the gorillas. Bwindi gorilla diet was overall more folivorous than chimpanzee diet, but was markedly more frugivorous than that of gorillas in the nearby Virunga Volcanoes. During 4 mo of the year Bwindi gorilla diet included more food species than that of the chimpanzees. Three factors in particular—seasonal consumption of fibrous foods by gorillas, interspecific differences in preferred fruit species, and meat consumption by chimpanzees—contributed to dietary divergence between the two species. When feeding on fruits, gorillas ate Myrianthus holstii more frequently than chimpanzees did, while chimpanzees included more figs in their annual diet. Chimpanzee diet included meat of duikers and monkeys; gorilla frequently consumed decaying wood.     

Signs from a green desert: a preliminary examination of the archaeobotanical remains from a Byzantine dovecote near Shivta, Israel

This paper presents a preliminary examination of archaeobotanical material from pigeon dung samples obtained from Byzantine period destruction levels of a dovecote near the site of Shivta, Israel. Such pigeon dung was a valuable fertilizer in antiquity and would not have been abandoned without a reason. The plant remains from the dung provide direct evidence of pigeon diet and the local environment during the Byzantine period. Eleven plant taxa, represented by either seeds and/or plant parts (cereal chaff material), including five wild taxa, one legume, four fruit/nut taxa and several unidentified seed fragments were recovered. The most common seeds found were from weeds of the genus Thymelaea sp., and Ficus (fig). The finds indicate that the birds in the dovecote consumed a mixed diet of wild seeds including Thymelaea sp. and Fumaria sp. (fumitory), and small fruits of Vitis (grape), Ficus (fig), Olea (olive) and Phoenix (date). The sample analyzed also included Rumex (dock), Carex (sedge) and Androsace which may not have come from the feed. Apparently the pigeons were free to forage in the desert, the fallowed fields and refuse piles or/and were intentionally fed agricultural by-products including wild plants.     

The significance of fibrous foods for Kibale Forest chimpanzees.

Four categories of plant food dominated the diet of chimpanzees in Kibale Forest, Uganda: non-fig tree fruits, fig tree fruits, herbaceous piths and terrestrial leaves. Fruit abundance varied unpredictably, more among non-figs than figs. Pith intake was correlated negatively with fruit abundance and positively with rainfall, whereas leaf intake was not influenced by fruit abundance. Piths typically have low sugar and protein levels. Compared with fruits and leaves they are consistently high in hemicellulose and cellulose, which are insoluble fibres partly digestible by chimpanzees. Herbaceous piths appear to be a vital resource for African forest apes, offering an alternative energy supply when fruits are scarce.     

Sexual specialization in two tropical dioecious figs

Ficus species (figs) and their species-specific pollinator wasps are involved in an intimate mutualism in which wasps lay eggs in some ovaries of the closed inflorescences (syconia), and mature, inseminated offspring carry pollen from mature syconia to fertilize receptive inflorescences. In monoecious species, each syconium produces seeds and wasps. In functionally dioecious fig species, making up approximately half the figs worldwide, male and female functions are separated; hermaphrodite (functionally male) trees produce wasps and pollen only, while female trees produce seeds only. This sexual separation allows selection to act independently on the reproductive biology of each sex. Examining sexual specialization in a tight mutualism allows us to determine aspects of the mutualism that are flexible and those that are canalized. In this study, we quantified the phenology of two species of dioecious figs, F. exasperata and F. hispida, for 2 years by following the fates of several thousand syconia over time. In studying each of these species in a dry and a wet site in south India, we tested specific predictions of how dioecious figs might optimize sexual function. On female trees of both species, more inflorescences matured during the wet (monsoon) season than in any other season; this fruiting period enabled seeds to be produced during the season most suitable for germination. In F. exasperata, functionally male trees released most wasps from mature syconia in the dry season, during peak production of receptive female syconia, and thus maximized successful pollination. In F. hispida, “male” trees produced more syconia in the dry and monsoon seasons than in the post-monsoon season. In both species, male and female trees abscised more unpollinated, young inflorescences than pollinated inflorescences, but abscission appeared to be more likely due to resource- rather than pollinator- limitation. The phenology of F. exasperata requires that male inflorescences wait in receptive phase for scarce pollinators to arrive. As expected, male inflorescences of this species had a longer receptive phase than female inflorescences. In F. hispida, where pollinators are rarely scarce, duration of receptive phase was the same for both sexes. Duration of developing phase was longer in female syconia of both species than in male syconia, most likely because they need a longer period of investment in a fleshy fruit. Variation in developing phase of female syconia in one species (F. exasperata) was also greater than that in male syconia, and enabled female trees to sample a variety of germination environments in time. The strong sexual differences in both fig species support the hypothesis that selection for sexual specialization has strongly influenced the reproductive biology of these species. Received: 28 May 1997 / Accepted: 2 February 1998     

Seasonality of Leaf and Fig Production in Ficus squamosa,a Fig Tree with Seeds Dispersed by Water

The phenology of plants reflects selection generated by seasonal climatic factors and interactions with other plants and animals, within constraints imposed by their phylogenetic history. Fig trees (Ficus) need to produce figs year-round to support their short-lived fig wasp pollinators, but this requirement is partially de-coupled in dioecious species, where female trees only develop seeds, not pollinator offspring. This allows female trees to concentrate seed production at more favorable times of the year. Ficus squamosa is a riparian species whose dispersal is mainly by water, rather than animals. Seeds can float and travel in long distances. We recorded the leaf and reproductive phenology of 174 individuals for three years in Chiang Mai, Northern Thailand. New leaves were produced throughout the year. Fig production occurred year-round, but with large seasonal variations that correlated with temperature and rainfall. Female and male trees initiated maximal fig crops at different times, with production in female trees confined mainly to the rainy season and male figs concentrating fig production in the preceding months, but also often bearing figs continually. Ficus squamosa concentrates seed production by female plants at times when water levels are high, favouring dispersal by water, and asynchronous flowering within male trees allow fig wasps to cycle there, providing them with potential benefits by maintaining pollinators for times when female figs become available to pollinate.     

西双版纳地区4种榕树果实的资源分配及其种子萌发特性

为了解榕树果实的资源分配和种子萌发特征,对西双版纳地区的垂叶榕(Ficus benjamina)、高山榕(F.altissima)、聚果榕(F.racemosa)和无柄雅榕(F.concinna)的果实营养、繁殖资源分配,以及种子萌发特性进行了研究。结果表明,4种榕果的营养分配存在差异,榕果重的种类果肉亦多,榕果大者果腔较大,且种子数量较多,但雄花和榕小蜂数量不一定多。榕果分配给雌性功能及雄性功能繁殖的小花比例也不相同,聚果榕在雌性功能上的分配显著大于雄性功能,其它3种榕果则分配于雄性功能的资源比雌性功能的多,但程度不同。此外,4种榕树的种子大小及重量存在显著的种间差异,以至于影响到种子的萌发率及根茎生长,果实最大的聚果榕种子最多,萌发率高,但由于种子最小,其根茎生长缓慢且短;果实适中的垂叶榕和高山榕种子较大,不但萌发率较高,其根茎生长也较快;果实和种子都小的无柄雅榕,种子萌发率低,根茎生长慢。生长在水热环境好的榕树比生长在相对贫瘠环境的榕树,由于极少面临水热光照等限制,更容易进化出较小的种子,萌发能力和速度也相对较差。这说明不同生境中榕树进化出了不同的繁殖分配机制及资源利用策略来适应环境变化。     

Phenology of Tree Species in Bolivian Dry Forests

Phenological characteristics of 453 individuals representing 39 tree species were investigated in two dry forests of the Lomerío region, Department of Santa Cruz, Bolivia. The leaf, flower, and fruit production of canopy and sub–canopy forest tree species were recorded monthly over a two–year period. Most canopy species lost their leaves during the dry season, whereas nearly all sub–canopy species retained their leaves. Peak leaf fall for canopy trees coincided with the peak of the dry season in July and August. Flushing of new leaves was complete by November in the early rainy season. Flowering and fruiting were bimodal, with a major peak occurring at the end of the dry season (August–October) and a minor peak during the rainy season (January). Fruit development was sufficiently long in this forest that fruiting peaks actually tended to precede flowering peaks by one month. A scarcity of fruit was observed in May, corresponding to the end of the rainy season. With the exception of figs (Ficus), most species had fairly synchronous fruit production. Most canopy trees had small, wind dispersed seeds or fruits that matured during the latter part of the dry season, whereas many sub–canopy tree species produced larger animal– or gravity–dispersed fruits that matured during the peak of the rainy season. Most species produced fruit annually. Lomerio received less rainfall than other tropical dry forests in which phenological studies have been conducted, but rainfall can be plentiful during the dry season in association with the passage of Antarctic cold fronts. Still, phenological patterns in Bolivian dry forests appear to be similar to those of other Neotropical dry forests.     

A comparison of growth and reproduction, under laboratory conditions, of males and females of a dioecious fig tree

The interaction between Ficus spp. (Moraceae) and their pollinating wasps (Chalcidoidae: Agaonidae) is a highly co-evolved mutualism. Approximately half of all fig species are monoecious and produce a mixture of wasps and seeds within the same fig. In functionally dioecious fig trees male and female functions are separate. Figs on male trees produce wasps and pollen, whereas figs borne on female plants produce only seeds. Dioecious fig phenology provides an excellent opportunity to investigate the effect of sexual specialization on the obligate fig?Cfig wasp interaction and the non-pollinators associated with the system. Here we describe laboratory studies of phenological variation between two sexes in terms of vegetative growth and fig production in a dioecious fig tree Ficus montana. We also describe reproductive output in terms of wasp production in males and seeds in females. Intrasexual asynchrony was observed for the plants, with synchrony between the sexes with year-round production of figs. Male plants grew more rapidly, but leaf phenology was very similar. Crop sizes and development times were the same for males and females. Seasonal effects were strong for leaf phenology and fig initiation, but had a very limited effect on fig composition. The results show that the phenological differences described for other dioecious figs do not apply to all species.     

Influence of fruit availability on macronutrient and energy intake by female chimpanzees

Daily energy intake of adult female mammals is influenced by environmental conditions and physiological requirements, including reproduction. We examined the effects of fruit availability on macronutrient and metabolisable energy (ME) intake by adult female chimpanzees (Pan troglodytes schweinfurthii) of the Kanyawara community in Kibale National Park, Uganda, from January 2014 through June 2015. Drupe fruits were abundant for 4 months, whereas the other 14 months were dominated by fig fruits. The mean daily intake of food (dry matter) and ME did not differ between drupe‐months and fig‐months. However, foraging costs were higher during fig‐months, as indicated by a 20% increase in feeding time. Furthermore, during drupe‐months female chimpanzees ingested more water‐soluble carbohydrates and lipids, and less available protein and neutral detergent fibre. Although ME intake did not differ consistently between drupe‐months and fig‐months, they consumed more on days when ripe fruit dominated the diet than when leaves and pithy stems dominated the diet. Our data suggest that differences in diet quality between drupes and figs can have important effects on frugivore foraging and that they influence net energy gain more by their effects on macronutrient composition or foraging cost than by their direct impact on energy intake.     

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.     

Plasticity and diversity of the phenology of dioecious Ficus species in Taiwan

While Ficus present a series of traits often associated with dioecy, the prevalence of dioecy in Ficus is atypical. In Asian floras, dioecious Ficus species generally outnumber monoecious ones. Further this is also true in relatively northerly locations for Ficus such as the island of Taiwan. Ficus are pollinated by species-specific wasps that use fig flowers as breeding sites. In dioecious fig species, pollinators develop only in the inflorescences of male fig trees. In this study, we investigated the reproductive phenology of four dioecious Ficus species with distinct ecologies in several locations in northern and southern Taiwan. The two first species (Ficus erecta and Ficus septica) were investigated in four locations. Reproductive phenology was quite different among sites, even within a single species. For example, F. erecta presented well-defined crops at the population level in its usual high-elevation habitat but continuous fig production at low elevations, especially in South Taiwan. The two other fig species (Ficus pedunculosa var. mearnsii and Ficus tinctoria subsp. swinhoei), are shrubs growing together along seashores in exposed locations on coral reef remnants. These two species presented quite different traits allowing the survival of pollinating wasp populations. Ficus pedunculosa var. mearnsii produced figs continuously so that fresh receptive figs were always available for the pollinating wasps while F. tinctoria subsp. swinhoei extended the period of receptivity of its figs, so that receptive figs that had been waiting for pollinating wasps were almost always available. In summary, dioecious figs in Taiwan showed remarkable variation in their phenology, within species among locations or among species within location. Nevertheless, despite this variation, the phenology of the trees always allowed survival of pollinating wasp populations. Dioecious figs seem to have adopted a differentiated set of strategies which result in high resilience of pollinator populations. This resilience could help explain the atypical prevalence of dioecy in Ficus.