Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

Directed seed dispersal towards areas with low conspecific tree density by a scatter‐hoarding rodent

Scatter‐hoarding animals spread out cached seeds to reduce density‐dependent theft of their food reserves. This behaviour could lead to directed dispersal into areas with lower densities of conspecific trees, where seed and seedling survival are higher, and could profoundly affect the spatial structure of plant communities. We tested this hypothesis with Central American agoutis and Astrocaryum standleyanum palm seeds on Barro Colorado Island, Panama. We radio‐tracked seeds as they were cached and re‐cached by agoutis, calculated the density of adult Astrocaryum trees surrounding each cache, and tested whether the observed number of trees around seed caches declined more than expected under random dispersal. Seedling establishment success was negatively dependent on seed density, and agoutis carried seeds towards locations with lower conspecific tree densities, thus facilitating the escape of seeds from natural enemies. This behaviour may be a widespread mechanism leading to highly effective seed dispersal by scatter‐hoarding animals.     

Cooperation‐mediated plasticity in dispersal and colonization

Kin selection theory predicts that costly cooperative behaviors evolve most readily when directed toward kin. Dispersal plays a controversial role in the evolution of cooperation: dispersal decreases local population relatedness and thus opposes the evolution of cooperation, but limited dispersal increases kin competition and can negate the benefits of cooperation. Theoretical work has suggested that plasticity of dispersal, where individuals can adjust their dispersal decisions according to the social context, might help resolve this paradox and promote the evolution of cooperation. Here, we experimentally tested the hypothesis that conditional dispersal decisions are mediated by a cooperative strategy: we quantified the density‐dependent dispersal decisions and subsequent colonization efficiency from single cells or groups of cells among six genetic strains of the unicellular Tetrahymena thermophila that differ in their aggregation level (high, medium, and low), a behavior associated with cooperation strategy. We found that the plastic reaction norms of dispersal rate relative to density differed according to aggregation level: highly aggregative genotypes showed negative density‐dependent dispersal, whereas low‐aggregation genotypes showed maximum dispersal rates at intermediate density, and medium‐aggregation genotypes showed density‐independent dispersal with intermediate dispersal rate. Dispersers from highly aggregative genotypes had specialized long‐distance dispersal phenotypes, contrary to low‐aggregation genotypes; medium‐aggregation genotypes showing intermediate dispersal phenotype. Moreover, highly aggregation genotypes showed evidence for beneficial kin‐cooperation during dispersal. Our experimental results should help to resolve the evolutionary conflict between cooperation and dispersal: cooperative individuals are expected to avoid kin‐competition by dispersing long distances, but maintain the benefits of cooperation by dispersing in small groups.     

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.     

Short‐term variations in gene flow related to cyclic density fluctuations in the common vole

In highly fluctuating populations with complex social systems, genetic patterns are likely to vary in space and time due to demographic and behavioural processes. Cyclic rodents are extreme examples of demographically instable populations that often exhibit strong social organization. In such populations, kin structure and spacing behaviour may vary with density fluctuations and impact both the composition and spatial structure of genetic diversity. In this study, we analysed the multiannual genetic structure of a cyclic rodent, Microtus arvalis, using a sample of 875 individuals trapped over three complete cycles (from 1999 to 2007) and genotyped at 10 microsatellite loci. We tested the predictions that genetic diversity and gene flow intensity vary with density fluctuations. We found evidences for both spatial scale‐dependant variations in genetic diversity and higher gene flow during high density. Moreover, investigation of sex‐specific relatedness patterns revealed that, although dispersal is biased toward males in this species, distances moved by both sexes were lengthened during high density. Altogether, these results suggest that an increase in migration with density allows to restore the local loss of genetic diversity occurring during low density. We then postulate that this change in migration results from local competition, which enhances female colonization of empty spaces and male dispersal among colonies.     

Density‐dependent fitness,not dispersal movements,drives temporal variation in spatial genetic structure in dark‐eyed juncos (Junco hyemalis)

Some studies have found that dispersal rates and distances increase with density, indicating that density‐dependent dispersal likely affects spatial genetic structure. In an 11‐year mark–recapture study on a passerine, the dark‐eyed junco, we tested whether density affected dispersal distance and/or fine‐scale spatial genetic structure. Contrary to expectations, we found no effect of predispersal density on dispersal distance or the proportion of locally produced juveniles returning to the population from which they hatched. However, even though density did not affect dispersal distance or natal return rates, we found that density still did affect spatial genetic structure. We found significant positive spatial genetic structure at low densities of (postdispersal) adults but not at high densities. In years with high postdispersal (adult) densities that also had high predispersal (juvenile) densities in the previous year, we found negative spatial genetic structure, indicating high levels of dispersal. We found that density also affected fitness of recruits, and fitness of immigrants, potentially linking these population parameters with the spatial genetic structure detected. Immigrants and recruits rarely nested in low postdispersal density years. In contrast, in years with high postdispersal density, recruits were common and immigrants had equal success to local birds, so novel genotypes diluted the gene pool and effectively eliminated positive spatial genetic structure. In relation to fine‐scale spatial genetic structure, fitness of immigrants and new recruits is poorly understood compared to dispersal movements, but we conclude that it can have implications for the spatial distribution of genotypes in populations.     

Density‐dependent pre‐dispersal seed predation and fruit set in a tropical tree

Negative density‐dependent demographic processes operating at post‐dispersal seed, seedling, and juvenile stages are the dominant explanation for the coexistence of high numbers of tree species in tropical forests. At adult stages, the effect of pollinators and pre‐dispersal fruit predators are often dependent on the density or abundance of flowers and fruit in the canopy, but each have opposite effects on individual realized reproduction. We studied the effect of density on total and mature fruit set and pre‐dispersal predation rates within individual tree canopies in a common canopy tree species, Jacaranda copaia in a 50‐ha forest census plot in central Panama. We sampled all reproductive sized trees in the plot (n = 188) across three years and estimated fruit set and predation rates. Population‐wide pre‐dispersal seed predation averaged between 6–37% across years. Using linear mixed effects models, we found that increased density and fecundity of conspecific neighbours increased focal tree fruit set, but also the rate of pre‐dispersal predation. An interaction between individual and neighbourhood fruit production predicted lower predation rates at high individual and neighbourhood fecundities, which suggests predator satiation at high fruit abundance levels. However, the rate at which fruit set increased with conspecific neighbour fruit production was greater than the rate at which fruit were lost to predation, resulting in an overall positive effect of neighbour density on mature fruit production in focal trees. Our results run counter to the expectation of a uniformly negative effect of density across all life stages in tropical trees and suggest further exploration of the role of spatial clumping, pollen dispersal limitation, and predation at pre‐dispersal adult stages in maintenance of species diversity in plant communities.     

FINE‐SCALE GENETIC STRUCTURE IN A WILD BIRD POPULATION: THE ROLE OF LIMITED DISPERSAL AND ENVIRONMENTALLY BASED SELECTION AS CAUSAL FACTORS

Individuals are typically not randomly distributed in space; consequently ecological and evolutionary theory depends heavily on understanding the spatial structure of populations. The central challenge of landscape genetics is therefore to link spatial heterogeneity of environments to population genetic structure. Here, we employ multivariate spatial analyses to identify environmentally induced genetic structures in a single breeding population of 1174 great tits Parus major genotyped at 4701 single‐nucleotide polymorphism (SNP) loci. Despite the small spatial scale of the study relative to natal dispersal, we found multiple axes of genetic structure. We built distance‐based Moran's eigenvector maps to identify axes of pure spatial variation, which we used for spatial correction of regressions between SNPs and various external traits known to be related to fitness components (avian malaria infection risk, local density of conspecifics, oak tree density, and altitude). We found clear evidence of fine‐scale genetic structure, with 21, seven, and nine significant SNPs, respectively, associated with infection risk by two species of avian malaria (Plasmodium circumflexum and P. relictum) and local conspecific density. Such fine‐scale genetic structure relative to dispersal capabilities suggests ecological and evolutionary mechanisms maintain within‐population genetic diversity in this population with the potential to drive microevolutionary change.     

Conspecific density dependence and community structure: Insights from 11 years of monitoring in an old‐growth temperate forest in Northeast China

Forest community structure may be influenced by seedling density dependence, however, the effect is loosely coupled with population dynamics and diversity in the short term. In the long term the strength of conspecific density dependence may fluctuate over time because of seedling abundance, yet few long‐term studies exist. Based on 11 years of seedling census data and tree census data from a 25‐ha temperate forest plot in Northeast China, we used generalized linear mixed models to test the relative effects of local neighborhood density and abiotic factors on seedling density and seedling survival. Spatial point pattern analysis was used to determine if spatial patterns of saplings and juveniles, in relation to conspecific adults, were in accordance with patterns uncovered by conspecific negative density dependence at the seedling stage. Our long‐term results showed that seedling density was mainly positively affected by conspecific density, suggesting dispersal limitation of seedling development. The probability of seedling survival significantly decreased over 1 year with increasing conspecific density, indicating conspecific negative density dependence in seedling establishment. Although there was variation in conspecific negative density dependence at the seedling stage among species and across years, a dispersed pattern of conspecific saplings relative to conspecific adults at the local scale (<10 m) was observed in four of the 11 species examined. Overall, sapling spatial patterns were consistent with the impacts of conspecific density on seedling dynamics, which suggests that conspecific negative density dependence is persistent over the long term. From the long‐term perspective, conspecific density dependence is an important driver of species coexistence in temperate forests.     

Fine‐scale environmental control of hybridization in oaks

Natural hybridization is attracting much interest in modern speciation and conservation biology studies, but the underlying mechanisms remain poorly understood. In particular, it is unclear why environmental changes often increase hybridization rates. To study this question, we surveyed mating events in a mixed oak stand and developed a spatially explicit individual‐based hybridization model. This model, where hybridization is frequency‐dependent, pollen is nonlimiting and which allows immigrant pollen to compete with local pollen, takes into account species‐specific pollen dispersal and sexual barriers to hybridization. The consequences of pollen limitation on hybridization were studied using another simple model. The results indicate that environmental changes could increase hybridization rates through two distinct mechanisms. First, by disrupting the spatial organization of communities, they should decrease the proportion of conspecific pollen available for mating, thus increasing hybridization rates. Second, by decreasing the density of conspecifics, they should increase pollen limitation and thus hybridization rates, as a consequence of chance pollination predominating over deterministic pollen competition. Altogether, our results point to a need for considering hybridization events at the appropriate level of organization and provide new insights into why hybridization rates generally increase in disturbed environments.     

Active dispersal is differentially affected by inter‐ and intraspecific competition in closely related nematode species

Competition is one of the main drivers of dispersal, which can be an important mechanism to achieve permanent or temporal coexistence of multiple species. This coexistence can be achieved by a dispersal‐competition tradeoff, spatial store effects or neutral dynamics. Here we test the effect of inter‐ and intraspecific competition on dispersal of four species of the marine nematode species complex Litoditis marina. A previous study in closed microcosms without a possibility for dispersal had demonstrated pronounced interspecific competition, leading to the exclusion of one species. We now investigated whether 1) the dispersal is affected by interspecific interactions, by intraspecific competition (density) or by food availability, 2) the dispersal dynamics influence assemblage composition and can lead to co‐occurrence of the species, and 3) the abiotic environment (here salinity) can affect these dynamics. We show that density is the main driver for dispersal in two of the four species. Dispersal of a third species always started at the same time irrespective of density, whereas in the fourth species interspecific interactions accelerated dispersal. Remarkably, this fourth species was not a strong competitor, suggesting that a dispersal–competition tradeoff does not explain the observed coexistence. Salinity did not alter the timing of dispersal when interspecific interactions were present but did affect assemblage composition. Consequently, spatial store effects may influence coexistence. All four species co‐occurred in fairly stable abundances throughout the present experiment indicating the importance of species specific dispersal strategies for coexistence. Co‐occurrence can be facilitated because competition is postponed or avoided by dispersal. Neutral dynamics also played a role as intra‐ and interspecific competition were of similar importance in three of the four species. We conclude that dispersal is a driver of the coexistence of closely related nematode species, and that population density and interspecific interactions shape these dynamics.     

The influence of breeding phenology on the genetic structure of four pond‐breeding salamanders

Understanding metapopulation dynamics requires knowledge about local population dynamics and movement in both space and time. Most genetic metapopulation studies use one or two study species across the same landscape to infer population dynamics; however, using multiple co‐occurring species allows for testing of hypotheses related to different life history strategies. We used genetic data to study dispersal, as measured by gene flow, in three ambystomatid salamanders (Ambystoma annulatum , A. maculatum , and A. opacum ) and the Central Newt (Notophthalmus viridescens louisianensis ) on the same landscape in Missouri, USA . While all four salamander species are forest dependent organisms that require fishless ponds to reproduce, they differ in breeding phenology and spatial distribution on the landscape. We use these differences in life history and distribution to address the following questions: (1) Are there species‐level differences in the observed patterns of genetic diversity and genetic structure? and (2) Is dispersal influenced by landscape resistance? We detected two genetic clusters in A. annulatum and A. opacum on our landscape; both species breed in the fall and larvae overwinter in ponds. In contrast, no structure was evident in A. maculatum and N. v. louisianensis , species that breed during the spring. Tests for isolation by distance were significant for the three ambystomatids but not for N. v. louisianensis . Landscape resistance also contributed to genetic differentiation for all four species. Our results suggest species‐level differences in dispersal ability and breeding phenology are driving observed patterns of genetic differentiation. From an evolutionary standpoint, the observed differences in dispersal distances and genetic structure between fall breeding and spring breeding species may be a result of the trade‐off between larval period length and size at metamorphosis which in turn may influence the long‐term viability of the metapopulation. Thus, it is important to consider life history differences among closely related and ecologically similar species when making management decisions.     

Forest amount,not structure,influences fruit removal of two pioneer species in Atlantic forest remnants

Fruit removal is a key component of the seed dispersal process with direct consequences for plant recruitment. Anthropogenic disturbances might affect removal rates by changing frugivore diversity and their behavior. Here, we investigated the effects of local forest structure and landscape context on fruit removal of two common pioneer species of Melastomataceae family, Clidemia capitellata and Henriettea succosa in 14 Atlantic forest sites. We evaluated local forest structure, measured as canopy openness and conspecific density, and the amount of forest cover at the landscape scale measured in 2000 m buffer radius. Our results showed that the landscape context was more important than local features to explain fruit removal. The percentage of fruits removed from both species sharply decreases with forest loss and increases again in highly deforested landscapes. We demonstrate changes in fruit removal due to forest loss and suggest that both Melastomataceae species rely on the absence of other plant species, and consequently, in a less competitive environment to increase fruit removal and seed dispersal. Abstract in Portuguese is available with online material.     

Female‐biased dispersal in a bat with a female‐defence mating strategy

The ultimate causes for predominant male‐biased dispersal (MBD) in mammals and female‐biased dispersal (FBD) in birds are still subject to much debate. Studying exceptions to general patterns of dispersal, for example, FBD in mammals, provides a valuable opportunity to test the validity of proposed evolutionary pressures. We used long‐term behavioural and genetic data on individually banded Proboscis bats (Rhynchonycteris naso) to show that this species is one of the rare mammalian exceptions with FBD. Our results suggest that all females disperse from their natal colonies prior to first reproduction and that a substantial proportion of males are philopatric and reproduce in their natal colonies, although male immigration has also been detected. The age of females at first conception falls below the tenure of males, suggesting that females disperse to avoid father–daughter inbreeding. Male philopatry in this species is intriguing because Proboscis bats do not share the usual mammalian correlates (i.e. resource‐defence polygyny and/or kin cooperation) of male philopatry. They have a mating strategy based on female defence, where local mate competition between male kin is supposedly severe and should prevent the evolution of male philopatry. However, in contrast to immigrant males, philopatric males may profit from acquaintance with the natal foraging grounds and may be able to attain dominance easier and/or earlier in life. Our results on Proboscis bats lent additional support to the importance of inbreeding avoidance in shaping sex‐biased dispersal patterns and suggest that resource defence by males or kin cooperation cannot fully explain the evolution of male philopatry in mammals.     

Limited influence of local and landscape factors on finescale gene flow in two pond‐breeding amphibians

Dispersal and gene flow within animal populations are influenced by the composition and configuration of the landscape. In this study, we evaluated hypotheses about the impact of natural and anthropogenic factors on genetic differentiation in two amphibian species, the spotted salamander (Ambystoma maculatum) and the wood frog (Lithobates sylvaticus) in a commercial forest in central Maine. We conducted this analysis at two scales: a local level, focused on factors measured at each breeding pond, and a landscape level, focused on factors measured between ponds. We investigated the effects of a number of environmental factors in six categories including Productivity, Physical, Land Composition, Land Configuration, Isolation and Location. Embryos were sampled from 56 spotted salamander breeding ponds and 39 wood frog breeding ponds. We used a hierarchical Bayesian approach in the program GESTE at each breeding pond and a random forest algorithm in conjunction with a network analysis between the ponds. We found overall high genetic connectivity across distances up to 17 km for both species and a limited effect of natural and anthropogenic factors on gene flow. We found the null models best explained patterns of genetic differentiation at a local level and found several factors at the landscape level that weakly influenced gene flow. This research indicates multiscale investigations that incorporate local and landscape factors are valuable for understanding patterns of gene flow. Our findings suggest that dispersal rates in this system are high enough to minimize genetic structuring and that current forestry practices do not significantly impede dispersal.     

Environmental drivers of femaleness of an inter‐Andean monoecious shrub

Hetero‐and conspecific interactions, nutrient availability, climate, habitat heterogeneity, and disturbances can generate variation and spatial patterns of femaleness in plants. We assessed whether year, site, plant size, plant density, and canopy area of conspecific neighbors influenced the expression and spatial aggregation of femaleness in Croton aff. wagneri, a monoecious shrub from dry shrublands of the inter‐Andean valleys in Ecuador. We georeferenced in two sites (1,700 and 1,400 m.a.s.l) in five 10 × 10 m plots, within each site, the position of each Croton reproductive plant during first part of flowering season in two years, and measured their height, length, and width. The femaleness index of each plant was determined by the number of female and male buds and flowers. Plant density was determined for each plot, along with the number of neighbors and the summed canopy area of conspecific neighbors (at 1.0, 2.0, and 2.5 m radius, and the five closest plants) from each focal plant. Croton´s femaleness at the lower elevation site was greater than at the higher elevation site and increased with plant size and with canopy of the closest five neighbors. Soil at the lower elevation site had higher temperatures and lower water content. Aggregate patterns of femaleness were found in more plots at the lower elevation site. Our results indicate that location, plant size, and canopies of conspecific neighbors of Croton can affect femaleness and its aggregation and support the hypothesis that femaleness can be influenced by facilitative interactions. Abstract in Spanish is available with online material.     

Natal habitat effects drive density—dependent scaling of dispersal decisions

The dispersal behavior of a species is critical for the stability and persistence of its populations across a landscape. How population density affects dispersal decisions is important for predicting these dynamics, as the form of density‐dependent dispersal influences the stability and persistence of populations. Natal habitat experience often has strong impacts on individual dispersal behavior as well, but its influence on density‐dependent dispersal behaviors remains unexplored. Here we address this conceptual gap in two experiments separately examining habitat selection and emigration from recently colonized patches for two species of flour beetle Tribolium sp. We found that interactions between the quality of habitat experienced during natal development and current habitat for dispersal capable adults can strongly affect the form of density dependence, including reversing the direction of nonlinearities (accelerating to decelerating), or even negating the influence of population density for individual dispersal decisions. Across heterogeneous landscapes, where individuals from different populations may experience different natal habitats, this altering of density‐dependent relationships is predicted by theory to fundamentally influence regional population dynamics. Our results indicate that species which occur across heterogeneous environments, such as during conservation reintroductions, or as invasive species spread, have much potential for natal experience to interact with density dependence and influence local and regional population dynamics.     

Mating Behavior Drives Seed Dispersal by the Long‐wattled Umbrellabird Cephalopterus penduliger

Frugivores exhibit considerable variation in the seed dispersal services they provide. Understanding what drives these differences is a key goal for ecologists because of the central role seed dispersal plays in shaping ecological and genetic diversity in plant populations. The lek‐mating system of the Long‐wattled Umbrellabird (Cephalopterus penduliger) provides a powerful lens to examine how mating behavior may impact seed dispersal outcomes. As in all lek‐breeding species, male Umbrellabirds congregate in traditional sites (leks) to display, whereas females are solitary and visit leks only rarely. This study demonstrates how differences in mating behavior between the sexes drive distinctive seed movement and deposition patterns by male vs. female Umbrellabirds. Using radio tracking and gut retention trials, we documented divergent movement patterns between the sexes that are directly attributable to mating behavior differences. These movement differences led males to disperse seeds long distances from source trees and to deposit the majority of seeds they ingested within the lek; females dispersed seeds shorter distances and more evenly across the landscape. We empirically confirmed that the density of dispersed seeds was higher in leks than in control areas outside the lek, yet found no evidence that this higher density of seeds in leks reduced probability of seedling establishment. This research not only provides a mechanistic explanation for long dispersal distances and high levels of genetic diversity previously reported for seeds in Umbrellabird leks, but also highlights the importance of explicitly considering behavior in studies of animal‐mediated seed dispersal.     

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.     

Heterogeneity in local density allows a positive evolutionary relationship between self‐fertilisation and dispersal

Despite empirical evidence for a positive relationship between dispersal and self‐fertilization (selfing), theoretical work predicts that these traits should always be negatively correlated, and the Good Coloniser Syndrome of high dispersal and selfing (Cf. Baker's Law) should not evolve. Critically, previous work assumes that adult density is spatiotemporally homogeneous, so selfing results in identical offspring production for all patches, eliminating the benefit of dispersal for escaping from local resource competition. We investigate the joint evolution of dispersal and selfing in a demographically structured metapopulation model where local density is spatiotemporally heterogeneous due to extinction‐recolonization dynamics. Selfing alleviates outcrossing failure due to low local density (an Allee effect) while dispersal alleviates competition through dispersal of propagules from high‐ to low‐density patches. Because local density is spatiotemporally heterogeneous in our model, selfing does not eliminate heterogeneity in competition, so dispersal remains beneficial even under full selfing. Hence the Good Coloniser Syndrome is evolutionarily stable under a broad range of conditions, and both negative and positive relationships between dispersal and selfing are possible, depending on the environment. Our model thus accommodates positive empirical relationships between dispersal and selfing not predicted by previous theoretical work and provides additional explanations for negative relationships.