Unravelling seed dispersal through fragmented landscapes: Frugivore species operate unevenly as mobile links

Seed dispersal constitutes a pivotal process in an increasingly fragmented world, promoting population connectivity, colonization and range shifts in plants. Unveiling how multiple frugivore species disperse seeds through fragmented landscapes, operating as mobile links, has remained elusive owing to methodological constraints for monitoring seed dispersal events. We combine for the first time DNA barcoding and DNA microsatellites to identify, respectively, the frugivore species and the source trees of animal‐dispersed seeds in forest and matrix of a fragmented landscape. We found a high functional complementarity among frugivores in terms of seed deposition at different habitats (forest vs. matrix), perches (isolated trees vs. electricity pylons) and matrix sectors (close vs. far from the forest edge), cross‐habitat seed fluxes, dispersal distances and canopy‐cover dependency. Seed rain at the landscape‐scale, from forest to distant matrix sectors, was characterized by turnovers in the contribution of frugivores and source‐tree habitats: open‐habitat frugivores replaced forest‐dependent frugivores, whereas matrix trees replaced forest trees. As a result of such turnovers, the magnitude of seed rain was evenly distributed between habitats and landscape sectors. We thus uncover key mechanisms behind “biodiversity–ecosystem function” relationships, in this case, the relationship between frugivore diversity and landscape‐scale seed dispersal. Our results reveal the importance of open‐habitat frugivores, isolated fruiting trees and anthropogenic perching sites (infrastructures) in generating seed dispersal events far from the remnant forest, highlighting their potential to drive regeneration dynamics through the matrix. This study helps to broaden the “mobile‐link” concept in seed dispersal studies by providing a comprehensive and integrative view of the way in which multiple frugivore species disseminate seeds through real‐world landscapes.     

Spatial–temporal distribution of ornithochorous seeds from an Elaeagnus umbellata community dominating a riparian habitat

The dispersal efficiency and potential distribution of ornithochorous seeds of Elaeagnus umbellata in a riparian habitat were evaluated to clarify this species' establishment site in relation to the disturbance regime of the floodplain. Fruit removal by avian frugivores was monitored using fruit bags, and the spatial distribution of excreted seeds was quantified by seed traps set randomly on a gravel bar as an isolated seed source in the Yoshino River throughout an autumn fruiting season. Although more than 45% of the fruits remained on the twigs in the fruit bags, almost all fruits on the control twigs without fruit bags were exploited by the beginning of January. The fruit removal rate and seed dispersal distance were positively correlated with an increase in wintering bird species and their abundance. Intact bird‐dispersed seeds of E. umbellata were trapped within a 400‐m range and damaged seeds were limited to traps set within 50 m from the seed source. Frugivore behavior, such as feces excretion on rocks near water drinking sites and perching on surrounding woodland, greatly influenced the spatial and temporal dispersal pattern of the seed rain. In the present study, the avian frugivores showed upstream seed dispersal; thus, in years with stochastic autumnal floods, secondary dispersal via hydrochory downstream may be facilitated. The intensive seed dispersal in E. umbellata indicates that the present distribution of parent trees in the restricted elevation range of the gravel bars is the result of survival through disturbance, rather than seed dispersal limitation.     

Enhanced seed dispersal of Prunus africana in fragmented and disturbed forests?

Forest destruction and disturbance can have long-term consequences for species diversity and ecosystem processes such as seed dispersal. Understanding these consequences is a crucial component of conserving vulnerable ecosystems. In the heavily fragmented and disturbed Kakamega Forest, western Kenya, we studied seed dispersal of Prunus africana (Rosaceae). In the main forest, five forest fragments, and differently disturbed sites, we quantified the overall frugivore community as an indicator for species diversity. Furthermore, we determined the frugivores on 28 fruiting P. africana trees, estimated seed dispersal, crop size and the general fruit availability of surrounding trees. During the overall frugivore census we recorded 49 frugivorous species; 36 of them were observed visiting P. africana trees and feeding on their fruits. Although overall frugivore species richness was 1.1 times lower in fragments than in main forest sites and 1.02 times higher in highly disturbed than in less disturbed sites, P. africana experienced 1.1 times higher numbers of frugivores in fragments than in main forest sites and 1.5 times higher numbers of frugivores in highly disturbed than in less disturbed sites. Correspondingly, seed dispersal was 1.5 times higher in fragments than in main forest sites and 1.5 times higher in more disturbed than less disturbed sites. Fruit availability of surrounding trees and crop size influenced the number of visitors to some degree. Thus, the number of dispersed seeds seemed to be slightly higher in fragmented and highly disturbed sites. This indicates that loss of single species does not necessarily lead to a decrease of ecosystem services. However, loss of diversity could be a problem in the long term, as a multitude of species might act as buffer against future environmental change.     

Impact of avian frugivores on dispersal and recruitment of the invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> in an agricultural landscape

Although seed dispersal is considered to be a key process determining the spatial structure and spread of non-native plant populations, few studies have explicitly addressed the link between dispersal vector behaviour, seed distribution and seedling recruitment to gain insight into the process of exotic species invasion within a fragmented landscape context. The present study analyses the relationship between avian frugivory and spatial patterns of seed deposition and seedling recruitment for an expanding population of the invasive Prunus serotina in a hedgerow network landscape in Flanders, Belgium. We quantified fruit production, observed frugivores, and determined the spatial distribution of bird droppings and P. serotina seedlings. A relatively diverse assemblage of frugivores visited P. serotina seed trees, with Columba palumbus and Turdus merula being by far the most important dispersers. Landscape structure strongly affected dispersal vector behaviour and the spatial distribution of perching birds, droppings and seedlings. Frugivorous birds non-randomly dispersed seeds to perching sites and an association between perching birds, seed deposition and seedling recruitment was found. Results indicate that landscape structure contributes to non-random seed deposition of P. serotina by common local frugivores. Cutting the larger seed trees is proposed as the most feasible measure to slow down the invasion rate.     

Monitoring seed dispersal at isolated standing trees in tropical pastures: consequences for local species availability

The tropical rain forest landscape has been transformed to a mosaic composed of patches of crops, secondary vegetation and remnant forest fragments of different shapes and sizes. Isolation of patches and fragments is a critical issue in the maintenance of local species diversity. In this study we focus on the dispersal of propagules by birds to understant the movement of plants between landscape components. Seed deposition and the behavior of frugivorous birds were monitored at four isolated fig trees (Ficus yoponensis and F. aurea) in man-made pastures. Seed deposition was measured by trapping seeds under canopy trees for six months and by direct observation of bird visits to the four trees for one year. Seed deposition densities were 465, 614, 632 and 1097 seeds/m² accumulated over six months under each of the four trees. We recorded 8268 seeds of 107 species under the trees, among them, 6726 seeds (81%) were of 56 species dispersed by vertebrate frugivores. Seeds of tree species accounted for 26% of the total species. Seventy-three species of birds perched in the observed trees, and 3344 visits were made by 47 species of frugivores. Frugivorous birds occurred in two groups: resident species nesting in the pastures and resident species nesting elsewhere. Propagule exchange between landscape components is clearly influenced by the behavior of these two groups. Structure and dynamics of the landscape depend on plant species availability within the mosaic. This availability is high and suggests possibilities for the management of the local species diversity of tropical rain forests.     

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’.     

Avian frugivory and seed dispersal of a large fruited tree in an Indian moist deciduous forest

Seed dispersal systems in degraded areas can be compromised following the decline of large-bodied frugivore populations responsible for their dispersal. In this context we examined the seed dispersal ecology of a large fruited deciduous tree (Dillenia pentagyna) along a forest degradation gradient in India. We examined the effect of structural components of vegetation and frugivore foraging behavior on D. pentagyna seed dispersal. Depauperate mammalian community and declined large avian frugivores e.g. hornbills in our study site make this system a specialized one and currently dependent on only two large bodied avian frugivores. Seed dispersal followed an overall leptokurtic pattern and the seed dispersal kernels were best explained by an inverse power function. Seed dispersal kernels in dense forest indicated longer dispersal distances than moderately dense forest and degraded forest. In degraded areas, no dispersal away from the crown was recorded for D. pentagyna and it occurred at low density. Canopy foliage abundance of the surrounding vegetation of the focal trees was best explained by quantity of seed dispersal by large avian frugivores. The number of avian frugivore species those are effective disperser of D. pentagyna decreased along the degradation gradient. Avian frugivore behavior in terms of visitation and seed swallowed is a determining factor that controls quantity of seed dispersal. Our study underscores deleterious impact of forest degradation on avian disperser community which in turn would affect regeneration capacity of degraded forest.     

Seed Banks in Savanna,Forest Fragments,and Continuous Forest Edges Differ in a Tropical Dry Forest in Madagascar

Rapid deforestation has fragmented habitat across the landscape of Madagascar. To determine the effect of fragmentation on seed banks and the potential for forest regeneration, we sampled seed viability, density and diversity in 40 plots of 1 m² in three habitat types: forest fragments, the near edge of continuous forest, and deforested savanna in a highly fragmented dry deciduous forest landscape in northwestern Madagascar. While seed species diversity was not different between forest fragments and continuous forest edge, the number of animal‐dispersed seeds was significantly higher in forest fragments than in continuous forest edge, and this pattern was driven by a single, small‐seeded species. In the savanna, seeds were absent from all but three of the 40 plots, indicating that regeneration potential is low in these areas. Several pre‐ and post‐dispersal biotic and abiotic factors, including variation in the seed predator communities and edge effects could explain these findings. Understanding the extent to which seed dispersal and seed banks influence the regeneration potential of fragmented landscapes is critical as these fragments are the potential sources of forest expansion and re‐connectivity.     

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).     

Forest cover and fruit crop size differentially influence frugivory of select rainforest tree species in Western Ghats,India

Forest fragmentation and habitat loss are major disruptors of plant–frugivore interactions, affecting seed dispersal and altering recruitment patterns of the dependent tree species. In a heterogeneous production landscape (primarily tea and coffee plantations) in the southern Western Ghats, India, we examined effects of surrounding forest cover and fruit crop size on frugivory of four rainforest bird-dispersed tree species (N = 131 trees, ≥30 trees per species, observed for 623 hr). Frugivore composition differed among the four tree species with the large-seeded Canarium strictum and Myristica dactyloides being exclusively dependent on large-bodied avian frugivores, whereas medium-seeded Persea macrantha and Heynea trijuga were predominantly visited by small-bodied and large-bodied avian frugivores, respectively. Using the seed-dispersal-effectiveness framework, we identified effective frugivores and examined their responses to forest cover and fruit crop size. Results were idiosyncratic and were governed by plant and frugivore traits. Visitations to medium-seeded Persea had a positive relationship with forest cover but the relationship was negative for the large-seeded Myristica. In addition, two of the three effective frugivores for Persea responded to the interactive effect of forest cover and fruit crop size. Frugivore visitations to Heynea were not related to forest cover or fruit crop, and there were too few visitations to Canarium to discern any trends. These results highlight the context-specific responses of plant–frugivore interactions to forest cover and fruit crop size influenced by plant and frugivore traits.     

Effects of site and fruit size on the composition of avian frugivore assemblages in a fragmented Afrotropical forest

Factors influencing the interaction between fruiting trees and their frugivorous seed dispersers in fragmented Afrotropical landscapes are poorly known. With the use of Mantel statistics we analysed assemblages of frugivorous birds on 58 individual trees belonging to 11 species growing in seven Kenyan cloud forest fragments. Overall, frugivores showed little specialization on particular trees. Fruit size explained a substantial amount of the variation in frugivore assemblages among different tree species at the same site. In addition, frugivore assemblages on conspecific trees were significantly more similar when the trees occurred at the same site. This location effect was attributable to the different sites and forest fragments (of different sizes and disturbance levels) varying in the densities and composition of their avian frugivores, vegetation composition and tree fruiting phenologies. It was consolidated further by the low mobility of most of these avian frugivores, particularly their reluctance to cross between forest fragments. Habitat disturbance and fragmentation may therefore have affected fruit selection, with implications for both seed dispersal and regeneration.     

Toucans (Ramphastos ambiguus) facilitate resilience against seed dispersal limitation to a large‐seeded tree (Virola surinamensis) in a human‐modified landscape

Large‐seeded plants may suffer seed dispersal limitation in human‐modified landscapes if seed dispersers are absent or unable to disperse their seeds. We investigated dispersal limitation for the large‐seeded tree Virola surinamensis in a human‐modified landscape in southern Costa Rica. During two fruiting seasons, we monitored crop size, seed removal rates, the number of fruiting conspecifics within 100 m, and feeding visitation rates by frugivores at trees located in high and low forest disturbance conditions. Seed removal rates and the total number of seeds removed were high regardless of the disturbance level, but these parameters increased with tree crop size and decreased with the number of fruiting V. surinamensis trees within a 100 m radius. Trees at low disturbance levels were more likely to be visited by seed dispersers. Black mandibled toucans (Ramphastos ambiguus) and spider monkeys (Ateles geoffroyi) were the most important seed dispersers, based on visitation patterns and seed removal rates. Spider monkey feeding visits were more frequent at high disturbance levels, but the monkeys preferentially visited isolated trees with large yields and surrounded by a low number of fruiting Virola trees within 100 m. Toucan visitation patterns were not constrained by any of the predictors and they visited trees equally across the landscape. We suggest that isolated and highly fecund Virola trees are an important food resource for spider monkeys in human‐modified landscapes and that toucans can provide resilience against seed dispersal limitations for large‐seeded plants in human‐modified landscapes in the absence of hunting.     

Can remnant frugivore species effectively disperse tree seeds in secondary tropical rain forests?

Seed dispersal by frugivores in tropical rain forests is important for maintaining viable tree populations. Over the years, vertebrate assemblages in tropical forests have been altered by anthropogenic disturbances, leading to concerns about the ability of remnant vertebrates to substitute for the lost or declining vertebrate populations. We compared vertebrate composition and frugivore visitation rates as an indirect measure of rate of seed dispersal in three tropical rain forests in Uganda, namely Mabira, Budongo and Kibale Forests. Mabira is highly disturbed, Kibale is little and Budongo is intermediate. The aim was to determine whether vertebrate assemblages in differentially disturbed forests had comparable abilities to disperse seeds and whether tree species were equally vulnerable to loss of seed dispersers. Assemblages of forest generalist species were similar in all forests, but specialists were less abundant in the heavily disturbed forest. Remnant frugivores in the heavily disturbed forest were mainly small-bodied species that spat seeds beneath fruiting trees compared to large-bodied species observed in the less disturbed forests that ingested and carried away the seeds. We postulate that the quantity of seeds dispersed in heavily disturbed forests is much reduced due to low visitation rates of frugivores and the absence of large frugivores that consume large quantities of fruit. The quality of seed dispersal is affected as well by the distance over which seeds are moved. Assessment of vulnerability of trees shows no evidence for disperser substitution for trees producing large fruits. Fruit trees with low nutritional contents and digestibility were least visited in frugivore-impoverished forests. The loss of large specialist frugivores is likely to affect recruitment of many trees, especially of species that cannot establish beneath adult conspecifics.     

Patterns of Seed Dispersal and Dispersal Failure in a Hawaiian Dry Forest Having Only Introduced Birds

Dry forests are among the most endangered natural communities in the Hawaiian Islands. Most have been reduced to isolated trees and small forest fragments in which native tree species reproduce poorly. The replacement of native birds by introduced generalists may be contributing to dry forest decline through modification of seed dispersal patterns. To document seed dispersal by introduced birds, we conducted foraging observations on fleshy-fruited trees and measured seed rain under trees and in adjacent open areas for 1 year in a dry forest dominated by native trees. Although trees covered only 15.2 percent of the study area, 96.9 percent of the bird-dispersed seeds were deposited beneath them. The Japanese white-eye (Zosterops japonicus) was the principal dispersal agent. Among bird-dispersed seeds, those of the invasive tree Bocconia frutescens accounted for 75 percent of all seeds collected beneath trees (14.8 seeds/m²/yr) and the invasive shrub Lantana camara accounted for 17 percent. Although nearly 60 percent of the reserve's native woody species possess fleshy fruits, introduced birds rarely disperse their seeds. Native trees accounted for <8 percent of all bird-dispersed seeds and are consequently experiencing dispersal failure by falling directly under parent trees. Smaller-seeded non-native plants, in contrast, may be benefiting from dispersal by introduced birds. Current dispersal patterns suggest that these readily disseminated non-native plants may eventually replace the remaining native flora.     

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.     

Early fate of Myristica hypargyraea seeds dispersed by Ducula pacifica in Tonga,Western Polynesia

Abstract Although pigeons from the genus Ducula are considered among the best avian dispersers of large seeds in Asia and the Pacific, little has been documented on their role. The early fate of dispersed and undispersed seeds of the large‐seeded tree Myristica hypargyraea A. Gray was studied in order to understand the advantage of seed dispersal by the Pacific Pigeon, Ducula pacifica Gmelin in Tonga. Frequency of visits by frugivores to fruiting trees and dispersal distance of seeds were measured. Pre‐dispersal vertebrate seed predation was assessed, then post‐dispersal predation was measured over 160 days. Overall, pre‐dispersal seed predation by parrots was low but variable among trees sampled. Most seeds (54.7%) in the study area were estimated to be dispersed by D. pacifica; 79.7% of those ingested were expelled directly beneath conspecific fruiting crowns, 20% were dispersed locally and < 0.3% were dispersed more than 300 m into a different forest type. Flying foxes (Pteropus tonganus Quoy and Gaimard) dispersed very few seeds (0.7%) and all were dropped below fruiting crowns. Between 4% and 39% of dispersed and undispersed seeds were still viable, or had established seedlings after 160 days. Most seeds had been removed or killed by rats, and seed survival was highest for locally dispersed seeds (approx. 20 m from source trees and within the M. hypargyraea forest). Although D. pacifica was the only frugivore observed to disperse seeds into this higher zone of survival, overall they did not confer a great advantage to seed survival since significant numbers of seeds/seedlings also persisted under fruiting crowns (27% under crowns compared with 39% locally dispersed). Nevertheless, D. pacifica was the only vector by which seeds were regularly moved within the M. hypargyraea forest and over longer distances, and hence, D. pacifica still plays a significant role in the regeneration of M. hypargyraea.     

The role of avian frugivores in germination of seeds of fleshy-fruited invasive alien plants

Many highly invasive plant species have fleshy fruits which are eaten by native frugivorous animals. These frugivores play an important role in long-distance seed dispersal, and may also affect germination success. The aim of this study was to determine whether generalist frugivores enhance or decrease seed germination of invasive alien species through pulp removal or seed coat abrasion, besides serving as dispersal agents. Fruits of four fleshy-fruited invasive alien plant species, namely Solanum mauritianum, Cinnamomum camphora, Lantana camara and Psidium guajava, were fed to three generalist avian frugivorous species, which have been observed feeding on these fruits in the wild. Seed retention time was recorded as this affects dispersal distance and the duration that seeds are exposed to the effects of the gut. Seeds removed from excreta, seeds from manually de-pulped fruit, and whole fruit were planted in soil trays housed in a greenhouse. Daily germination counts were done. Seed retention times differed significantly between bird species for all fruits, except those of C. camphora. However, all frugivores had a similar effect on the germination success of seeds of S. mauritianum, L. camara and P. guajava, showing that gut retention time was not important. Germination of seeds from manually de-pulped fruits did not differ from that of ingested seeds of all plant species, suggesting that seed coat abrasion was also not important. Pulp removal resulted in significantly higher germination rates, both in the two species with larger, multi-seeded fruit (S. mauritianum and P. guajava), and in the two species having single-seeded fruit with waxy exocarps (C. camphora and L. camara). Pulp removal also resulted in significantly earlier germination of L. camara and P. guajava seeds. Therefore, frugivores not only accelerate dispersal, but also greatly enhance seed germination of all fleshy-fruited invasive alien species in this study.     

Landscape‐level tree cover predicts species richness of large‐bodied frugivorous birds in forest fragments

Large‐bodied frugivorous birds play an important role in dispersing large‐sized seeds in Neotropical rain forests, thereby maintaining tree species richness and diversity. Conversion of contiguous forest land to forest fragments is thought to be driving population declines in large‐bodied frugivores, but the mechanistic drivers of this decline remain poorly understood. To assess the importance of fragment‐level versus local landscape attributes in influencing the species richness of large‐bodied (>100 g) frugivorous birds, we surveyed 15 focal species in 22 forest fragments (2.7 to 33.6 ha, avg. = 16.0 ha) in northwest Ecuador in 2014. Fragment habitat variables included density of large trees, canopy openness and height, and fragment size; landscape variables included elevation and the proportion of tree cover within a 1 km radius of each fragment. At both the individual species level, and across the community of 12 species of avian frugivore we detected, there was higher richness and probability of presence in fragments with more tree cover on surrounding land. This tendency was particularly pronounced among some endangered species. These findings corroborate the idea that partially forested land surrounding fragments may effectively increase the suitable habitat for forest‐dwelling frugivorous birds in fragmented landscapes. These results can help guide conservation priorities within fragmented landscapes, with particular reference to retaining trees and reforesting to attain high levels of tree cover in areas between forest patches.     

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.     

Forest edges and fire ants alter the seed shadow of an ant-dispersed plant

Exotic species invade fragmented, edge-rich habitats readily, yet the distinct impacts of habitat edges and invaders on native biota are rarely distinguished. Both appear detrimental to ant-dispersed plants such as bloodroot, Sanguinaria canadensis. Working in northeastern Georgia (USA), an area characterized by a rich ant-dispersed flora, fragmented forests, and invasions by the red imported fire ant, Solenopsis invicta , I monitored the interactions between ants and S. canadensis seeds in uninvaded forest interiors, uninvaded forest edges, invaded forest interiors, and invaded forest edges. I observed 95% of the seed dispersal events that occurred within the 60-min observation intervals. Seed collection rates were similar among all four (habitat × invasion) groups. The presence of invasive ants had a strong effect on seed dispersal distance: S. invicta collected most seeds in invaded sites, but was a poorer disperser than four of five native ant taxa. Habitat type (interior versus edge) had no effect on seed dispersal distance, but it had a strong effect on seed dispersal direction. Dispersal towards the edge was disproportionately rare in uninvaded forest edges, and ants in those habitats moved the average dispersed seed approximately 70 cm away from that edge. Dispersal direction was also skewed away from the edge in uninvaded forest interiors and invaded forest edges, albeit non-significantly. This biased dispersal may help explain the rarity of myrmecochorous plants in younger forests and edges, and their poor ability to disperse between fragments. This is the first demonstration that forest edges and S. invicta invasion influence seed dispersal destination and distance, respectively. These forces act independently.