Seasonal Food Use Strategies of <Emphasis Type="Italic">Semnopithecus vetulus nestor</Emphasis>, at Panadura and Piliyandala,Sri Lanka

Field studies on Semnopithecus vetulus have shown high folivory and the ability to feed heavily on mature leaves, which are constantly available. In research spanning 19 mo, I examined the feeding behavior of 2 free-ranging groups of Semnopithecus vetulus nestor in home gardens and rubber monocultures at Panadura (PT1 group) and Piliyandala (R1 group), Sri Lanka. Overall, results showed that >80% of their diet comprised seasonal plant parts, largely fruits. Despite differences in spatial and temporal food availability in their respective habitats, seasonal plant parts dominated the diets of both groups except briefly (2 mo) for R1 when mature leaf petioles were the main plant food. Both groups increased their use of seasonal foods with heightened seasonal food availability, and increased dietary diversity with declining use of their highest-ranked specific item of diet: fruits of Artocarpus heterophyllus (jakfruit, Moraceae). PT1, which was in a species-rich environment, maintained a high intake of seasonal foods year round by exploiting a large number of species, mainly for fruits. In contrast, R1, in a habitat with significantly lower tree species richness, had a less diverse diet but maintained an equally high intake of seasonal foods, primarily fruits and seeds, by exploiting a few species heavily. My study also highlights the dietary flexibility of a single colobine species in space and time. Such information is useful for conservation planning because rapidly occurring changes are taking place in natural colobine habitats.     

Primate Diet and Biomass in Relation to Vegetation Composition and Fruiting Phenology in a Rain Forest in Gabon

To test the hypothesis that primate populations are limited by food resources, we studied the feeding ecology of three cercopithecines and one colobine in a rain forest in central Gabon. Simultaneously, we monitored the fruiting phenology of trees and estimated the biomass of the monkey community. The Makandé Forest is dominated by Caesalpiniaceae and characterized by a lack of secondary vegetation and of trees species producing fleshy fruits. Fruit production was irregular intra- and interannually. Fruiting peaks of dry fruits (mainly Caesalpiniaceae) and of fleshy fruits occurred at the same period. However, interseasonal and interannual variability was greater in Caesalpiniaceae than in other families. As a result, the Makandé forest is subject to bottlenecks when food is scarce. On an annual basis, seeds (primarily Caesalpiniaceae) dominated the diet of all monkeys. On a seasonal basis, cercopithecines preferentially consumed fleshy fruits as long as they were available, whereas colobines increased consumption of young leaves when seed availability declined. The consumption of mature leaves was low. The monkey community biomass (ca. 204 kg/km²) is one of the lowest in Central Africa. We suggest that both cercopithecine and colobine populations are limited as a result of the combined effect of the dominance of Caesalpiniaceae, which provide dry fruits according to a mast-fruiting pattern and mature leaves of low quality, and the lack of seral successional stages, which provide fleshy fruit on a more regular pattern and leaves of better quality. During the period of food scarcity, cercopithecines should suffer from the low availability of fleshy fruit, which are their favorite food. At the same period, colobines should be limited by the low availability of edible leaves. Similar low primate biomasses are found in forests dominated by Caesalpiniaceae or Lecythidaceae in South America and in Dipterocarpaceae forests in South Asia, which suggests that their biological characteristics, in particular dry fruits and mast fruiting, are unfavorable to monkey populations. Our results confirm that habitat mosaics may support larger populations of primary consumers than homogeneous primary forests can.     

Purple-faced Langurs in Human-Modified Environments Feeding on Cultivated Fruits: A Comment to Dela (2007, 2012)

Recently Dela (International Journal of Primatology, 28 (2007): 607–626; International Journal of Primatology, 33 (2012): 40–72) published her study from the mid-1980s on the diet of purple-faced langurs Trachypithecus (Semnopithecus) vetulus in village gardens and rubber plantations. Unlike studies from the 1970s that reported the species, like other colobines, to be largely folivorous with few ripe fruits eaten, Dela found them to be largely frugivorous. The frequent feeding on ripe fruits challenges the paradigm that colobine digestive adaptations restrict the use of ripe fleshy fruits. No reference was made to any other post-1970s study on the species. Here I provide a concise overview of more than a dozen studies conducted in the 1990s and 2000s that show that 1) other populations live in similar human-modified environments showing feeding adaptations as reported by Dela, 2) these populations rely largely on cultivated crops and feed heavily on fruits, 3) living in these situations introduces them to additional threats. Especially in western Sri Lanka little natural habitat remains and deforestation has led the langurs to exchange the forest jungle for the urban jungle, with power lines, fences, walls, and roofs being used instead of trees. The main fruits that provide a staple for langurs in these areas are jackfruit (Artocarpus heterophyllus), banana (Musa spp.), and mango (Mangifera indica); studies in undisturbed habitats and indeed Dela’s own study suggest the heavy use of human-edible fruits by langurs may not necessarily indicate preferential selection of these food sources. Living in human-modified environments makes the langurs more prone to infection with gastrointestinal parasites, and may lead to death by electrocution or being killed by guard dogs. The large degree of agreement between studies suggest that feeding on ripe fruits from cultivars is not unique to Dela’s two study groups and shows that some langur groups are able to survive for extended periods on uncolobine-like diets when they cannot access their preferred foods.     

Craniodental mechanics and diet in Asian colobines: Morphological evidence of mature seed predation and sclerocarpy

Folivory has been accepted as the general dietary pattern for colobines. However, recent ecological studies have revealed that extensive seed eating is found in some colobine species. The ripeness of foraged seeds is also reported to differ between seed eaters. As seeds are generally stress‐limited and may pose greater mechanical demands, seed‐eating species are predicted to exhibit morphological features adaptive for seed predation. In addition, species that feeds on seeds from unripe fruits with hard pericarp is predicted to exhibit increased leverage for anterior dentition. To test these hypotheses, we compared the craniodental morphology of seed‐eating Asian colobines (Presbytis rubicunda and Trachypithecus phayrei) with those of species that rarely exploit seeds (Presbytis comata, Trachypithecus obscurus, and Semnopithecus vetulus). The results show that the seed‐eating colobines possess a masticatory system with enhanced leverage at postcanine bite points. The sclerocarpic forager P. rubicunda also exhibits markedly greater masticatory leverage at anterior dental bite points, while the mature‐seed‐eating T. phayrei shows no such advantage for canine and incisor use. These observations suggest that P. rubicunda is well adapted to husking the resistant pericarps of unripe fruits, using the anterior dentition and to gain access to the immature seeds, whereas such sclerocarpic feeding behavior may be less important for T. phayrei. Our findings indicate that the distinctive craniodental variations of colobines may be linked to mature and/or immature seed eating and suggest the significance of seed predation for the evolution of colobine monkeys. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

The feeding ecology and activity budget of proboscis monkeys

A group of proboscis monkeys (Nasalis larvatus) consisting of an alpha‐male, six adult females, and several immatures was observed from May 2005–2006. We collected over 1,968 hr of focal data on the adult male and 1,539 hr of focal data on the six females in a forest along the Menanggul River, Sabah, Malaysia. Availability and seasonal changes in plant species consumed by the focal monkeys were determined by vegetation surveys carried out across an area of 2.15 ha along 200–500 m trails in riverine forest. A total of 188 plant species were consumed by the focal monkeys. The activity budget of members of our study group was 76.5% resting, 19.5% feeding, and 3.5% moving. Young leaves (65.9%) and fruits (25.9%) accounted for the majority of feeding time. Over 90% of fruit feeding involved the consumption of unripe fruits and in the majority of case both the fruit flesh and seeds were eaten. Although fruit eating was rare in some months, during other times of the year time fruit feeding exceeded the time devoted to young leaves. We found that monthly fruit availability was positively related to monthly fruit eating and feeding activity, and seasonal fluctuations in dietary diversity were significantly affected by fruit eating. These results suggest that fruit availability and fruit‐eating behaviors are key factors that influence the activity budget of proboscis monkeys. Earlier assumptions that colobine monkeys are obligate folivores do not apply well to proboscis monkeys and certain other colobines. Our findings may help contribute to a better understanding of the dietary adaptations and feeding ecology of Asian colobines. Am. J. Primatol. 71:478–492, 2009. © 2009 Wiley‐Liss, Inc.     

The diet of the capped langur (Presbytis pileata) in a moist deciduous forest in Bangladesh

Capped langurs (Presbytis pileata) in Madhupur National Park in north-central Bangladesh have an annual diet that is comprised largely of mature leaves (42% of 20,460 total feeding records). Among colobine monkeys, only some populations of red colobus (Procolobus badius) have a diet richer in mature leaves. New leaves (11%) and fruit (24%) are the other annually important dietary items. Seasonal breakdown of this diet, however, revealed that during the monsoon months of May through September theP. pileata diet is approximately 50% fruit, including pulpy ripe fruit. This is also the period of maximum fruit availability. Data on diet and food availability indicate that while capped langurs subsist on mature leaves during the dry season (80% of diet from November to March), they select fruit and new leaves and switch to these foods whenever they are available. These data support the hypothesis that the colobine feeding strategy is adapted to cope with seasonal food scarcity.     

Relationships between the fossil colobine Mesopithecus pentelicus and extant cercopithecoids, based on dental metrics

Maxillary dental measurements from six specimens of Mesopithecus pentelicus, 64 cercopithecines, 59 African colobines, and 64 Asian colobines were analyzed by means of a "nested research design" that was specifically designed to explore the affiliation between fossils and extant cercopithecoids at different systematic levels. The results showed that the variation among taxonomic groups was mainly associated with size; however, in addition, interesting shape differences emerged, molars were shown to be important discriminators, Mesopithecus was confirmed as a colobine (as expected) and found to be closer to Asian species than to African ones, and the odd-nosed colobines were found to share more dental similarities with Mesopithecus than other colobines. The last finding is in contrast to previous studies, in which it was proposed that M. pentelicus is morphologically closely related to the African colobus and the gray langur (Semnopithecus).     

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 diet and feeding behavior of the silver leaf monkey (Trachypithecus auratus sondaicus) in Indonesia

I describe the diet and feeding behavior of silver leaf monkeys (Trachypithecus auratus sondaicus) in the Pangandaran Nature Reserve, West Java, Indonesia, and compare a group living in old secondary rain forest with a group living in mixed plantation/secondary forest to determine intraspecific variation in feeding behavior and the importance of the plantation species in the diet of the monkeys. Young leaves and leaf buds made up slightly less than half of their diets, with both groups showing a preference for a few species when seasonally available. Fruits and flowers of a few species were also preferentially selected when available. These included sweet, fleshy fruits, which most other colobines tend to avoid. Young leaf intake was greatest in months when fruit intake was low. Mature leaves were rarely eaten. Both groups spent approximately 20% of feeding time foraging on Moraceae species. Differences in the diet of the two groups were related largely to differences in vegetational composition and the availability and abundance of food items for the species common to both sites. Teak (Tectona grandis) was the top food species of the group living in mixed plantation/secondary forest, with the midribs of young leaves preferentially selected. Young leaves ofT. grandis, available throughout the study, provided a staple food and were eaten when preferred foods were scarce. More favored food items were available to the group living in old secondary forest, though none was a staple food.     

Theoretical and functional complexity of white variegation of unripe fleshy fruits

In many plant species, the bright colors of ripe fruit serve to attract frugivores to enable efficient seed dispersal. Here I show that the fleshy fruit of several dozens of species originating from Asia (southeastern, eastern and central), the Middle East, Africa, America (South, Central and North), Australia, Polynesia and Micronesia, with fruit usually larger than 1 cm, have white or light green spots while they are still unripe. In many of these species, while the spots are conspicuous, the unripe fruit is known to be poisonous, bitter or sour. I propose that this fruit syndrome may signal frugivores that the fruit is still unripe. Similarly to the succulent leaves of window-plants, these spots form windows that enable light to penetrate deeper into the photosynthetic layers in the developing fruit. This seems to be a solution to overcome the limitations of light harvest because of the high volume to surface ratio of developing fleshy fruits. The white or whitish variegation in these unripe fleshy fruits may serve at least five functions: 1) Windows for photosynthesis, 2) camouflage, 3) signaling to frugivores that they are not ripe (possibly sometimes a type of mutualism with frugivores), 4) signaling to frugivores that they are poisonous - aposematism, and 5) mimicking insect eggs to reduce egg laying. All these functions may be partly or fully simultaneous. Because these white spots appear in plants of diverse geographical and taxonomic origin, it is probably an old adaptation, and such a syndrome has appeared and been selected for many times.     

Alternative seed-handling strategies in primates: seed-spitting by long-tailed macaques (Macaca fascicularis)

Summary The seeds in fruits consumed by primates may be chewed and digested, swallowed and defecated intact, or separated from the flesh and spat out. We show by a combination of close field observations and experiments with caged animals, that long-tailed macaques (Macaca fascicularis) have a remarkably low threshold of 3–4 mm for swallowing seeds and also that wild macaques rarely break them. The seeds of 69% of the ripe fruit species eaten are spat out intact or cleaned outside the mouth and dropped. Seed-spitting significantly reduces the swallowed food bulk and may lessen the risk of releasing seed toxins during mastication. However, it requires that even small fruits are processed in the mouth one or a few at a time. We suggest that fruit storage in the cheek pouches of cercopithecine monkeys allows them to spit seeds individually without excessively slowing fruit intake while feeding on patchily distributed fruit. In contrast, Apes and New World monkeys apparently swallow and defecate most ripe seeds in their diet and colobine monkeys break and digest them, detoxifying seed defenses by bacterial fermentation.     

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.     

Diet ofColobus polykomos on Tiwai Island: Selection of food in relation to its seasonal abundance and nutritional quality

A group of Colobus polykomosat Tiwai, Sierra Leone, demonstrated seasonal flexibility in its diet, with seeds, young leaves, and mature leaves each dominating the diet at different times. Comparison of food consumption with phenological data indicates that seeds are eaten whenever available and are preferred to other foods, while young leaves are preferred to mature leaves. Colobus polykomosalso prefer liane to tree leaves, despite the relatively high quality of mature tree foliage at the Tiwai site. Analysis of protein, fiber, and energy values of foods selected and items available, but not eaten, suggests that preference is related to protein and energy maximization. Leguminous plants, especially Papilionaceae and Mimosaceae, are highlighted as important food sources for C. polykomos;seeds and leaves from these families have a high nitrogen content, and the protein content of leguminous seeds often equals or exceeds that found in leaves. It is predicted, therefore, that colobines living in habitats with a high density of legumes will feed heavily on seeds, subject to constraints such as seasonal availability. Where suitable leguminous species are less common, a mixture of fruits, seeds, and young or mature leaves or both is likely to be selected. The results of this and other recent studies of colobines do not support the notion that colobines are specialist folivores.     

Digestive strategies and food choice in mantled howler monkeys Alouatta palliata mexicana: bases of their dietary flexibility

Mantled howler monkeys (Alouatta palliata) occupy a wide variety of tropical habitats and are the most folivorous of New World primates. However, their diet may include fruits, buds, petioles, and flowers, as well as leaves, suggesting they must cope with variations in the nutrient composition of their food. We studied the physiological basis of the dietary flexibility of these monkeys by comparing food choice, digestive performance and patterns of digesta flow in six adults, fed diets of either leaves or a mixture of fruit and leaves. Although monkeys ate similar amounts of the two diets, they ingested more digestible protein when offered the leaf diet, on which they lost body mass, but they ingested much more soluble sugars when offered fruit and leaves on which they gained mass. Digestibilities of dry matter, fat, energy and fibre did not differ between diets, but those of crude protein, soluble sugars and minerals were higher on the fruit–leaf diet. Mean retention times in the gut of solute (Co-EDTA) and particulate markers (Cr-mordanted cell walls) did not differ between diets, but on both diets the monkeys retained the particulate marker (mean retention time ca 55 h) for longer than they did the solute marker (MRT ca 50 h). A lack of selective retention of solutes and small particles in the gastro-intestinal tract of howler monkeys probably restricts them to mixed diets but their digestive strategy is sufficiently flexible to allow them to feed on a diet of leaves when fruit is unavailable.     

Feeding Ecology of Guerezas in the Kakamega Forest,Kenya: The Importance of Moraceae Fruit in Their Diet

Eastern black-and-white colobus (Colobus guereza), or guerezas, have long been considered to be one of the most folivorous primates. I conducted a study of the feeding ecology of two guereza groups (T and O) over an annual cycle in the Kakamega Forest of western Kenya. I found that the annual diets of both groups comprised mostly of leaves (T: 48%, O: 57%) though fruit (T: 44%, O: 33%) also accounted for a substantial portion of the diet. In the six months when fruit was most abundant, fruit consumption constituted an average of 58% of T-group's monthly diet and 42% of O-group's monthly diet. In contrast to most previous studies of colobines, in which seeds were the primary fruit item consumed, almost all of the fruit eaten by guerezas at Kakamega consisted of whole fruits. At least 72% of the whole fruits consumed by T-and O-groups were whole fruits from trees in the Moraceae family, which dominates the tree family biomass at Kakamega. Unlike at sites where guerezas consumed fruit primarily when young leaves were scarce, at Kakamega guerezas ate fruit in accordance with its availability and irrespective of the availability of young leaves. My findings demonstrate that guerezas are more dietarily flexible than was previously known, which may help to explain why the species can survive in such a wide variety of forested habitats across equatorial Africa.     

Nutritional and Morphological Traits of Invasive and Exotic Fleshy‐fruits in South Africa

While fleshy‐fruited invasive alien plants are recognized as some of the worst invaders on a global scale, until recently, little consideration has been given to the frugivores that feed on these fruits and, more specifically, the fruit traits, which may influence this. We investigated a series of morphological and nutritive fruit traits for ca 30 species of fleshy‐fruited invasive alien and exotic species in South Africa. Invasive alien fruit traits were compared with comparable traits of a similar sample size of indigenous fleshy fruits, which occur in the same area. Finally, the similarity of traits for the same invasive alien species was compared with those fruits in Australia. Invasive alien fleshy fruits were similar in morphology, but greater in some nutritive aspects when compared with indigenous fruits. Furthermore, they were very similar in all aspects to their counterparts in Australia. Most seeds of invasive fleshy fruits were small and light, which may explain some of their invasive success, as benefits associated with small seededness may promote invasive potential. Nutritionally, most invasive alien fleshy‐fruits were hexose‐dominant, containing low lipid and nitrogen content. While frugivore preference trends remain to be formally investigated, this study provides insights into fruit traits, which may tentatively outline why invasive fruits are universally fed on and thus successfully spread.     

Mapping homology between human and black and white colobine monkey chromosomes by fluorescent in situ hybridization

We used in situ hybridization of chromosome specific DNA probes (“chromosome painting”) of all human chromosomes to establish homologies between the human and the white and black colobus (Colobus guereza 2n = 44). The 24 human paints gave 31 signals on the autosomes (haploid male chromosome set). Robertsonian translocations between chromosomes homologus to human 14 and 15, 21 and 22, form colobine chromosomes 6 and 16, respectively. Reciprocal translocations were found between human chromosomes 1 and 10, 1 and 17, as well as 3 and 19. The alternating hybridization signals between human 3 and 19 on Colobus chromosome 12 show that in this case a reciprocal translocation was followed by a pericentric inversion. The hybridization data show that in spite of the same diploid number and similar Fundamental Numbers, the black and white colobine monkey differs from Presbytis cristata, an Asian colobine, by 6 reciprocal translocations. Comparisons with the hybridization patterns in other primates show that some Asian colobines have a more derived karyotype with respect to African colobines, macaques, great apes, and humans. Chromosome painting also clearly shows that similarities in diploid number and chromosome morphology both between colobines and gibbons are due to convergence. Am. J. Primatol. 42:289–298, 1997. © 1997 Wiley-Liss, Inc.     

Exclusive frugivory and seed dispersal of Rhamnus alaternus in the bird breeding season

We studied avian frugivory and seed dispersal in a dioecious shrub, Rhamnus alaternus, focusing on the quantitative and qualitative components of effectiveness. The study took place at three locations in the northeast of the Iberian Peninsula, and examined bird behaviour, intensity of feeding, and the consequences for seedling emergence. The coincidence between the bird breeding season and fruit ripening of R. alaternus in the absence of other ripe fruit, generates a monospecific interaction. The extant frugivorous species were mainly legitimate seed dispersers and their abundance was low. Sylvia melanocephala and S. undata were the most important at one site whereas S. atricapilla, Erithacus rubecula and Turdus merula predominated at the other two sites. Fruit handling took place directly on the branches. Bird species used microhabitats differently as first post-feeding perch, which usually was a short distance away. The low density of frugivorous birds in all localities, among others factors, resulted in satiation of the disperser community and many mature fruits unconsumed. Both adults and juveniles feed upon the plants and their foraging patterns are similar. Adults of S. melanocephala were observed to feed fruit to nestlings and consequently a second phase of dispersal potentially arises from the transport of fecal sacs. Pulp removal and passage through the digestive tract increased the probability of seedling emergence. This plant-dispersal interaction has important consequences, both positive and negative for the plant. Positively, the fruiting of R. alaternus at a time when other ripe fruits are not available avoids interspecific competition for seed dispersers. In addition, a low density of seed rain may reduce intraspecific competition. Negatively, the low density and small size of the breeding frugivorous bird community limit fruit handling and removal away from the parent plants, while the territorial behaviour of birds at that time of the year reduces the potential distances of seed dispersal.     

脊椎动物传播植物肉质果中的次生物质及其生态作用

在种子植物-动物的互惠关系中,植物果实成熟后需要吸引种子传播者取食果实,传播其种子至适宜萌发的生境,同时又要防御种子捕食者过度消耗种子。果实内的次生物质(如:配糖生物碱、大黄素、辣椒素)在此过程中起到重要的调控作用。依赖脊椎动物传播的肉质果中往往含有与植物茎、叶内相同的次生物质,其种类繁多,主要分为含氮化合物、酚类化合物和萜类化合物。未成熟果实内富含次生物质(如:单宁、大黄素),主要保护未成熟种子不被潜在的捕食者和食果动物取食,这些次生物质的含量通常随果实成熟而降低;其它次生物质(如:脱辅基类胡萝卜素)的含量随果实成熟而增多,可能起到吸引食果动物的作用。在对脊椎动物捕食的抵御中,果实内不同类型的次生物质促使成熟果实对所有脊椎动物都有毒性(专毒性)或者仅对种子捕食者有毒性(泛毒性)。肉质果内的次生物质对植物-食果动物相互关系的调控作用,还可以通过调节动物取食频次和数量、抑制和促进种子萌发、改变种子在肠道的滞留时间、吸引传播者等生态作用而实现。某种次生物质往往集多种生态作用于一身。目前对肉质果内次生物质与脊椎动物相互关系的探讨还不够深入。未来研究需要综合考虑植物次生物质与果实生理生化、形态学等特征对食果者的综合调控机理;次生物质在种子传播后的调控作用对植物种群或群落结构和分布格局的影响;从动植物协同进化角度探讨植物次生物质的产生、防御和吸引策略与脊椎动物对果实的选择和消费之间的关系等。开展脊椎动物传播肉质果实中次生物质的研究,对完善种子传播机制、植物繁殖和更新格局,丰富动植物相互作用、协同进化理论具有重要的意义。     

Asian colobine social structure: Ecological and evolutionary constraints

Both ecological and phylogenetic factors and their interaction influence primate social structure. Food resources, in particular, affect social structure parameters such as sex/age composition, group size, home range size, and individual density. As a group, the Asian colobines may be partially buffered from the impact of food constraints on social structure, given their specialized digestive physiology. Asian colobines are able to subsist on mature foliage, a relatively abundant and non-patchily distributed resource, when preferred foods are not available. In food limited populations, increases in group size or density should cause increases in home range size. We did not find this relationship for Asian colobines, however, and thus it would appear that the majority of Asian colobine populations are below carrying capacity. Food may be less of a limitation to populations than other factors, such as social stress. The formation of bands (associations between relatively stable groups) has been documented for bothNasalis andRhinopithecus, and based on recent data for severalPresbytis andTrachypithecus species, we predict that band formation is the norm for the Asian colobines, includingPygathrix. Group home ranges forPresbytis andTrachypithecus species typically overlap and both intergroup tolerance and intergroup aggression are observed. This suggests that groups form differentiated relationships, tolerating some groups but not others. Such differentiated relationships are the foundation of band formation.