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).     

Forest cover,hunting pressure,and fruit availability influence seed dispersal in a forest‐savanna mosaic in the Congo Basin

Forest fragmentation, reduced forest cover, and hunting pressure are the main threats affecting animal‐mediated seed dispersal. However, their combined effects on seed dispersal rates have been simultaneously investigated only rarely, and never in Africa. We aimed to disentangle the effects of forest cover, hunting pressure, frugivore abundance, and fruit availability at the local and landscape scales on the seed dispersal rates of Staudtia kamerunensis (Myristicaceae). To estimate the percentages of seed dispersal failure (undispersed seeds), we quantitated fruit remains below fruiting trees distributed across five contrasting sites in a semi‐natural forest‐savanna mosaic in the Democratic Republic of Congo. We used statistical analyses accounting for spatial autocorrelation and found that forest cover in the surrounding landscape, hunting level, the associated abundance of dispersers, and fruit availability all had significant effects on the percentage of seed dispersal failure. The combination of high fruit availability and reduced abundance of seed dispersers could accelerate seed disperser satiation, causing the seed dispersal system to be saturated. Our study highlights how two major factors associated with anthropogenic activities, forest cover and hunting, affect seed dispersal by animals. These findings could have far‐reaching implications for our understanding of tree‐frugivore interactions and the conservation of tropical communities.     

Effects of zoochory on the spatial genetic structure of plant populations

Spatial genetic structure (SGS) of plants results from the nonrandom distribution of related individuals. SGS provides information on gene flow and spatial patterns of genetic diversity within populations. Seed dispersal creates the spatial template for plant distribution. Thus, in zoochorous plants, dispersal mode and disperser behaviour might have a strong impact on SGS. However, many studies only report the taxonomic group of seed dispersers, without further details. The recent increase in studies on SGS provides the opportunity to review findings and test for the influence of dispersal mode, taxonomic affiliation of dispersers and their behaviour. We compared the proportions of studies with SGS among groups and tested for differences in strength of SGS using Sp statistics. The presence of SGS differed among taxonomic groups, with reduced presence in plants dispersed by birds. Strength of SGS was instead significantly influenced by the behaviour of seed dispersal vectors, with higher SGS in plant species dispersed by animals with behavioural traits that result in short seed dispersal distances. We observed high variance in the strength of SGS in plants dispersed by animals that actively or passively accumulate seeds. Additionally, we found SGS was also affected by pollination and marker type used. Our study highlights the importance of vector behaviour on SGS even in the presence of variance created by other factors. Thus, more detailed information on the behaviour of seed dispersers would contribute to better understand which factors shape the spatial scale of gene flow in animal‐dispersed plant species.     

Seed fate and decision‐making processes in scatter‐hoarding rodents

A mechanistic understanding of seed movement and survival is important both for the development of theoretical models of plant population dynamics, spatial spread, and community assembly, and for the conservation and management of plant communities under global change. While models of wind‐borne seed dispersal have advanced rapidly over the past two decades, models for animal‐mediated dispersal have failed to make similar progress due to their dependence on interspecific interactions and complex, context‐dependent behaviours. In this review, we synthesize the literature on seed dispersal and consumption by scatter‐hoarding, granivorous rodents and outline a strategy for development of a general mechanistic seed‐fate model in these systems. Our review decomposes seed dispersal and survival into six distinct sub‐processes (exposure, harvest, allocation, preparation, placement, and recovery), and identifies nine intermediate (latent) variables that link physical state variables (e.g. seed and animal traits, habitat structure) to decisions regarding seed allocation to hoarding or consumption, cache placement and management, and deployment of radicle‐pruning or embryo excision behaviours. We also highlight specific areas where research on these intermediate relationships is needed to improve our mechanistic understanding of scatter‐hoarder behaviour. Finally, we outline a strategy to combine detailed studies on individual functional relationships with seed‐tracking experiments in an iterative, hierarchical Bayesian framework to construct, refine, and test mechanistic models for context‐dependent, scatter‐hoarder‐mediated seed fate.     

Retention time variability as a mechanism for animal mediated long-distance dispersal

Long-distance dispersal (LDD) events, although rare for most plant species, can strongly influence population and community dynamics. Animals function as a key biotic vector of seeds and thus, a mechanistic and quantitative understanding of how individual animal behaviors scale to dispersal patterns at different spatial scales is a question of critical importance from both basic and applied perspectives. Using a diffusion-theory based analytical approach for a wide range of animal movement and seed transportation patterns, we show that the scale (a measure of local dispersal) of the seed dispersal kernel increases with the organisms' rate of movement and mean seed retention time. We reveal that variations in seed retention time is a key determinant of various measures of LDD such as kurtosis (or shape) of the kernel, thinkness of tails and the absolute number of seeds falling beyond a threshold distance. Using empirical data sets of frugivores, we illustrate the importance of variability in retention times for predicting the key disperser species that influence LDD. Our study makes testable predictions linking animal movement behaviors and gut retention times to dispersal patterns and, more generally, highlights the potential importance of animal behavioral variability for the LDD of seeds.     

Destination‐based seed dispersal homogenizes genetic structure of a tropical palm

As the dominant seed dispersal agents in many ecosystems, frugivorous animals profoundly impact gene movement and fine‐scale genetic structure of plants. Most frugivores engage in some form of destination‐based dispersal, in that they move seeds towards specific destinations, resulting in clumped distributions of seeds away from the source tree. Molecular analyses of dispersed seeds and seedlings suggest that destination‐based dispersal may often yield clusters of maternal genotypes and lead to pronounced local genetic structure. The long‐wattled umbrellabird Cephalopterus penduliger is a frugivorous bird whose lek mating system creates a species‐specific pattern of seed dispersal that can potentially be distinguished from background dispersal processes. We used this system to test how destination‐based dispersal by umbrellabirds into the lek affects gene movement and genetic structure of one of their preferred food sources Oenocarpus bataua, a canopy palm tree. Relative to background dispersal processes, umbrellabird mating behaviour yielded more diverse seed pools in leks that included on average five times more seed sources and a higher incidence of long‐distance dispersal events. This resulted in markedly lower fine‐scale spatial genetic structure among established seedlings in leks than background areas. These species‐specific impacts of destination‐based dispersal illustrate how detailed knowledge of disperser behaviour can elucidate the mechanistic link driving observed patterns of seed movement and genetic structure.     

Seed dispersal in heterogeneous landscapes: linking field observations with spatially explicit models

Seed dispersal by animals drives persistence and colonization of the majority of fleshy‐fruited plants. Different factors have been identified as important in shaping patterns of seed deposition. These factors include habitat heterogeneity, movement patterns of frugivore species, and their feeding behavior. Most studies, however, have analysed the effect of one factor at the time, either with a modelling approach or from field observations. Here, we combine empirical data with spatially‐explicit models to explore the contribution of habitat availability and patterns of frugivore post‐feeding flight distances to the spatial patterns of seed dispersal. We found that both factors, distance from the mother tree and microhabitat identity, were shaping those patterns. Our results show that seed dispersal is tremendously heterogeneous and complex in space. The observed seed shadow is not the result of a single process but instead an intricate combination of distance and habitat selection. We suggest that the apparent combination of processes results from the simultaneous effects of distinct functional groups of frugivores.     

植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

What's your move? Movement as a link between personality and spatial dynamics in animal populations

Recent studies have established the ecological and evolutionary importance of animal personalities. Individual differences in movement and space‐use, fundamental to many personality traits (e.g. activity, boldness and exploratory behaviour) have been documented across many species and contexts, for instance personality‐dependent dispersal syndromes. Yet, insights from the concurrently developing movement ecology paradigm are rarely considered and recent evidence for other personality‐dependent movements and space‐use lack a general unifying framework. We propose a conceptual framework for personality‐dependent spatial ecology. We link expectations derived from the movement ecology paradigm with behavioural reaction‐norms to offer specific predictions on the interactions between environmental factors, such as resource distribution or landscape structure, and intrinsic behavioural variation. We consider how environmental heterogeneity and individual consistency in movements that carry‐over across spatial scales can lead to personality‐dependent: (1) foraging search performance; (2) habitat preference; (3) home range utilization patterns; (4) social network structure and (5) emergence of assortative population structure with spatial clusters of personalities. We support our conceptual model with spatially explicit simulations of behavioural variation in space‐use, demonstrating the emergence of complex population‐level patterns from differences in simple individual‐level behaviours. Consideration of consistent individual variation in space‐use will facilitate mechanistic understanding of processes that drive social, spatial, ecological and evolutionary dynamics in heterogeneous environments.     

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 tracking,frugivore satiation,and their consequences for seed dispersal

Vertebrate frugivore communities are easily satiated by abundant fruit supplies and, contrary to abiotic dispersal agents, typically disperse only part of the available seed pool. This frugivore satiation is likely to be a widespread phenomenon and should be an influential predictor of plants’ ability to disperse their offspring to suitable establishment sites; yet it has never been systematically quantified. Here I investigate patterns of fruit abundance, frugivore activity and frugivore satiation, and their consequences for seed dispersal in the fleshy-fruited tree Frangula alnus. Based on constant-effort seed trapping conducted over 3 years, I assess densities of total and frugivore-consumed seedfall across two spatial (within/between populations) and two temporal (within/between ripening seasons) scales. Furthermore, I examine relationships between fruit abundance and the amount of seeds that are actually dispersed away from fruiting trees. Frugivore activity tightly matched fruit abundance, although some differences existed between scales. This marked fruit tracking did not prevent a significant frugivore satiation, however, and only 53% of the available fruit crops were actually consumed. The extent of satiation varied most at the within-population level, likely due to the territorial behaviour of important frugivore species. In contrast, levels of satiation remained remarkably invariable through time, suggesting that frugivores behave as opportunists and closely adjust the composition of their diet to the available food supply. Overall, greater fruit abundance resulted in a higher proportion of seeds falling beneath fruiting trees, but it also helped increase the (absolute) number of seeds dispersed. This study shows that frugivore satiation can be an important phenomenon even when frugivores tightly track fruit abundance. Its negative effects on recruitment may be attenuated, however, if greater fruit crops help increase population-wide frugivore activity and the amount of seeds being dispersed to suitable establishment sites.     

Exploring the interaction of avian frugivory and plant spatial heterogeneity and its effect on seed dispersal kernels using a simulation model

Seed dispersal by avian frugivores is one of the key processes influencing plant spatial patterns, but may fail if there is disruption of plant–frugivore mutualisms, such as decline in abundance of dispersers, fragmentation of habitat, or isolation of individual trees. We used simulation model experiments to examine the interaction between frugivore density and behaviour and the spatial arrangement of fruiting plants and its effect on seed dispersal kernels. We focussed on two New Zealand canopy tree species that produce large fruits and are dispersed predominantly by one avian frugivore (Hemiphaga novaeseelandiae). Although the mean seed dispersal distance decreased when trees became more aggregated, there were more frugivore flights between tree clusters, consequently stretching the tails of the dispersal kernels. Conversely, when trees were less aggregated in the landscape, mean dispersal distances increased because seeds were deposited over larger areas, but the kernels had shorter tails. While there were no statistically meaningful changes in kernel parameters when frugivore density changed, decreases in density did cause a proportional reduction in the total number of dispersed seeds. However, birds were forced to move further when fruit availability and fruit ripening were low. Sensitivity analysis showed that dispersal kernels were primarily influenced by the model parameters relating to disperser behaviour, especially those determining attractiveness based on distance to candidate fruiting trees. Our results suggest that the spatial arrangement of plants plays an important role in seed dispersal processes – although tree aggregation curbed the mean seed dispersal distance, it was accompanied by occasional long distance events, and tree dispersion caused an increase in mean dispersal distance, both potentially increasing the probability of seeds finding suitable habitats for germination and growth. Even though low frugivore densities did not cause dispersal failure, there were negative effects on the quantity of seed dispersal because fewer seeds were dispersed.     

Seed Dispersal by Avian Frugivores: Non‐random Heterogeneity at Fine Scales

Seed dispersal studies have primarily examined dispersal as a function of distance from the parent tree and/or heterogeneity in dispersal due to animal use of nesting, roosting and sleeping sites. However, non‐random heterogeneity in seed dispersal is also likely to result from the post‐foraging behavior and movement of frugivores which prefer certain trees. To characterize variation in seed rain at fine scales, we studied the dispersal curve of Prunus ceylanica, a primarily bird‐dispersed species. We compared seed rain at conspecifics, heterospecific fruiting trees with similar frugivore assemblages, emergent trees, and the landscape surrounding these trees. Seed rain of P. ceylanica was found to peak globally under the canopy of conspecifics but to peak locally under the canopy and immediate neighborhood of heterospecific fruiting trees. Our results demonstrate that seed rain is highly clumped even at fine spatial scales. A large proportion of seeds are dispersed in specific, localized regions. This variation can have important implications for plant population dynamics and might significantly alter the impact of post‐dispersal processes. Seed dispersal models may need to incorporate this heterogeneity to explain manifestations of spatially explicit dynamics like mixed species ‘orchards’.     

Changes of effective gene dispersal distances by pollen and seeds across successive life stages in a tropical tree

Pollen and seed dispersal are the two key processes in which plant genes move in space, mostly mediated by animal dispersal vectors in tropical forests. Due to the movement patterns of pollinators and seed dispersers and subsequent complex spatial patterns in the mortality of offspring, we have little knowledge of how pollinators and seed dispersers affect effective gene dispersal distances across successive recruitment stages. Using six highly polymorphic microsatellite loci and parentage analyses, we quantified pollen dispersal, seed dispersal, and effective paternal and maternal gene dispersal distances from pollen‐ and seed‐donors to offspring across four recruitment stages within a population of the monoecious tropical tree Prunus africana in western Kenya. In general, pollen‐dispersal and paternal gene dispersal distances were much longer than seed‐dispersal and maternal gene dispersal distances, with the long‐distance within‐population gene dispersal in P. africana being mostly mediated by pollinators. Seed dispersal, paternal and maternal gene dispersal distances increased significantly across recruitment stages, suggesting strong density‐ and distance‐dependent mortality near the parent trees. Pollen dispersal distances also varied significantly, but inconsistently across recruitment stages. The mean dispersal distance was initially much (23‐fold) farther for pollen than for seeds, yet the pollen‐to‐seed dispersal distance ratio diminished by an order of magnitude at later stages as maternal gene dispersal distances disproportionately increased. Our study elucidates the relative changes in the contribution of the two processes, pollen and seed dispersal, to effective gene dispersal across recruitment. Overall, complex sequential processes during recruitment contribute to the genetic make‐up of tree populations. This highlights the importance of a multistage perspective for a comprehensive understanding of the impact of animal‐mediated pollen and seed dispersal on small‐scale spatial genetic patterns of long‐lived tree species.     

The relative contributions of seed and pollen dispersal to gene flow and genetic diversity in seedlings of a tropical palm

Seed and pollen dispersal shape patterns of gene flow and genetic diversity in plants. Pollen is generally thought to travel longer distances than seeds, but seeds determine the ultimate location of gametes. Resolving how interactions between these two dispersal processes shape microevolutionary processes is a long‐standing research priority. We unambiguously isolated the separate and combined contributions of these two dispersal processes in seedlings of the animal‐dispersed palm Oenocarpus bataua to address two questions. First, what is the spatial extent of pollen versus seed movement in a system characterized by long‐distance seed dispersal? Second, how does seed dispersal mediate seedling genetic diversity? Despite evidence of frequent long‐distance seed dispersal, we found that pollen moves much further than seeds. Nonetheless, seed dispersal ultimately mediates genetic diversity and fine‐scale spatial genetic structure. Compared to undispersed seedlings, seedlings dispersed by vertebrates were characterized by higher female gametic and diploid seedling diversity and weaker fine‐scale spatial genetic structure for female gametes, male gametes and diploid seedlings. Interestingly, the diversity of maternal seed sources at seed deposition sites (N _em) was associated with higher effective number of pollen sources (N _ep), higher effective number of parents (N _e) and weaker spatial genetic structure, whereas seed dispersal distance had little impact on these or other parameters we measured. These findings highlight the importance maternal seed source diversity (N _em) at frugivore seed deposition sites in driving emergent patterns of fine‐scale genetic diversity and structure.     

Patterns of resource tracking by avian frugivores at multiple spatial scales: two case studies on discordance among scales

Scaling is relevant for the analysis of plant‐frugivore interaction, since the ecological and evolutionary outcomes of seed dispersal depend on the spatial and temporal scale at which frugivory patterns emerge. We analyse the relationship between fruit abundance and frugivore activity at local and landscape spatial scales in two different systems composed, respectively, by the bird‐dispersed woody plants Juniperus communis and Bursera fagaroides, and their frugivore assemblages. We use a hierarchical approach of nested patchiness of fruit‐resource, where patches are defined by individual plants within site, at the local scale, and by sites within region, at the landscape scale. The structure of patches is also described in terms of contrast (differences in fruit availability among patches) and aggregation (spatial distribution of patches). For J. communis, frugivore activity was positively related to fruit availability at the landscape scale, this pattern seldom emerging at the local scale; conversely, B. fagaroides showed a general trend of positive local pattern that disappeared at the landscape scale. These particular trends might be partially explained by differences in contrast and aggregation. The strong contrast among plants within site together with a high aggregation among sites would promote the B. fagaroides pattern to be only local, whereas in J. communis, low aggregation among sites within region would favour a sharp landscape‐scale pattern. Both systems showed discordant patterns of fruit‐resource tracking among consecutive spatial scales, but the sense of discordance differed among systems. These results, and the available multi‐scale frugivory data, suggest that discordance among successive scales allows to link directly frugivory patterns to resource‐tracking mechanisms acting at particular scales, resulting, thus, more informative than concordance observational data, in which landscape patterns might result from accumulated effect of local mechanisms. In this context, we propose new methodological approaches for a better understanding of the hierarchical behavioural mechanisms underpinning the multi‐scale resource tracking by frugivores.     

Variation in seed dispersal effectiveness: the redundancy of consequences in diversified tropical frugivore assemblages

Plant–frugivore mutualistic assemblages frequently combine multiple, complementary or not (i.e. redundant), distinct effects of animal species. To a large extent, the outcomes of these interactions crucially depend on the delayed consequences of frugivore effectiveness on plant recruitment. We evaluated seed dispersal effectiveness for three plant species in a Brazilian Atlantic forest with a marked habitat heterogeneity defined by bamboo and non‐bamboo patches. Twenty one, 23 and 14 bird species ate fruits of Euterpe edulis, Sloanea guianensis and Virola bicuhyba trees, respectively. For both Euterpe and Virola, visitation rate was the variable contributing for most variance across frugivore species in the quantitative component of effectiveness (QC, which depends on the combined effects of interaction frequency and per‐interaction effect), while the number of fruits manipulated/visit had the greatest contribution in Sloanea. By combining observational data and experimental seed addition for Euterpe we tested for consistent functional patterns among species in the frugivore assemblage, extending beyond the fruit removal stage. Rankings of QC across Euterpe frugivores remained consistent with their relative contributions to fruit removal and, importantly, with their contributions to seedling establishment. Yet, QC of effectiveness across Euterpe frugivores were more homogeneous at the fruit removal and dispersal stages (contribution to seed dispersal) than for the delayed, dissemination and post‐dispersal effects on recruitment. High complementarity of diversified frugivore assemblages may increase through added variance in their delayed effects related to qualitative components of effectiveness. Our results underscore the importance of assessing how dispersal services provided by mutualistic frugivores play complementary, rather than redundant, roles in seed dispersal within heterogeneous landscapes. Such ecological outcomes highlight the value of combining observational and experimental field designs to assess functional diversity patterns of tropical frugivore assemblages and delayed effects of their interactions with plants.     

Fragmentation, habitat composition and the dispersal/predation balance in interactions between the Mediterranean myrtle and avian frugivores

Human‐induced fragmentation and disturbance of natural habitats can shift abundance and composition of frugivore assemblages, which may alter patterns of frugivory and seed dispersal. However, despite their relevance to the functioning of ecosystems, plant‐frugivore interactions in fragmented areas have been to date poorly studied. I investigated spatial variation of avian frugivore assemblages and fruit removal by dispersers and predators from Mediterranean myrtle shrubs (Myrtus communis) in relation to the degree of fragmentation and habitat features of nine woodland patches (72 plants). The study was conducted within the chronically fragmented landscape of the Guadalquivir Valley (SW Spain), characterized by ~1% of woodland cover. Results showed that the abundance and composition of the disperser guild was not affected by fragmentation, habitat features or geographical location. However, individual species and groups of resident/migrant birds responded differently: whereas resident dispersers were more abundant in large patches, wintering dispersers were more abundant in fruit‐rich patches. Predator abundances were similar between patches, although the guild composition shifted with fragmentation. The proportion of myrtle fruits consumed by dispersers and predators varied greatly between patches, but did not depend on bird abundances. The geographical location of patches determined the presence or absence of interactions between myrtles and seed predators (six predated and three non‐predated patches), a fact that greatly influenced fruit dispersal success. Moreover, predation rates were lower (and dispersal rates higher) in large patches with fruit‐poor heterospecific environments (i.e. dominated by myrtle). Predator satiation and a higher preference for heterospecific fruits by dispersers may explain these patterns. These results show that 1) the frugivore assemblage in warm Mediterranean lowlands is mostly composed of fragmentation‐tolerant species that respond differently to landscape changes; and 2) that the feeding behaviour of both dispersers and predators influenced by local fruit availability may be of great importance for interpreting patterns of frugivory throughout the study area.     

Antelope mating strategies facilitate invasion of grasslands by a woody weed

Intra and interspecific variation in frugivore behaviour can have important consequences for seed dispersal outcomes. However, most information comes from among‐species comparisons, and within‐species variation is relatively poorly understood. We examined how large intraspecific differences in the behaviour of a native disperser, blackbuck antelope Antilope cervicapra, influence dispersal of a woody invasive, Prosopis juliflora, in a grassland ecosystem. Blackbuck disperse P. juliflora seeds through their dung. In lekking blackbuck populations, males defend clustered or dispersed mating territories. Territorial male movement is restricted, and within their territories males defecate on dung‐piles. In contrast, mixed‐sex herds range over large areas and do not create dung‐piles. We expected territorial males to shape seed dispersal patterns, and seed deposition and seedling recruitment to be spatially localized. Territorial males had a disproportionately large influence on seed dispersal. Adult males removed twice as much fruit as females, and seed arrival was disproportionately high on territories. Also, because lek‐territories are clustered, seed arrival was spatially highly concentrated. Seedling recruitment was also substantially higher on territories compared with random sites, indicating that the local concentration of seeds created by territorial males continued into high local recruitment of seedlings. Territorial male behaviour may, thus, result in a distinct spatial pattern of invasion of grasslands by the woody P. juliflora. An ex situ experiment showed no beneficial effect of dung and a negative effect of light on seed germination. We conclude that large intraspecific behavioural differences within frugivore populations can result in significant variation in their effectiveness as seed dispersers. Mating strategies in a disperser could shape seed dispersal, seedling recruitment and potentially plant distribution patterns. These mating strategies may aid in the spread of invasives, such as P. juliflora, which could, in turn, negatively influence the behaviour and ecology of native dispersers.