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。     

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.     

Dependence of broad-scale geographical variation in fleshy-fruited plant species richness on disperser bird species richness

Aim We analysed the interdependence of avian frugivore‐ and fruited plant‐species richness at the scale of major river basins across Europe, taking into account several environmental factors along different spatial gradients. Location Continental Europe and the British Isles. Methods We focused on wintering birds and autumn/winter fruiting plants, and used major river basins as geographical units and Structural Equation Modelling as the principal analytical tool. Results The statistical influence of disperser species richness on fleshy‐fruited plant species richness is roughly double that of the reverse. Broad‐scale variation in frugivore richness is more dependent on environmental factors than on fruited plant richness. However, the influence of disperser richness on plant richness is four times higher than the influence of environmental factors. Environmental influences on both birds and plants are greater than purely spatial influences. Main conclusions Our results are interpreted as indicating that biotic dispersal of fruits strongly affects broad‐scale geographical trends of fleshy‐fruited plant species richness, whereas richness of fruited plants moderately affects frugivore richness.     

Evolution of angiosperm seed disperser mutualisms: the timing of origins and their consequences for coevolutionary interactions between angiosperms and frugivores

The origins of interactions between angiosperms and fruit‐eating seed dispersers have attracted much attention following a seminal paper on this topic by Tiffney (1984). This review synthesizes evidence pertaining to key events during the evolution of angiosperm–frugivore interactions and suggests some implications of this evidence for interpretations of angiosperm–frugivore coevolution. The most important conclusions are: (i) the diversification of angiosperm seed size and fleshy fruits commenced around 80 million years ago (Mya). The diversity of seed sizes, fruit sizes and fruit types peaked in the Eocene around 55 to 50 Mya. During this first phase of the interaction, angiosperms and animals evolving frugivory expanded into niche space not previously utilized by these groups, as frugivores and previously not existing fruit traits appeared. From the Eocene until the present, angiosperm–frugivore interactions have occurred within a broad frame of existing niche space, as defined by fruit traits and frugivory, motivating a separation of the angiosperm–frugivore interactions into two phases, before and after the peak in the early Eocene. (ii) The extinct multituberculates were probably the most important frugivores during the early radiation phase of angiosperm seeds and fleshy fruits. Primates and rodents are likely to have been important in the latter part of this first phase. (iii) Flying frugivores, birds and bats, evolved during the second phase, mainly during the Oligocene and Miocene, thus exploiting an existing diversity of fleshy fruits. (iv) A drastic climate shift around the Eocene–Oligocene boundary (around 34 Mya) resulted in more semi‐open woodland vegetation, creating patchily occurring food resources for frugivores. This promoted evolution of a ‘flying frugivore niche’ exploited by birds and bats. In particular, passerines became a dominant frugivore group worldwide. (v) Fleshy fruits evolved at numerous occasions in many angiosperm families, and many of the originations of fleshy fruits occurred well after the peak in the early Eocene. (vi) During periods associated with environmental change altering coevolutionary networks and opening of niche space, reciprocal coevolution may result in strong directional selection formative for both fruit and frugivore evolution. Further evidence is needed to test this hypothesis. Based on the abundance of plant lineages with various forms of fleshy fruits, and the diversity of frugivores, it is suggested that periods of rapid coevolution in angiosperms and frugivores occurred numerous times during the 80 million years of angiosperm–frugivore evolution.     

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.     

Frugivores and cheap fruits make fruiting fruitful

Animal seed dispersal provides an important ecosystem service by strongly benefiting plant communities. There are several theoretical studies on the ecology of plant–animal seed–disperser interactions, but few studies have explored the evolution of this mutualism. Moreover, these studies ignore plant life history and frugivore foraging behaviour. Thus, it remains an open question what the conditions for the diversification of fruit traits are, in spite of the multitude of empirical studies on fruit trait diversity. Here, we study the evolution of fruit traits using a spatially explicit individual‐based model, which considers the costs associated with adaptations inducing dispersal by frugivory, as well as frugivore foraging behaviour and abundance. Our model predicts that these costs are the main determinants of the evolution of fruit traits and that when the costs are not very high, the evolution of larger fruit traits (e.g. fleshy/colourful fruits) is controlled by the choosiness and response thresholds of the frugivores as well as their numerical abundance.     

Reliability of macaques as seed dispersers

Seed dispersal is an ecological process crucial for forest regeneration and recruitment. To date, most studies on frugivore seed dispersal have used the seed dispersal effectiveness framework and have documented seed-handling mechanisms, dispersal distances and the effect of seed handling on germination. In contrast, there has been no exploration of “disperser reliability” which is essential to determine if a frugivore is an effective disperser only in particular regions/years/seasons or across a range of spatio-temporal scales. In this paper, we propose a practical framework to assess the spatial reliability of frugivores as seed dispersers. We suggest that a frugivore genus would be a reliable disperser of certain plant families/genera if: (a) fruits of these plant families/genera are represented in the diets of most of the species of that frugivore, (b) these are consumed by the frugivore genus across different kinds of habitats, and (c) these fruits feature among the yearly staples and preferred fruits in the diets of the frugivore genus. Using this framework, we reviewed frugivory by the genus Macaca across Asia to assess its spatial reliability as seed dispersers. We found that the macaques dispersed the seeds of 11 plant families and five plant genera including at least 82 species across habitats. Differences in fruit consumption/preference between different groups of macaques were driven by variation in plant community composition across habitats. We posit that it is essential to maintain viable populations of macaques across their range and keep human interventions at a minimum to ensure that they continue to reliably disperse the seeds of a broad range of plant species in the Anthropocene. We further suggest that this framework be used for assessing the spatial reliability of other taxonomic groups as seed dispersers.     

Fruit traits of vertebrate-dispersed alien plants: smaller seeds and more pulp sugar than indigenous species

Vertebrates play a major role in dispersing seeds of fleshy-fruited alien plants. However, we know little of how the traits of alien fleshy fruits compare with indigenous fleshy fruits, and how these differences might contribute to invasion success. In this study, we characterised up to 38 fruit morphology, pulp nutrient and phenology traits of an assemblage of 34 vertebrate-dispersed alien species in south-eastern Queensland, Australia. Most alien fruits were small (81% < 15 mm in mean width), and had watery fruit pulps that were high in sugars and low in nitrogen and lipids. When compared to indigenous species, alien fruits had significantly smaller seeds. Further, alien fruit pulps contained more sugar and more variable (and probably greater) nitrogen per pulp wet weight, and species tended to have longer fruiting seasons than indigenous species. Our analyses suggest that fruit traits could be important in determining invasiveness and could be used to improve pre- and post-border weed risk assessment.     

Fruit–frugivore interactions in two southern hemisphere forests: allometry,phylogeny and body size

The size of fleshy fruits spans several orders of magnitude. However, the evolution of fruit size diversity is poorly understood. Fruit size diversity is hypothesised to result from several potential processes. The frugivore hypothesis postulates that different‐sized animal fruit consumers select for different‐sized fruits. The correlated selection hypothesis postulates that fruit size is allometrically related to other plant traits (e.g. leaf size, plant height); therefore differences in fruit size result from correlated evolution with other plant traits. We tested the frugivore and correlated selection hypotheses as potential explanations for fruit size diversity in two New Zealand study sites. We observed birds foraging for fruits over two fruiting seasons at each site and measured fruit size, leaf size and plant height in a total of 32 plant species. Relationships between average fruit size, leaf size, plant size and the average size of birds consuming each fruit species were then evaluated using phylogenetically independent contrasts. Similar results were obtained in both study sites. Fruit size was correlated with the size of avian fruit consumers, but was unrelated to leaf size or plant height. Therefore, results falsified the correlated selection hypothesis but failed to falsify the frugivore hypothesis. Although results suggest that frugivores may have influenced the evolution of fruit size in New Zealand, further study is needed to generate a mechanistic understanding of how frugivores may have selected for interspecific variation in fruit size.     

Removal of vertebrate-dispersed fruits in vegetation on fertile and infertile soils

Summary The hypothesis that more plant species with vertebrate-dispersed fruits occur on fertile soils because there is a greater probability of fruit removal from the parent plant was tested at 16 sites around Sydney, Australia. Removal rates from artificial fruit spikes were two and a half times greater on fertile than infertile soil sites, although this was not quite statistically significant. High variability in removal rate between sites was evident irrespective of fertility. Most removal occurred during the day indicating that birds were important consumers, rather than nocturnal mammals. Bird abundance and diversity did not differ between soil types. More frugivorous species were found in plant communities growing on fertile soil. Two models could explain the patterns observed. Firstly, plants with vertebrate-dispersed fruits could be favoured on fertile soils because of a high abundance of frugivorous birds accomplishing seed dispersal. Alternatively, plants with vertebrate-dispersed fruits could be favoured on fertile soil sites for some other reason and frugivorous birds could be attracted to these areas of abundant food. The correlation between soil fertility and the percentage of vertebrate-dispersed fruits was stronger than the correlation between soil fertility and removal rates and suggests that the second model is more likely to be true. Frugivorous birds are unlikely to be responsible for the high percentage of species with vertebrate-dispersed fruits in fertile soil environments.     

Global patterns in fruiting seasons

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

Water and fish select for fleshy fruits in tropical wetland forests

Adjacent floodplain and upland tropical forests experience the same temperature and precipitation regimes, but differ substantially in plant species composition and biotic interactions because of extensive flooding. We hypothesize that flooded forests filter fruiting traits linked to seed dispersal by water and fishes, such that selection by water and fish led to (1) trees that synchronize the timing of fruiting with annual floods, and (2) the evolution of fleshy tissues on fruits to improve buoyancy and increase fruit consumption rates by fish. To test this hypothesis, we compared plant communities in seasonally flooded forests and adjacent upland forest in terms of fruiting phenology, the frequency of trees bearing fleshy fruit, and the role of fleshy tissues in buoyancy and seed viability. Beta‐diversity in this system is high, with significant differences in species composition across habitats. As predicted, the production of ripe fleshy fruits was significantly greater in flooded than upland forests during the flood season. Furthermore, we found that trees with fleshy fruit were significantly more abundant in flooded forests even though species richness of fleshy fruit‐bearing trees was proportionally similar in flooded and upland forests. Additionally, fleshy pulp increased buoyancy. Likewise, time afloat decreased for denser fruit and those with high seed to pulp ratios. In concert, these results suggest that fleshy fruits in Neotropical floodplain forests facilitated hydrochory and ichthyochory. Once established, water and fish became important agents of selection on fruiting traits.     

Positive relationship between fruit removal by animals and seedling recruitment in a tropical forest

Fleshy-fruited plants rely on animal frugivores to disperse their seeds, and seed removal by frugivores may leave an imprint on seedling recruitment. However, to what extent plant–frugivore interactions are related to seedling recruitment has rarely been quantified at the community level, especially in species-rich tropical forests. In this study, we tested the effect of different plant traits on fruit removal by frugivores and tested the relative importance of fruit removal, plant traits and abiotic factors for seedling recruitment. We quantified plant–frugivore interactions of 22 fleshy-fruited plant species consumed by 56 diurnal frugivore species, and counted the number of seedlings that emerged along an elevational gradient in the Colombian Andes. We measured a set of plant traits (i.e., crop size; fruit size; seed load and mass; fruit nutritional contents), estimated the density of adult plants and recorded relevant abiotic factors (light, temperature and humidity). We found that fruit removal by frugivores was positively associated with crop size, but negatively associated with fruit length and unrelated to seed load and fruit nutritional content. Seedling densities were positively related to the density of adult plants, seed mass and fruit removal by animals. We found no relationship between abiotic factors and seedling recruitment. Our results indicate that fruit abundance and morphology are important determinants of fruit removal and that fruit removal is positively associated with seedling recruitment accounting for effects of species abundance and plant traits. We conclude that plant traits shape fruit removal and seedling recruitment at the community level, while these two crucial processes of forest regeneration are directly linked by seed dispersal of animals.     

Seed mass and the evolution of fleshy fruits in angiosperms

Fleshy fruits, like drupes and berries, have evolved many times through angiosperm history. Two hypotheses suggest that fleshy fruit evolution is related to changes in the seed mass fitness landscape. The reduced dispersal capability following from an increase in seed mass may be counterbalanced by evolution of traits mediating seed dispersal by animals, such as fleshy fruits. Alternatively, increasing availability and capabilities of frugivores promote evolution of fleshy fruits and allow an increase in seed size. Both these hypotheses predict an association between evolution of fleshy fruits and increasing seed size. We investigated patterns of fruit and seed evolution by contrasting seed mass between fleshy and non‐fleshy fruited sister clades. We found a consistent association between possession of fleshy fruits and heavier seeds. The direction of fruit type change did not alter this pattern; seed mass was higher in clades where fleshy fruits evolved and lower in clades where non‐fleshy fruits evolved, as compared to their sister clades. These patterns are congruent with the predictions from the two hypotheses, but other evidence is needed to distinguish between them. We emphasize the need to integrate studies of seed disperser effectiveness, seed morphology, and plant recruitment success to better understand the frugivores’ role in fleshy fruit evolution.     

Can Functional Traits Predict Ecological Interactions? A Case Study Using Rain forest Frugivores and Plants in Australia

In rain forest, the large numbers of species of fleshy-fruited plants and frugivorous animals result in a large number of potential fruit–frugivore interactions, which are challenging to survey in the field. Yet, knowledge of these relationships is needed to predict consequences of changes in the frugivore assemblage for seed dispersal. In the absence of comprehensive dietary information, it may be possible to delineate between frugivores that disperse different plants using ‘functional traits,’ or morphological and behavioral attributes of frugivores that interact with differences in salient characteristics of plant species. Here we use data on the consumption of 244 Australian rain forest plant species by 38 bird species to test for associations between patterns of frugivory and birds': (1) degree of frugivory, (2) gape width, and (3) seed treatment (seed crushing or seed dispersing). Degree of frugivory and gape width explain 74 percent of the variation in the sizes of fruits consumed by frugivorous birds. Among birds that consume a substantial dietary proportion of fruit, birds with wider gapes consume larger fruits. In contrast, this relationship was not shown by birds for which fruit is only a minor dietary component. Degree of frugivory and gape width, together with seed treatment, also strongly predict the overall taxonomic composition and diversity of plants consumed by bird species. Functional classifications of frugivore species may prove useful in developing a predictive understanding of fruit–frugivore interactions in other rain forest regions where detailed dietary information is not available for most frugivores.     

The role of fruit traits of bird-dispersed plants in invasiveness and weed risk assessment

Aim  Birds play a major role in the dispersal of seeds of many fleshy-fruited invasive plants. The fruits that birds choose to consume are influenced by fruit traits. However, little is known of how the traits of invasive plant fruits contribute to invasiveness or to their use by frugivores. We aim to gain a greater understanding of these relationships to improve invasive plant management.
Location  South-east Queensland, Australia.
Methods  We measure a variety of fruit morphology, pulp nutrient and phenology traits of a suite of bird-dispersed alien plants. Frugivore richness of these aliens was derived from the literature. Using regressions and multivariate methods, we investigate relationships between fruit traits, frugivore richness and invasiveness.
Results  Plant invasiveness was negatively correlated to fruit size, and all highly invasive species had quite similar fruit morphology [smaller fruits, seeds of intermediate size and few (< 10) seeds per fruit]. Lower pulp water was the only pulp nutrient trait associated with invasiveness. There were strong positive relationships between the diversity of bird frugivores and plant invasiveness, and in the diversity of bird frugivores in the study region and another part of the plants' alien range.
Main conclusions  Our results suggest that weed risk assessments (WRA) and predictions of invasive success for bird-dispersed plants can be improved. Scoring criteria for WRA regarding fruit size would need to be system-specific, depending on the fruit-processing capabilities of local frugivores. Frugivore richness could be quantified in the plant's natural range, its invasive range elsewhere, or predictions made based on functionally similar fruits.     

What makes a good neighborhood? Interaction of spatial scale and fruit density in the predator satiation dynamics of a masting juniper tree

Spatio-temporal variability in fruit production (masting) has been regarded as a key mechanism to increase plant fitness by reducing seed predation. However, considerably more effort has been devoted into understanding the consequences of temporal rather than spatial variations in fruit crop for plant fitness. In order to simultaneously evaluate both components, we quantify fruit production and pre-dispersal damage by three arthropod species (mites, chalcid wasps and moths) in the Spanish juniper (Juniperus thurifera) during 3 years in a spatially explicit context. Our aims were to assess (1) the interaction between fruit production and pre-dispersal fruit damage by arthropods, (2) the potential interference or competition between arthropods, and (3) the form of the phenotypic selection exerted by arthropods on fruit traits considering the spatial context. Arthropods damaged a substantial fraction of fruits produced by Spanish juniper with levels of damage showing sharp inter-annual variations. Fruit damage by mites was negatively related to yearly fruit crop and positively correlated at individual trees fruiting in consecutive years. Increased interspecific interference was an additional consequence of reduced fruit availability during small crop years. During a masting year, fruit damage by less mobile species such as mites was negatively affected by tree crop size, and no spatial structure was observed for mite damage. The incidence of chalcid wasps was low, so the spatial pattern of seed predation was unclear, and no preferences for fruit or seed traits were detected. Conversely, moths selected larger fruits and their incidence on trees was spatially aggregated up to 20 m, with predation levels being negatively affected by fruit abundance at the patch level, suggesting a positive density-dependent effect of neighbors on fruit output. These results highlight the importance of including the spatial component to understand complex species interactions at local scales.     

Functional heterogeneity in a plant–frugivore assemblage enhances seed dispersal resilience to habitat loss

The ability of ecosystems to maintain their functions after disturbance (ecological resilience) depends on heterogeneity in the functional capabilities among species within assemblages. Functional heterogeneity may affect resilience by determining multiplicity between species in the provision of functions (redundancy) and complementarity between species in their ability to respond to disturbances (response diversity), but also by promoting the maintenance of biological information that enables ecosystems to reorganize themselves (ecological memory). Here, we assess the role of the components of the functional heterogeneity of a plant–frugivore assemblage on the resilience of seed dispersal to habitat loss. For three years, we quantified the distributions of fruits, frugivorous thrushes (Turdus spp.) and dispersed seeds, as well as frugivore diet and movement, along a gradient of forest cover in N Spain. The abundances and the spatial distributions of fruits and birds varied between years. The different thrushes showed similar diets but differed in spatial behavior and response to habitat loss, suggesting the occurrence of both functional redundancy and response diversity. Forest cover and fruit availability affected the spatial distribution of the whole frugivore assemblage. Fruit tracking was stronger in years when fruits were scarcer but more widespread across the whole fragmented landscape, entailing larger proportions of seeds dispersed to areas of low forest cover and open microhabitats. Rather than depending on redundancy and/or response diversity, seed dispersal resilience mostly emerged from the ecological memory conferred by the inter‐annual variability in fruit production and the ability of thrushes to track fruit resources across the fragmented landscape. Ecological memory also derived from the interaction of plants and frugivores as source organisms (trees in undisturbed forest), mobile links (birds able to disperse seeds into the disturbed habitat), and biological legacies (remnant trees and small forest patches offering scattered fruit resources across the landscape).     

Frugivory on Persea lingue in temperate Chilean forests: interactions between fruit availability and habitat fragmentation across multiple spatial scales

Habitat degradation and fragmentation are expected to reduce seed dispersal rates by reducing fruit availability as well as the movement and abundance of frugivores. These deleterious impacts may also interact with each other at different spatial scales, leading to nonlinear effects of fruit abundance on seed dispersal. In this study we assessed whether the degradation and fragmentation of southern Chilean forests had the potential to restrict seed dispersal the lingue (Persea lingue) tree, a fleshy-fruited tree species. Of five frugivore bird species, the austral thrush (Turdus falcklandii) and the fire-eyed diucon (Xolmis pyrope) were the only legitimate seed dispersers as well as being the most abundant species visiting lingue trees. The results showed little or no direct effect of habitat fragmentation on seed dispersal estimates, possibly because the assemblage of frugivore birds was comprised habitat-generalist species. Instead, the number of fruits removed per focal tree exhibited an enhanced response to crop size, but only in the more connected fragments. In the fruit-richer fragment networks, there was an increased fragment-size effect on the proportion of fruits removed in comparison to fruit-poor networks in which the fragment size effect was spurious. We suggest that such nonlinear effects are widespread in fragmented forest regions, resulting from the link between the spatial scales over which frugivores sample resources and the spatial heterogeneity in fruiting resources caused by habitat fragmentation and degradation.     

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.