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

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

Host-mediated effects on the reproductive phenological asynchrony of a generalist mistletoe in China

一种泛性桑寄生植物繁殖物候异步性的寄主介导效应 寄主介导效应被认为会导致半寄生性的桑寄生植物的繁殖物候异步性，并由此为与桑寄生植物互惠共生的传粉者和种子散布者提供更长时间的食物资源供应，但目前关于此方面的研究还缺乏相关的实证数据。本研究以广泛分布于中国西南西双版纳地区的一种泛性桑寄生科植物五蕊寄生(Dendrophthoe pentandra)为材料，每周监测其开花、结果物候，检测了其开花和结果物候是否呈季节性格局，量化了五蕊寄生繁殖物候的异步程度，并检测了影响该植物始花期早晚的因素。最后，本研究还检验了五蕊寄生繁殖物候的异步性随寄主种类数量变化的效应。研究结果表明：(i)在连续两年的物候观测中五蕊寄 生的花期和果期都呈单峰分布格局；(ii)始花期显著受到植物大小和光照强度的影响，即冠幅越大和受光程度越高的植物个体有更早的始花期和更长的花期和果期；(iii)不同的寄主种类对五蕊寄生的繁殖物候有显著的影响，但与假设相反的是，随着寄主种类数量的增加，五蕊寄生繁殖物候的异步性没有显著提高。这项研究表明，在解释泛性桑寄生植物的繁殖物候异步性及寄主种类的数量对其影响的生态学意义还需进行更深入的探究。     

Bird Assemblage and Visitation Pattern at Fruiting Elmerrillia tsiampaca (Magnoliaceae) Trees in Papua New Guinea

Most tropical trees produce fleshy fruits that attract frugivores that disperse their seeds. Early demography and distribution for these tree species depend on the effects of frugivores and their behavior. Anthropogenic changes that affect frugivore communities could ultimately result in changes in tree distribution and population demography. We studied the frugivore assemblage at 38 fruiting Elmerrillia tsiampaca, a rain forest canopy tree species in Papua New Guinea. Elmerrillia tsiampaca is an important resource for frugivorous birds at our study site because it produces abundant lipid-rich fruits at a time of low fruit availability. We classified avian frugivores into functional disperser groups and quantified visitation rates and behavior at trees during 56 canopy and 35 ground observation periods. We tested predictions derived from other studies of plant–frugivore interactions with this little-studied frugivore assemblage in an undisturbed rain forest. Elmerrillia tsiampaca fruits were consumed by 26 bird species, but most seeds were removed by eight species. The most important visitors (Columbidae, Paradisaeidae and Rhyticeros plicatus) were of a larger size than predicted based on diaspore size. Columbidae efficiently exploited the structurally protected fruit, which was inconsistent with other studies in New Guinea where structurally protected fruits were predominantly consumed by Paradisaeidae. Birds vulnerable to predation foraged for short time periods, consistent with the hypothesis that predator avoidance enhances seed dispersal. We identified seven functional disperser groups, indicating there is little redundancy in disperser groups among the regular and frequent visitors to this tropical rain forest tree species.     

Habitat Characteristics of the Endangered Hoolock Gibbons of Bangladesh: The Role of Plant Species Richness

Hoolock gibbons ( Hoolock hoolock ) are endangered small apes occurring in Bangladesh, India, Myanmar, and south China. All known populations have undergone declines primarily due to habitat destruction or alteration. We examined the influence of area of natural forests, area of plantations, area of agricultural land and edible plant species richness on the hoolock gibbon numbers from 22 sites in Bangladesh using multiple regression analysis. Area of natural forests and plantations were not linked to hoolock gibbon numbers, whereas area of agricultural land and edible plant species richness were both significantly linked. Higher edible plant species richness was directly related to hoolock gibbon numbers, while the area of agricultural land was inversely related. This is the first quantitative study illustrating the importance of these two variables with hoolock gibbon numbers. We also analyzed the existing habitat characteristics in two of the largest hoolock gibbon populations in Bangladesh: Lawachara (42 individuals) and Kaptai (84 individuals). Trees in Lawachara were generally smaller compared to those in Kaptai. Artocarpus , Tectona , and Dipterocarpus were the three most important genera in both sites, although Artocarpus was more important in Lawachara. Each site also had an abundant variety of tree species utilized by hoolock gibbons for food. We discuss the current context of primate conservation in Bangladesh and suggest that the current conservation priority for gibbons in the country should be protection and improvement of habitat.     

Nutrients in fruits as determinants of resource tracking by birds

Fruit pulp is an important source of nutrients for many bird species. Fruit‐eating birds use a variety of strategies to cope with changes in the availability of fruits, exhibiting a remarkable ability to track resources. We assessed the role of nutrient availability in the fruiting environment as a factor driving resource tracking by fruit‐eating birds. Fruit consumption by the four most common frugivorous species in a 6‐ha plot in the Southern Yungas montane forest of Argentina was assessed. We determined the content of selected nutrients (soluble carbohydrates, proteins, phenols, ascorbic acid and essential minerals) in 22 fruiting plant species eaten by birds, and measured fruit–frugivore interactions and the availability of nutrients and dry fruit pulp mass over 2 years. There was strong temporal covariation in the availability of the selected nutrients in fruits across the study period. Similarly, the availability of nutrients in the fruiting environment covaried with pulp mass. Fruit consumption by the four commonest bird species and the abundance of most species were positively associated with nutrient availability and dry pulp mass. Nutrient availability was a good predictor of temporal fruit tracking by three of the four commonest frugivores. Despite large differences in particular nutrient concentrations in fruits, overall nutrient (and pulp) quantity in the fruiting environment played a greater role in fruit tracking than did the nutritional quality of individual fruits. While overall nutrient availability (i.e. across fruit) and total pulp mass were important determinants of fruit tracking, we suggest that plant species‐specific differences in fruit nutrient concentration may be important in short‐term foraging decisions involved in fruit choice and nutritional balance of birds.     

Opposed latitudinal patterns of network‐derived and dietary specialization in avian plant–frugivore interaction systems

Latitudinal patterns of biodiversity have been studied for centuries, but it is only during the last decades that species interaction networks have been used to examine the proposed latitudinal gradient of biotic specialization. These studies have given idiosyncratic results, which may either be because of genuine biological differences between systems, different concepts and scales used to quantify biotic specialization or because the methodological approaches used to compare interaction networks were inappropriate. Here we carefully examine the latitudinal specialization gradient using a global dataset of avian plant–frugivore assemblages and interaction networks. In particular, we test whether network‐derived specialization patterns differ from patterns based on assemblage‐level information on avian dietary preferences on specific food types. We found that network‐derived measures of specialization (complementary specialization H₂′ and < d’>, modularity Q) increased with latitude, i.e. frugivorous birds divide the niche of fruiting plants most finely at high latitudes where they also formed more modular interaction networks than at tropical latitudes. However, the strength and significance of the relationship between specialization metrics and latitude was influenced by the methodological approach. On the other hand, assemblage‐level information on avian specialization on fruit diet (i.e. the proportion of obligate frugivorous bird species feeding primarily on fruit) revealed an opposed latitudinal pattern as more bird species were specialized on fruit diet in tropical than in temperate assemblages. This difference in the latitudinal specialization gradient reflects that obligate frugivores require a high diversity of fruit plants, as observed in tropical systems, and fulfil more generalized roles in plant–frugivore networks than bird species feeding on different food types. Future research should focus on revealing the underlying ecological, historical and evolutionary mechanisms shaping these patterns. Our results highlight the necessity of comparing different scales of biotic specialization for a better understanding of geographical patterns of specialization in resource–consumer interactions.     

Fruit preferences by fishes in a Neotropical floodplain

The diets of frugivorous animals result from the interaction between feeding preference and ecological factors such as availability of alternative resources and interactions with other frugivores. A better understanding of frugivore diets will enable predictions about the vulnerability of plant populations to anthropogenic or natural environmental changes. In addition, frugivores with greater variation in diet (generalists) are potentially more resilient to habitat changes than specialists. Here, we combined data on diets of frugivorous fishes with visual censuses on fruit availability to evaluate if fishes are selective or opportunistic in their use of fruits in floodplain habitats. We found a high diversity of fruit species (74 species) frequently consumed by four common species of frugivorous fishes during the flooding season. The identity of preferred fruits, rather than their availability, had a stronger effect on the patterns of fruit consumed by fishes in this system. Fruit preferences by fish may influence the long-term persistence and the maintenance of plant diversity in floodplain habitats. Possible disturbances of fish populations with selective consumption and the plant populations on which they depend could increase the vulnerability of both populations involved. Thus, selectivity may affect ecological and evolutionary processes associated with frugivory by fish.     

Latitudinal cline of flowering and fruiting phenology in Fagaceae in Asia

The seasonal timing of flowering and fruiting is crucial for the reproductive success of plants and for resource availability to animals. Although plants synchronize their reproductive timing to coincide with appropriate seasons by responding to environmental cues, seasonal variations in temperature and precipitation vary minimally in very wet tropical environments. To explore the latitudinal cline in the reproductive phenology of the Fagaceae in Asia, we analyzed phenology data for a total of 94, 121, and 219 species from Thailand, Malesia, and China, respectively, in the three genera of Fagaceae, Quercus, Castanopsis, and Lithocarpus. We found that Quercus and Castanopsis showed flowering peaks in April in China. In Thailand, the peak shifted to an earlier month, and the peak disappeared in Malesia. The flowering period lengthened with decreasing latitude in the animal-pollinated genera Castanopsis and Lithocarpus. However, this was not the case for the wind-pollinated genus Quercus. The fruiting period lengthened with decreasing latitude in all three genera. We examined the relationship between reproductive phenology and climatic factors. The combination of monthly temperature and precipitation best explained the monthly change in the proportion of flowering and fruiting species in China in all three genera. However, climatic factors had almost no impact on the predictive ability of the model in Malesia. Our results on phenological shifts in the family Fagaceae, from the temperate climates and seasonal tropics to the humid tropics, provide valuable information for predicting phenological changes in future climate change.     

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

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

海南长臂猿破碎化生境中食物资源可利用性分析

食物种类及其可获得性是动物生存的基本条件,也是生境质量评价的重要指标。2013年1月至2014年7月,对海南长臂猿 (Nomascus hainanus) C群生境内食源植物种类、数量、食物可获得性及主要取食植物的径级结构进行调查分析,共记录到64种食源植物 (标记胸径 ≥ 5 cm的乔木1 484株)。海南长臂猿以高大粗壮的食源植物为主要采食对象。这些乔木型食源植物结果率超过50.0%的有15种,占C群全部乔木型采食植物种数的28.8%;结果率最高的是海南单籽暗罗 (Monoon laui, 76.7%),最低的是鹅掌柴 (Heptapleurum heptaphyllum, 9.6%)。18种长臂猿主食食源乔木中,白肉榕 (Ficus vasculosa)、斜叶榕 (Ficus tinctoria) 等15种食源植物的植株呈增长型结构。南酸枣 (Choerospondias axillaria)、二色波罗蜜 (Artocarpus styracifolius) 为稳定型结构,而桃榄 (Pouteria annamensis) 为衰退型结构。提示海南长臂猿食物供应植物仍处于年轻状态,但这些食源植物并非每年结果,每年都能结果的仅37种 (71.2%),采食树种的结果率高低不受海拔影响,而与树高和胸径显著相关。海南长臂猿在果实稀少的旱季会采食一定量的树叶 (15.6%),这种不稳定的食物供应将影响海南长臂猿的种群复壮。     

Seasonal variation of diet and time budget of Eastern hoolock gibbons (Hoolock leuconedys) living in a northern montane forest

Most gibbons dwell in the tropical forests of Southeastern Asia, but eastern hoolock gibbons (Hoolock leuconedys) survive in high montane forest ranging from 1,600 to 2,700 m a.s.l. in Gaoligongshan (>24°30′N), Yunnan, China. To assess the behavioral adaptations of hoolock gibbons to the montane forest, we related temperature and food availability within the habitat to the seasonal behavioral patterns of a family group and a solitary female between August 2010 and September 2011 in Nankang, Gaoligongshan National Nature Reserve. The maximum temperature was 29.2 °C and the minimum temperature was ?0.3 °C during the period. The monthly mean temperature was <10 °C between December and February, making Nankang the coldest gibbon habitat reported so far. Nonfig fruit and fig availability declined to nearly zero in cold months. The family group increased resting and decreased travel and social behaviors when the monthly mean temperature was low. Compared with other gibbon populations, the hoolock gibbons spent proportionally less time feeding on figs and other fruit than other gibbon populations except Nomascus concolor and Symphalangus syndactylus. Only 36 species of plants provided nonfig fruit or figs, which is less than the number of fruit species consumed by any other gibbon population observed during a similar period of time (about 1 year). Hoolock gibbons shifted their diet to leaves and increased feeding time when fruit was not available. We conclude that diet flexibility and an energy-conserving strategy during the cold season when fruit is scarce have enabled the hoolock gibbons to survive in a northern montane forest.     

Influence of Forest Seasonality on Gibbon Food Choice in the Rain Forests of Barito Ulu,Central Kalimantan

We describe the diet of two hybrid gibbon groups (Hylobates mulleri x H. agilis) in relation to forest seasonality. We collected data over 12 mo in lowland dipterocarp forest in the Barito Ulu research area, Central Kalimantan, Indonesia. Although non-fig fruit was the main dietary item (52–64% of diet), gibbon diet was most strongly influenced by the availability of flowers. During periods when flowers were most abundant and the gibbons increased consumption of them, they also ate figs or young leaves more often. We suggest that although flowers are nutritionally rich sources of food, providing relatively high levels of protein compared to fruit, they are unlikely to satiate gibbon hunger and they seek dietary bulk from figs or young leaves, because they are easily obtained. Rainfall also influenced food choice, and non-fig fruit availability had a weak influence on fruit selection for one group. The group concentrated feeding on the fruit of a few species when fruit was most abundant and ate a greater diversity of species when fruit was scarce. Gibbon diet appeared not to be influenced by changes in availability of figs, young leaves and diversity of fruiting species.     

Resource-related variables drive individual variation in flowering phenology and mediate population-level flowering responses to climate in an asynchronously reproducing palm

Many tropical plant species show wide intra-population variation in reproductive timing, resulting in the protracted presence of flowering and fruiting individuals. Various eco-evolutionary drivers have been proposed as ultimate causes for asynchronous phenology, yet little is known about the proximate factors that control reproductive onset among individuals or that influence the proportion of trees producing new inflorescences within a population. We employed a nine-year phenological record from 178 individuals of the hyperdominant, asynchronously flowering canopy palm, Oenocarpus bataua (Arecaceae)¸ to assess whether resource-related variables influence individual- and population-level flowering phenology. Among individuals, access to sunlight increased rates of inflorescence production, while the presence of resource sinks related to current investment in reproduction—developing infructescences—reduced the probability of producing new inflorescences. At the population level, climate anomalies induced by El Niño Southern Oscillation (ENSO) affected the proportion of the population producing inflorescences through time. Moreover, the effects of ENSO anomalies on flowering patterns depended on the prevalence of developing infructescences in the population, with stronger effects in periods of low developing-infructescence frequency. Taken together, these results suggest that resource-related variables can drive phenological differences among individuals and mediate population-level responses to larger-scale variables, such as climate anomalies. Consequently, a greater focus on the role of resource levels as endogenous cues for reproduction might help explain the frequent aseasonal phenological patterns observed among tropical plants, particularly those showing high intra-population asynchrony.     

Context‐dependency of a complex fruit–frugivore mutualism: temporal variation in crop size and neighborhood effects

The quantity of fruit consumed by dispersers is highly variable among individuals within plant populations. The outcome of such selection operated by frugivores has been examined mostly with respect to changing spatial contexts. The influence of varying temporal contexts on frugivore choice, and their possible demographic and evolutionary consequences is poorly understood. We examined if temporal variation in fruit availability across a hierarchy of nested temporal levels (interannual, intraseasonal, 120 h, 24 h) altered frugivore choice for a complex seed dispersal system in dry tropical forests of southern India. The interactions between Phyllanthus emblica and its primary disperser (ruminants) was mediated by another frugivore (a primate), which made large quantities of fruit available on the ground to ruminants. The direction and strength of crop size and neighborhood effects on this interaction varied with changing temporal contexts. Fruit availability was higher in the first of the two study years, and at the start of the season in both years. Fruit persistence on trees, determined by primate foraging, was influenced by crop size and conspecific neighborhood densities only in the high fruit availability year. Fruit removal by ruminants was influenced by crop size in both years and neighborhood densities only in the high availability year. In both years, these effects were stronger at the start of the season. Intraseasonal reduction in fruit availability diminished inequalities in fruit removal by ruminants and the influence of crop size and fruiting neighborhoods. All trees were not equally attractive to frugivores in a P. emblica population at all points of time. Temporal asymmetry in frugivore‐mediated selection could reduce potential for co‐evolution between frugivores and plants by diluting selective pressures. Inter‐dependencies formed between disparate animal consumers can add additional levels of complexity to plant–frugivore mutualistic networks and have potential reproductive consequences for specific individuals within populations.     

Determinants of fruit removal in <Emphasis Type="Italic">Geonoma pauciflora</Emphasis>, an understory palm of neotropical forests

Competition for seed dispersers is supposedly a selective force that drives the evolution of plant reproductive traits. In the understory of tropical forests, such competition should be especially severe among bird-dispersed plant species because (i) the production of copious fruit crops is limited by low light availability; (ii) there usually is a high density of fleshy fruited plants, and (iii) understory frugivorous birds are not abundant. In this paper, we took advantage of a high-density population of Geonoma pauciflora, a bird-dispersed palm species growing in the understory of the Brazilian Atlantic forest, to investigate the influence of plant traits and its immediate neighborhood on fruit removal. Intrinsic (crop and fruit sizes) and extrinsic traits (related to light availability) affected the fruit removal of G. pauciflora. Crop size had a greater influence than fruit size on the absolute number of fruits removed, whereas none of the investigated traits influenced decisively fruit removal efficiency (i.e., the proportion of an individual’s crop removed). The influence of light availability was mostly indirect, through its positive influence on fruit production. A significant positive spatial autocorrelation in removal efficiency occurred among neighboring plants within a 7-m radius, which is indicative of facilitation among neighboring individuals. The consequence of such positive spatial autocorrelation in removal efficiency for clonal plants such as G. pauciflora is that by attracting a frugivore a given ramet may promote the removal of fruits of other ramets, thus enhancing the reproductive output of the genet as a whole.     

Hyperdominance in fruit production in the Brazilian Atlantic rain forest: the functional role of plants in sustaining frugivores

The availability of fruits is critical for tropical forests, where the majority of plant species rely upon animal vectors for seed dispersal. However, we do not know how fruit production is temporally distributed over species and families. Two plant families are particularly important in floristic inventories of Atlantic rain forests: Arecaceae, a few species of which are highly abundant; and Myrtaceae, which is abundant and displays outstanding species diversity. In this context, we asked whether hyperdominance occurs in fruit production in the Atlantic rain forest, and whether it occurs in the abundant species of Arecaceae and Myrtaceae. We investigated whether the temporal fruit production patterns differ between Myrtaceae, Arecaceae, and the plant community as a whole. We also applied a functional dispersion index to assess the temporal fruit diversity over a 2‐yr period, with regard to morphological and phenological traits. We found that the phenomenon of hyperdominance occurs in fruit production: five species accounted for more than half of the pulp biomass. Arecaceae fruit biomass peaked at the end of wet season, overlapping with the community peak; whereas Myrtaceae species fruited throughout the year and were an important resource during periods of food scarcity. Myrtaceae filled more of the fruit morphospace over time because their fruits exhibit a large range of morphologies and phenological strategies. Our results demonstrated the importance of combining phenology and fruit morphology in the evaluation of resource availability, which revealed periods of high fruit diversity that could support a range of frugivore sizes and maintain overall ecosystem functionality.     

Gibbons (Nomascus gabriellae) provide key seed dispersal for the Pacific walnut (Dracontomelon dao), in Asia's lowland tropical forest

Understanding the mutualisms between frugivores and plants is essential for developing successful forest management and conservation strategies, especially in tropical rainforests where the majority of plants are dispersed by animals. Gibbons are among the most effective seed dispersers in South East Asia's tropical forests, but are also one of the highly threatened arboreal mammals in the region. Here we studied the seed dispersal of the Pacific walnut (Dracontomelon dao), a canopy tree which produces fruit that are common in the diet of the endangered southern yellow-cheeked crested gibbon (Nomascus gabriellae). We found that gibbons were the most effective disperser for this species; they consumed approximately 45% of the fruit crop, which was four times more than that consumed by macaques – the only other legitimate disperser. Gibbons tracked the temporal (but not spatial) abundance of ripe fruits, indicating this fruit was a preferred species for the gibbon. Both gibbons and macaques dispersed the majority (>90%) of the seeds at least 20 m away from parent crowns, with mean dispersal distances by gibbons measuring 179.3 ± 98.0 m (range: 4–425 m). Seeds defecated by gibbons germinated quicker and at greater rates than seeds spat by macaques, or in undispersed fruits. Gibbon-dispersed seeds were also more likely to be removed by unknown seed predators or unknown secondary dispersers. Overall, gibbons play a key role in the regeneration of the Pacific walnut. Our findings have significant implications both for the management of the Pacific walnut tree dominating tropical rainforest as well as the reintroduction program of the Southern yellow-cheeked crested gibbon.     

High Conservation Value of Forest Fragments for Plant and Frugivore Communities in a Fragmented Forest Landscape in South Africa

Fragmentation is a major threat factor for plant–frugivore communities in tropical and subtropical forests. Resulting changes in the distribution of traits within these communities, e.g., a loss in large‐bodied frugivores, may lead to strong changes in plant–frugivore interactions in fragmented forests. Yet, we still lack a thorough understanding of the interplay between forest fragmentation, the trait‐composition of communities and resulting plant–frugivore interactions on a community‐scale. In a fragmented South African landscape comprising different forest categories—i.e., continuous natural forest, forest fragments surrounded by natural grassland, and forest fragments surrounded by sugarcane—we investigated the relationship between communities of fruiting plants and their frugivore visitors in response to forest fragmentation, as well as the interactive effects of forest fragmentation and fruit size of the plants on the number of frugivore visitors and their body size. Neither the fruit size of plant nor the body mass of frugivore communities differed between natural forest sites and forest fragments. Moreover, in‐depth analyses of frugivore assemblages visiting plant species revealed no effect of forest category on the number of frugivore visits or their mean body mass. The number of visits and body mass of frugivores were merely determined by the crop and fruit size of the focal plant species. Overall, our results suggest that frugivory of plant species with differently sized fruits was not reduced in forest fragments. Thus, fragments with high fruit availability may be key elements maintaining the functional connectivity of a heterogeneous forest landscape.     

Frugivore Behavioural Details Matter for Seed Dispersal: A Multi-Species Model for Cantabrian Thrushes and Trees

Animal movement and behaviour is fundamental for ecosystem functioning. The process of seed dispersal by frugivorous animals is a showcase for this paradigm since their behaviour shapes the spatial patterns of the earliest stage of plant regeneration. However, we still lack a general understanding of how intrinsic (frugivore and plant species traits) and extrinsic (landscape features) factors interact to determine how seeds of a given species are more likely to be deposited in some places more than in others. We develop a multi-species mechanistic model of seed dispersal based on frugivore behavioural responses to landscape heterogeneity. The model was fitted to data from three-years of spatially-explicit field observations on the behaviour of six frugivorous thrushes and the fruiting patterns of three fleshy-fruited trees in a secondary forest of the Cantabrian range (N Spain). With such model we explore how seed rain patterns arise from the interaction between animal behaviour and landscape heterogeneity. We show that different species of thrushes respond differently to landscape heterogeneity even though they belong to the same genus, and that provide complementary seed dispersal functions. Simulated seed rain patterns are only realistic when at least some landscape heterogeneity (forest cover and fruit abundance) is taken into account. The common and simple approach of re-sampling movement data to quantify seed dispersal produces biases in both the distance and the habitat at which seeds arrive. Movement behaviour not only affects dispersal distance and seed rain patterns but also can affect frugivore diet composition even if there is no built-in preference for fruiting species. In summary, the fate of seeds produced by a given plant species is strongly affected by both the composition of the frugivore assemblage and the landscape-scale context of the plant location, including the presence of fruits from other plants (from the same or different species).     

Selection of Fruit by Gibbons (Hylobates muelleri × agilis) in the Rain Forests of Central Borneo

Gibbons (Hylobates spp.) are among the main frugivorous primates in Southeast Asia, yet little is known about the criteria by which they select fruit for consumption. We studied two gibbon groups for 14 mo in the lowland dipterocarp forests of Central Borneo to determine their selectivity for different fruit species and traits. Ideal gibbon fruit were yellow, large, with a juicy-soft pulp, thin skin and available in large crops. Gibbons ultimately sought seedless fruit, but when seeds were present they selected fruit with a single, well-protected seed. Given that few fruit exhibited all the desired traits, we also carried out a multiple regression using the selection ratios of the various fruit species and their associated fruit traits to determine which traits ultimately determined gibbon choice. The analysis was stratified to account for differences in fruit availability. Selection was strongest when fruit were abundant in the forest and was based on seed width (<21 mm), color (yellow-orange), and fruit weight (1–5 g). No selection is apparent when food abundance was intermediate, but when fruit were scarce they preferentially ate larger fruit (6–30 g).