Ecological consequences of forest elephant declines for Afrotropical forests

Poaching is rapidly extirpating African forest elephants (Loxodonta cyclotis) from most of their historical range, leaving vast areas of elephant‐free tropical forest. Elephants are ecological engineers that create and maintain forest habitat; thus, their loss will have large consequences for the composition and structure of Afrotropical forests. Through a comprehensive literature review, we evaluated the roles of forest elephants in seed dispersal, nutrient recycling, and herbivory and physical damage to predict the cascading ecological effects of their population declines. Loss of seed dispersal by elephants will favor tree species dispersed abiotically and by smaller dispersal agents, and tree species composition will depend on the downstream effects of changes in elephant nutrient cycling and browsing. Loss of trampling and herbivory of seedlings and saplings will result in high tree density with release from browsing pressures. Diminished seed dispersal by elephants and high stem density are likely to reduce the recruitment of large trees and thus increase homogeneity of forest structure and decrease carbon stocks. The loss of ecological services by forest elephants likely means Central African forests will be more like Neotropical forests, from which megafauna were extirpated thousands of years ago. Without intervention, as much as 96% of Central African forests will have modified species composition and structure as elephants are compressed into remaining protected areas. Stopping elephant poaching is an urgent first step to mitigating these effects, but long‐term conservation will require land‐use planning that incorporates elephant habitat into forested landscapes that are being rapidly transformed by industrial agriculture and logging.     

Functional nonredundancy of elephants in a disturbed tropical forest

Conservation efforts are often motivated by the threat of global extinction. Yet if conservationists had more information suggesting that extirpation of individual species could lead to undesirable ecological effects, they might more frequently attempt to protect or restore such species across their ranges even if they were not globally endangered. Scientists have seldom measured or quantitatively predicted the functional consequences of species loss, even for large, extinction‐prone species that theory suggests should be functionally unique. We measured the contribution of Asian elephants (Elephas maximus) to the dispersal of 3 large‐fruited species in a disturbed tropical moist forest and predicted the extent to which alternative dispersers could compensate for elephants in their absence. We created an empirical probability model with data on frugivory and seed dispersal from Buxa Tiger Reserve, India. These data were used to estimate the proportion of seeds consumed by elephants and other frugivores that survive handling and density‐dependent processes (Janzen‐Connell effects and conspecific intradung competition) and germinate. Without compensation, the number of seeds dispersed and surviving density‐dependent effects decreased 26% (Artocarpus chaplasha), 42% (Careya arborea), and 72% (Dillenia indica) when elephants were absent from the ecosystem. Compensatory fruit removal by other animals substantially ameliorated these losses. For instance, reductions in successful dispersal of D. indica were as low as 23% when gaur (Bos gaurus) persisted, but median dispersal distance still declined from 30% (C. arborea) to 90% (A. chaplasha) without elephants. Our results support the theory that the largest animal species in an ecosystem have nonredundant ecological functionality and that their extirpation is likely to lead to the deterioration of ecosystem processes such as seed dispersal. This effect is likely accentuated by the overall defaunation of many tropical systems.     

Incorporating animal behavior into seed dispersal models: implications for seed shadows

Seed dispersal fundamentally influences plant population and community dynamics but is difficult to quantify directly. Consequently, models are frequently used to describe the seed shadow (the seed deposition pattern of a plant population). For vertebrate-dispersed plants, animal behavior is known to influence seed shadows but is poorly integrated in seed dispersal models. Here, we illustrate a modeling approach that incorporates animal behavior and develop a stochastic, spatially explicit simulation model that predicts the seed shadow for a primate-dispersed tree species (Virola calophylla, Myristicaceae) at the forest stand scale. The model was parameterized from field-collected data on fruit production and seed dispersal, behaviors and movement patterns of the key disperser, the spider monkey (Ateles paniscus), densities of dispersed and non-dispersed seeds, and direct estimates of seed dispersal distances. Our model demonstrated that the spatial scale of dispersal for this V. calophylla population was large, as spider monkeys routinely dispersed seeds >100 m, a commonly used threshold for long-distance dispersal. The simulated seed shadow was heterogeneous, with high spatial variance in seed density resulting largely from behaviors and movement patterns of spider monkeys that aggregated seeds (dispersal at their sleeping sites) and that scattered seeds (dispersal during diurnal foraging and resting). The single-distribution dispersal kernels frequently used to model dispersal substantially underestimated this variance and poorly fit the simulated seed-dispersal curve, primarily because of its multimodality, and a mixture distribution always fit the simulated dispersal curve better. Both seed shadow heterogeneity and dispersal curve multimodality arose directly from these different dispersal processes generated by spider monkeys. Compared to models that did not account for disperser behavior, our modeling approach improved prediction of the seed shadow of this V. calophylla population. An important function of seed dispersal models is to use the seed shadows they predict to estimate components of plant demography, particularly seedling population dynamics and distributions. Our model demonstrated that improved seed shadow prediction for animal-dispersed plants can be accomplished by incorporating spatially explicit information on disperser behavior and movements, using scales large enough to capture routine long-distance dispersal, and using dispersal kernels, such as mixture distributions, that account for spatially aggregated dispersal.     

Introduced Birds and the Fate of Hawaiian Rainforests

Abstract:  The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.     

Molecular genetic and behavioral analysis of social organization in the Asian elephant (Elephas maximus)

We report on the genetic evaluation and behavioral study of social organization in the Asian elephant (Elephas maximus). Although Asian elephants and African elephants (Loxodonta africana) were previously thought to have similar social organizations, our results demonstrate a substantial difference in the complexity and structure of Asian elephant social groupings from that described for African savanna elephants. Photographic cataloging of individuals, radio telemetry, and behavioral observations in Ruhuna National Park, Sri Lanka, enabled us to assign associated females and young to four groups with overlapping ranges. Genetic sampling of individuals from the four groups in Ruhuna National Park and three other groups in surrounding areas, conducted through PCR amplification and sequencing of mitochondrial DNA from dung, supported the matriarchal nature of female groups and the lack of inter-group transfer of females. Behaviorally and genetically, the identified social groups were best described as ”family groups”. We did not find any evidence for the existence of social groups of higher complexity than family groups. Received: 25 March 2000 / Received in revised form: 28 March 2000 / Accepted: 1 April 2000     

Seed dispersal of desert annuals

We quantified seed dispersal in a guild of Sonoran Desert winter desert annuals at a protected natural field site in Tucson, Arizona, USA. Seed production was suppressed under shrub canopies, in the open areas between shrubs, or both by applying an herbicide prior to seed set in large, randomly assigned removal plots (10-30 m diameter). Seedlings were censused along transects crossing the reproductive suppression borders shortly after germination. Dispersal kernels were estimated for Pectocarya recurvata and Schismus barbatus from the change in seedling densities with distance from these borders via inverse modeling. Estimated dispersal distances were short, with most seeds traveling less than a meter. The adhesive seeds of P. recurvata went farther than the small S. barbatus seeds, which have no obvious dispersal adaptation. Seeds dispersed farther downslope than upslope and farther when dispersing into open areas than when dispersing into shrubs. Dispersal distances were short relative to the pattern of spatial heterogeneity created by the shrub and open space mosaic. This suggests that dispersal could contribute to local population buildup, possibly facilitating species coexistence. Overall, these results support the hypothesis that escape in time via delayed germination is likely to be more important for desert annuals than escape in space.     

Costs and benefits of capsaicin-mediated control of gut retention in dispersers of wild chilies

A fundamental way in which animal-dispersed plants can influence the viability and distribution of dispersed seeds is through control of retention time in the guts of dispersers. Using two species of wild chilies and their dispersers, we examined how chemical and physical properties of fruits and seeds mediate this interaction. Capsicum chacoense is polymorphic for pungency, occurs in Bolivia, and is dispersed mostly by elaenias. Capsicum annuum is not polymorphic, occurs in Arizona (USA), and is dispersed mostly by thrashers. We first tested whether capsaicin, the substance responsible for the pungency of chilies, affects gut retention time of seeds in primary dispersers. Capsaicin slowed gut passage of seeds but did so in a manner that differed greatly between bird species because the constipative effects of capsaicin occurred only after an 80-minute time lag. Elaenias in Bolivia held only 6% of C. chacoense seeds for > 80 minutes, whereas thrashers in Arizona held 78% of C. annuum seeds for > 80 minutes. Next we examined the effects of retention time on seed viability and germination. Increased retention resulted in a greater proportion of seeds germinating in C. annuum, had no effects on non-pungent C. chacoense, and had negative effects on pungent C. chacoense. These divergent effects are explained by differences in seed coat morphology: seed coats of pungent C. chacoense are 10-12% thinner than those of the other two types of seeds. Thus, longer retention times damaged seeds with the thinnest seed coats. In C. annuum, seed viability remained high regardless of retention time, but germination increased with retention, suggesting a role for scarification. Thus, in C. annuum, fruit chemistry appears well matched with seed morphology and disperser physiology: capsaicin extends gut retention for most seeds, resulting in greater seed scarification and higher germination rates. Increased retention of pungent C. chacoense seeds is detrimental, but because the primary consumers have short retention times, capsaicin slows only a small proportion of seeds, minimizing negative effects. These results illustrate the importance of context in studies of fruit secondary metabolites. The same chemical can have different impacts on plant fitness depending on its morphological, physiological, and ecological context.     

Postfledging Dispersal of White-crowned Pigeons: Implications for Conservation of Deciduous Seasonal Forests in the Florida Keys

From 1988 to 1991, we studied the postfledging dispersal of 31 radio-tagged White-crowned Pigeons ( Columba leucocephala ) from three natal keys in Florida Bay. Immature birds dispersed from the natal keys at 26–45 days after batching, and most young dispersed more than 20 km during the first 10 days postdispersal. Dispersing birds flew either north to the Florida mainland or east to northeast to the mainline Florida Keys. On the mainland, immature birds fed nearly exclusively within Everglades National Park or an adjacent state wildlife management area. On the mainline keys, White-crowned Pigeons selectively used 5.01–20 ha forest fragments (p < 0.10) during the first 72 hours postdispersal. After this period, dispersing birds showed no preference among fragment size classes but used deciduous seasonal forests more frequently than suburban habitat(p < 0.10). The spatial pattern of dispersal on the mainline keys suggests that, during the first 72 hours postdispersal. White-crowned Pigeons are not able to reach northern Key Largo, where 69% of the deciduous seasonal forests are protected in state or federal ownership. Protection of large forest fragments, especially on southern Key Largo, should be a priority for maintaining populations of White-crowned Pigeons. These forests provide a series of "stepping stones" that enable dispersing immature White-crowned Pigeons to fly to more distant areas where habitat availability is less restricted. This species is threatened in Florida and may play an important role in maintaining plant species diversity in the seasonal deciduous forests of south Florida by dispersing seeds of at least 37 species of trees and shrubs. Protection of sufficient habitat to allow successful postfledging dispersal of this important seed disperser will also protect the ecosystem's biodiversity.     

Geochemical and mineralogical characteristics of elephant geophagic soils in Udawalawe National Park,Sri Lanka

Geophagy or deliberate ingestion of soils was observed among Asian elephants (Elephas maximus) in the Udawalwe National Park, Sri Lanka, for several years. The geochemical and mineralogical composition of the clayey soil layers which are purposefully selected and eaten by elephants in the park were studied, in order to identify the possible reasons for elephant geophagy. The concentrations of major and trace elements were determined by means of X-ray fluorescence spectrometry in 21 soil samples from eight geophagic sites and six soil samples collected from four non-geophagic sites. The mineralogical composition of selected soil samples was investigated using X-ray diffractometry (XRD). These geochemical analyses revealed that geophagic soils in the study areas are deeply weathered and that most of the elements are leached from the soil layers under extreme weathering conditions. The XRD data showed that the soils of the area consisted mainly quartz, feldspar, and the clay minerals kaolinite, Fe-rich illite, and smectite. Although no significant geochemical differences were identified between geophagic and non-geophagic soils, a clear difference was observed in their clay mineralogical content. Soils eaten by elephants are richer in kaolinite and illite than non-geophagic soils, which contain a higher amount of smectite. It is suggested that elephants in Udawalawe National Park ingest soils mainly not to supplement the mineral contents of their forage but to detoxify unpalatable compounds in their diet.     

Experimental manipulation of seed shadows of an Afrotropical tree determines drivers of recruitment

The loss of animals in tropical forests may alter seed dispersal patterns and reduce seedling recruitment of tree species, but direct experimental evidence is scarce. We manipulated dispersal patterns of Manilkara mabokeensis, a monkey-dispersed tree, to assess the extent to which spatial distributions of seeds drive seedling recruitment. Based on the natural seed shadow, we created seed distributions with seeds deposited under the canopy ("no dispersal"), with declining density from the tree ("natural dispersal"), and at uniform densities ("good dispersal"). These distributions mimicked dispersal patterns that could occur with the extirpation of monkeys, low levels of hunting, and high rates of seed dispersal. We monitored seedling emergence and survival for 18 months and recorded the number of leaves and damage to leaves. "Good dispersal" increased seedling survival by 26%, and "no dispersal" decreased survival by 78%, relative to "natural dispersal." Using a mixed-effects survival model, we decoupled the distance and density components of the seed shadow: seedling survival depended on the seed density, but not on the distance from the tree. Although community seedling diversity tended to decrease with longer dispersal distances, we found no conclusive evidence that patterns of seed dispersal influence the diversity of the seedling community. Local seed dispersal does affect seedling recruitment and survival, with better dispersal resulting in higher seedling recruitment; hence the loss of dispersal services that comes with the reduction or extirpation of seed dispersers will decrease regeneration of some tree species.     

Birds as Suppliers of Seed Dispersal in Temperate Ecosystems: Conservation Guidelines from Real‐World Landscapes

Abstract: Seed dispersal by animals is considered a pivotal ecosystem function that drives plant‐community dynamics in natural habitats and vegetation recovery in human‐altered landscapes. Nevertheless, there is a lack of suitable ecological knowledge to develop basic conservation and management guidelines for this ecosystem service. Essential questions, such as how well the abundance of frugivorous animals predicts seeding function in different ecosystems and how anthropogenic landscape heterogeneity conditions the role of dispersers, remain poorly answered. In three temperate ecosystems, we studied seed dispersal by frugivorous birds in landscape mosaics shaped by human disturbance. By applying a standardized design across systems, we related the frequency of occurrence of bird‐dispersed seeds throughout the landscape to the abundance of birds, the habitat features, and the abundance of fleshy fruits. Abundance of frugivorous birds in itself predicted the occurrence of dispersed seeds throughout the landscape in all ecosystems studied. Even those landscape patches impoverished due to anthropogenic disturbance received some dispersed seeds when visited intensively by birds. Nonetheless, human‐caused landscape degradation largely affected seed‐deposition patterns by decreasing cover of woody vegetation or availability of fruit resources that attracted birds and promoted seed dispersal. The relative role of woody cover and fruit availability in seed dispersal by birds differed among ecosystems. Our results suggest that to manage seed dispersal for temperate ecosystem preservation or restoration one should consider abundance of frugivorous birds as a surrogate of landscape‐scale seed dispersal and an indicator of patch quality for the dispersal function; woody cover and fruit resource availability as key landscape features that drive seedfall patterns; and birds as mobile links that connect landscape patches of different degrees of degradation and habitat quality via seed deposition.     

Mechanistic understanding of human–wildlife conflict through a novel application of dynamic occupancy models

Crop and livestock depredation by wildlife is a primary driver of human–wildlife conflict, a problem that threatens the coexistence of people and wildlife globally. Understanding mechanisms that underlie depredation patterns holds the key to mitigating conflicts across time and space. However, most studies do not consider imperfect detection and reporting of conflicts, which may lead to incorrect inference regarding its spatiotemporal drivers. We applied dynamic occupancy models to elephant crop depredation data from India between 2005 and 2011 to estimate crop depredation occurrence and model its underlying dynamics as a function of spatiotemporal covariates while accounting for imperfect detection of conflicts. The probability of detecting conflicts was consistently <1.0 and was negatively influenced by distance to roads and elevation gradient, averaging 0.08–0.56 across primary periods (distinct agricultural seasons within each year). The probability of crop depredation occurrence ranged from 0.29 (SE 0.09) to 0.96 (SE 0.04). The probability that sites raided by elephants in primary period t would not be raided in primary period t + 1 varied with elevation gradient in different seasons and was influenced negatively by mean rainfall and village density and positively by distance to forests. Negative effects of rainfall variation and distance to forests best explained variation in the probability that sites not raided by elephants in primary period t would be raided in primary period t + 1. With our novel application of occupancy models, we teased apart the spatiotemporal drivers of conflicts from factors that influence how they are observed, thereby allowing more reliable inference on mechanisms underlying observed conflict patterns. We found that factors associated with increased crop accessibility and availability (e.g., distance to forests and rainfall patterns) were key drivers of elephant crop depredation dynamics. Such an understanding is essential for rigorous prediction of future conflicts, a critical requirement for effective conflict management in the context of increasing human–wildlife interactions.     

Noninvasive Stress and Reproductive Measures of Social and Ecological Pressures in Free-Ranging African Elephants

Abstract: The African elephant (   Loxodonta africana ) experienced a poaching-related 60% population decline between 1979 and 1988 that was inordinately concentrated on adults. This, coupled with political pressures to delist the elephant, has created a need for noninvasive physiological measures that can quantify the long-term effects of past mortality patterns of this long-lived species. We collected fresh fecal samples from 16 female elephants in three different groups over 23 months at Tarangire National Park, Tanzania, and analyzed them for fecal progesterone and cortisol metabolites. Social and ecological measures were collected concurrently. Fecal progesterone metabolite measures corresponded significantly with stage of gestation, and appear to be able to confirm pregnancy in female elephants from as early as 3 months of gestation. We found that progesterone metabolite concentrations were significantly lower during the dry season than during the wet season after controlling for stage of gestation. Fecal cortisol metabolite concentrations showed the opposite seasonal pattern, being significantly higher in the dry season and inversely correlated with rainfall across seasons. Fecal cortisol metabolite concentrations also increased with group size and were correlated positively with dominance rank in the largest group. Our results suggest that measures of progesterone and cortisol metabolites in feces provide indices of reproductive function and physiological stress that can quantify both natural and human disturbances in African elephants. These measures are ideally suited for monitoring the long-term effects of social disruption from poaching and a variety of other management concerns.     

The consequences of poaching and anthropogenic change for forest elephants

Poaching has devastated forest elephant populations (Loxodonta cyclotis), and their habitat is dramatically changing. The long‐term effects of poaching and other anthropogenic threats have been well studied in savannah elephants (Loxodonta africana), but the impacts of these changes for Central Africa's forest elephants have not been discussed. We examined potential repercussions of these threats and the related consequences for forest elephants in Central Africa by summarizing the lessons learned from savannah elephants and small forest elephant populations in West Africa. Forest elephant social organization is less known than the social organization of savannah elephants, but the close evolutionary history of these species suggests that they will respond to anthropogenic threats in broadly similar ways. The loss of older, experienced individuals in an elephant population disrupts ecological, social, and population parameters. Severe reduction of elephant abundance within Central Africa's forests can alter plant communities and ecosystem functions. Poaching, habitat alterations, and human population increase are probably compressing forest elephants into protected areas and increasing human–elephant conflict, which negatively affects their conservation. We encourage conservationists to look beyond documenting forest elephant population decline and address the causes of these declines when developing conversation strategies. We suggest assessing the effectiveness of the existing protected‐area networks for landscape connectivity in light of current industrial and infrastructure development. Longitudinal assessments of the effects of landscape changes on forest elephant sociality and behavior are also needed. Finally, lessons learned from West African elephant population loss and habitat fragmentation should be used to inform strategies for land‐use planning and managing human–elephant interactions.     

On the relative contributions of wind vs. animals to seed dispersal of four Sierra Nevada pines

Selective pressures that influence the form of seed dispersal syndromes are poorly understood. Morphology of plant propagules is often used to infer the means of dispersal, but morphology can be misleading. Several species of pines, for example, have winged seeds adapted for wind dispersal but owe much of their establishment to scatter-hoarding animals. Here the relative importance of wind vs. animal dispersal is assessed for four species of pines of the eastern Sierra Nevada that have winged seeds but differed in seed size: lodgepole pine (Pinus contorta murrayana, 8 mg); ponderosa pine (Pinus ponderosa ponderosa, 56 mg); Jeffrey pine (Pinus jeffreyi, 160 mg); and sugar pine (Pinus lambertiana, 231 mg). Pre-dispersal seed mortality eliminated much of the ponderosa pine seed crop (66%), but had much less effect on Jeffrey pine (32% of seeds destroyed), lodgepole pine (29%), and sugar pine (7%). When cones opened most filled seeds were dispersed by wind. Animals removed > 99% of wind-dispersed Jeffrey and sugar pine seeds from the ground within 60 days, but animals gathered only 93% of lodgepole pine seeds and 38% of ponderosa pine seeds during the same period. Animals gathered and scatter hoarded radioactively labeled ponderosa, Jeffrey, and sugar pine seeds, making a total of 2103 caches over three years of study. Only three lodgepole pine caches were found. Caches typically contained 1-4 seeds buried 5-20 mm deep, depths suitable for seedling emergence. Although Jeffrey and sugar pine seeds are initially wind dispersed, nearly all seedlings arise from animal caches. Lodgepole pine is almost exclusively wind dispersed, with animals acting as seed predators. Animals treated ponderosa pine in an intermediate fashion. Two-phased dispersal of large, winged pine seeds appears adaptive; initial wind dispersal helps to minimize pre-dispersal seed mortality whereas scatter hoarding by animals places seeds in sites with a higher probability of seedling establishment.     

Comparison of three chemical tests to assess seed viability: the seed dispersal system of the Macaronesian endemic plant <Emphasis Type="Italic">Rubia fruticosa</Emphasis> (Rubiaceae) as an example

Summary. Three chemical viability tests were evaluated in the seed dispersal system of Rubia fruticosa, in which three main groups of dispersers participate: reptiles, birds and mammals. Tetrazolium chloride (TTC) and indigo carmine (IC) indicated a lower viability of seeds from droppings of introduced rabbits (Oryctolagus cuniculus) than of those from control plants and the native dispersers, lizards and gulls. In the rabbit seed treatment, significant differences were observed between results obtained with TTC and IC tests. Interpretation of these data, due to the presence of doubtful embryo staining, was more difficult using the IC test. Furthermore, some seeds that were clearly dead had been underestimated. In contrast with results obtained from the two staining methods, the EC test did not confirm that viability of control seeds and those seeds consumed by native dispersers were clearly higher than in seeds ingested by O. cuniculus. Further, compared to the other two tests, the EC method requires more careful handling of the embryo during the extraction process to avoid errors in viability estimation, since this method measures concentration of electrolytes that are released through cellular membranes. Thus, TTC was the most reliable test to assess seed viability in the seed dispersal system of R. fruticosa, and these results agree with those obtained in previous germination experiments made on the same set of seeds given the same treatments.     

Disentangling the drivers of reduced long-distance seed dispersal by birds in an experimentally fragmented landscape

Seed dispersal is a crucial component of plant population dynamics. Human landscape modifications, such as habitat destruction and fragmentation, can alter the abundance of fruiting plants and animal dispersers, foraging rates, vector movement, and the composition of the disperser community, all of which can singly or in concert affect seed dispersal. Here, we quantify and tease apart the effects of landscape configuration, namely, fragmentation of primary forest and the composition of the surrounding forest matrix, on individual components of seed dispersal of Heliconia acuminata, an Amazonian understory herb. First we identified the effects of landscape configuration on the abundance of fruiting plants and six bird disperser species. Although highly variable in space and time, densities of fruiting plants were similar in continuous forest and fragments. However, the two largest-bodied avian dispersers were less common or absent in small fragments. Second, we determined whether fragmentation affected foraging rates. Fruit removal rates were similar and very high across the landscape, suggesting that Heliconia fruits are a key resource for small frugivores in this landscape. Third, we used radiotelemetry and statistical models to quantify how landscape configuration influences vector movement patterns. Bird dispersers flew farther and faster, and perched longer in primary relative to secondary forests. One species also altered its movement direction in response to habitat boundaries between primary and secondary forests. Finally, we parameterized a simulation model linking data on fruit density and disperser abundance and behavior with empirical estimates of seed retention times to generate seed dispersal patterns in two hypothetical landscapes. Despite clear changes in bird movement in response to landscape configuration, our simulations demonstrate that these differences had negligible effects on dispersal distances. However, small fragments had reduced densities of Turdus albicollis, the largest-bodied disperser and the only one to both regurgitate and defecate seeds. This change in Turdus abundance acted together with lower numbers of fruiting plants in small fragments to decrease the probability of long-distance dispersal events from small patches. These findings emphasize the importance of foraging style for seed dispersal and highlight the primacy of habitat size relative to spatial configuration in preserving biotic interactions.     

Nonredundancy in the dispersal network of a generalist tropical forest tree

Plant species with generalized dispersal mutualisms are considered to be robust to local frugivore extinctions because of redundancy between dispersal agents. However, real redundancy can only occur if frugivores have similar foraging and ranging patterns and if fruit is a limiting resource. We evaluated the quantitative and qualitative contributions of seed dispersers for an endochorus mast-fruiting species, Prunus javanica (Rosaceae) in Khao Yai National Park, Thailand, to evaluate the potential redundancy of dispersers. Data were collected from tree watches, seed/fruit traps, and seed transects under and away from fruiting trees, feeding and seed deposition by gibbons (Hylobates lar), and evaluations of seed and first-year seedling survival. We identified three clusters of dispersers within the network. Most (>80%) frugivore species observed were small birds and squirrels that were not functional dispersers, dropping most seeds under or very near the tree crown, where seedling survival was ultimately nil. Monkeys (Macaca leonina) were low-quality, short-range dispersers, but they dispersed large numbers of seeds and were responsible for 67% of surviving first-year seedlings. Gibbons and Oriental Pied Hornbills (Anthracoceros albirostris) handled few fruits, but they provided the highest quality service by carrying most seeds away from the canopy to medium and long distances, respectively. Although there was overlap in the deposition patterns of the functional dispersers, they displayed complementary, rather than redundant, roles in seed dispersal. Satiation of all functional dispersers further limited their capacity to "replace" one another. Redundancy must be evaluated at the community level because each type of disperser may shift to different species in the non-masting years of P. javanica. Our results underscore the need for research on broader spatial and temporal scales, which combines studies of dispersal and plant recruitment, to better understand mechanisms that maintain network stability.     

Genetic sub-structure and intermediate optimal outcrossing distance in the marine angiosperm <Emphasis Type="Italic">Zostera marina</Emphasis>

The spatial distribution of genetic variability depends on the spatial patterns of clonal and sexual reproduction, gene flow, genetic drift and natural selection. Species with restricted dispersal may exhibit genetic structuring within populations with immediate neighbours being close relatives, and may show differentiation among populations. Genetic structuring of a species may have important genetic, evolutionary and ecological consequences including distance-dependent mating success. In this study we used microsatellite markers to show that clones of Zostera marina in a population in the Ria Formosa, Portugal, were aggregated and covered distances of up to 3–4 m. Clones within 4 m of each other exhibited significant and positive coancestry values, reflecting the limited seed dispersal of this species. Hand-pollinations between near (0–10.9 m), intermediate (11–32 m) and far (15 km) individuals resulted in similar levels of seed set, although the near pollinations had higher, although not statistically significant, levels of seed abortion during maturation. Seeds from intermediate-distance pollinations had a significantly higher proportion of seeds germinate and shorter germination time than both the near and far seeds. Similarly, the average number of seedlings produced per pollination, used as an overall estimate of fitness, was significantly greater for the intermediate distance when compared to both near and far pollinations. These results suggest that the genetic structuring observed may result in both inbreeding and outbreeding depression, which gives rise to an intermediate optimal outcrossing distance.     

Meta‐Analysis of the Effects of Human Disturbance on Seed Dispersal by Animals

Animal‐mediated seed dispersal is important for sustaining biological diversity in forest ecosystems, particularly in the tropics. Forest fragmentation, hunting, and selective logging modify forests in myriad ways and their effects on animal‐mediated seed dispersal have been examined in many case studies. However, the overall effects of different types of human disturbance on animal‐mediated seed dispersal are still unknown. We identified 35 articles that provided 83 comparisons of animal‐mediated seed dispersal between disturbed and undisturbed forests; all comparisons except one were conducted in tropical or subtropical ecosystems. We assessed the effects of forest fragmentation, hunting, and selective logging on seed dispersal of fleshy‐fruited tree species. We carried out a meta‐analysis to test whether forest fragmentation, hunting, and selective logging affected 3 components of animal‐mediated seed dispersal: frugivore visitation rate, number of seeds removed, and distance of seed dispersal. Forest fragmentation, hunting, and selective logging did not affect visitation rate and were marginally associated with a reduction in seed‐dispersal distance. Hunting and selective logging, but not fragmentation, were associated with a large reduction in the number of seeds removed. Fewer seeds of large‐seeded than of small‐seeded tree species were removed in hunted or selectively logged forests. A plausible explanation for the consistently negative effects of hunting and selective logging on large‐seeded plant species is that large frugivores, as the predominant seed dispersers for large‐seeded plant species, are the first animals to be extirpated from hunted or logged forests. The reduction in forest area after fragmentation appeared to have weaker effects on frugivore communities and animal‐mediated seed dispersal than hunting and selective logging. The differential effects of hunting and selective logging on large‐ and small‐seeded tree species underpinned case studies that showed disrupted plant‐frugivore interactions could trigger a homogenization of seed traits in tree communities in hunted or logged tropical forests. Meta Análisis de los Efectos de la Perturbación Humana sobre la Dispersión de Semillas por Animales