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Seed dispersal is key to the persistence and spread of plant populations. Because the majority of plant species rely on animals to disperse their seeds, global change drivers that directly affect animals can cause cascading impacts on plant communities. In this review, we synthesize studies assessing how disperser loss alters plant populations, community patterns, multitrophic interactions, and ecosystem functioning. We argue that the magnitude of risk to plants from disperser loss is shaped by the combination of a plant species’ inherent dependence on seed dispersal and the severity of the hazards faced by their dispersers. Because the factors determining a plant species’ risk of decline due to disperser loss can be related to traits of the plants and dispersers, our framework enables a trait-based understanding of change in plant community composition and ecosystem functioning. We discuss how interactions among plants, among dispersers, and across other trophic levels also mediate plant community responses, and we identify areas for future research to understand and mitigate the consequences of disperser loss on plants globally., This work was supported by the National Socio-Environmental Synthesis Center’s Postdoctoral Fellowship to E.C.F. under funding received from the National Science Foundation (DBI1639145), the Alexander von Humboldt Foundation Fellowship to I.D., and the Spanish Ministry of Science to A.T. (project CGL2017-88122-P).

Food webs influence ecosystem diversity and functioning. Contemporary defaunation has reduced food web complexity, but simplification caused by past defaunation is difficult to reconstruct given the sparse paleorecord of predator-prey interactions. We identified changes to terrestrial mammal food webs globally over the past ~130,000 years using extinct and extant mammal traits, geographic ranges, observed predator-prey interactions, and deep learning models. Food webs underwent steep regional declines in complexity through loss of food web links after the arrival and expansion of human populations. We estimate that defaunation has caused a 53% decline in food web links globally. Although extinctions explain much of this effect, range losses for extant species degraded food webs to a similar extent, highlighting the potential for food web restoration via extant species recovery.

Many plants rely on animal mutualists for reproduction. Quantifying how animal mutualists impact plant performance provides a foundation for modelling how change in animal communities affects the composition and functioning of plant communities. We performed a meta-analysis of 2539 experiments, 6 times more than the last comprehensive meta-analysis, examining how gut passage by frugivores influences seed germination. We simultaneously analyzed multiple predictor variables related to study methodology, location, and frugivore identity to disentangle methodological from ecological impacts on effect sizes. We found that gut passage by birds, fish, reptiles, bats, primates, and other mammals on average increased seed germination, but that the magnitude varied across vertebrate groups. The positive effects of gut passage were largely explained by the de-inhibitory effects of pulp removal rather than by the scarification of seed tissues. Some previous studies and meta-analyses that found no effect of gut passage only tested scarification or did not distinguish between these tests of scarification and pulp removal. We found that, for a typical fleshy-fruited plant species, the lack of gut passage reduces germination by 60%. From an evolutionary perspective, this indicates a large risk associated with reliance on animal mutualists that is balanced against the benefits of animal-mediated seed dispersal. From a conservation perspective, this highlights the potential for large demographic consequences of frugivore declines on plant populations. Our database and findings advance quantitative predictions for the role of fruit-frugivore interactions in shaping plant communities in the Anthropocene.

Seed dispersal is an important ecological process that structures plant communities and influences ecosystem functioning. Loss of animal dispersers therefore poses a serious threat to forest ecosystems, particularly in the tropics where zoochory predominates. A prominent example is the near-total extinction of seed dispersers on the tropical island of Guam following the accidental introduction of the invasive brown tree snake (Boiga irregularis), negatively impacting seedling recruitment and forest regeneration. We investigated frugivory by a remnant population of Såli (Micronesian starling –Aplonis opaca) on Guam and two other island populations (Rota, Saipan) to evaluate their ecological role as a seed disperser in the Mariana archipelago. Using a combination of behavioural observations, nest contents and fecal samples, we documented frugivory of 37 plant species. Native plants comprised the majority (66%) of all species and 90% of all seeds identified in fecal and nest contents. Diet was highly similar across age classes and sampling years. In addition, plant species consumed by Såli comprised 88% of bird-dispersed adult trees and 54% of all adult trees in long-term forest monitoring plots, demonstrating the Såli’s broad diet and potential for restoring native forests. Overall, we provide the most comprehensive assessment to date of frugivory by the Såli and confirm its importance as a seed disperser on Guam and throughout the Marianas.

Substantial intra‐specific trait variation exists within plant communities, and in theory this variation could influence community dynamics. Although recent research has focused on intra‐specific variation in traits themselves, it is the influence of this variation on plant performance that makes intra‐specific trait variation relevant to ecological dynamics within or among species. Understanding the links between trait and performance variation, and the role of traits in mediating relationships among multiple components of performance, is critical for assessing the importance of intra‐specific trait variation for community dynamics. Seed size is thought to affect aspects of plant performance including fecundity, seedling growth, dispersal and tolerance of natural enemies. For two tropical tree species, we assessed how seed size was related to performance variation within each species and determined whether intra‐specific trait variation mediates intra‐specific performance tradeoffs. We used field seed rain collection to characterize size‐dependent outcomes of dispersal, sowed seeds of known size in soil collected near or far from conspecifics to characterize susceptibility to soil pathogens, and monitored growth of seedlings from seeds of known size. We found that intra‐specific seed size variation caused intra‐specific performance variation. The degree of trait‐based performance variation was consistently smaller than the degree of trait variation, and seed size influenced different components of performance for each species. One species exhibited a tradeoff in which small seeds had a fecundity advantage (more seedlings per unit reproductive mass) but produced smaller seedlings, whereas the other species exhibited a tradeoff in which small seeds dispersed to areas of low conspecific density but were less tolerant of density‐responsive natural enemies. Our results indicate that a single trait can influence multiple components of performance and can mediate different tradeoffs in co‐occurring species. Complex and heterogeneous effects of a single trait in multidimensional niche space may favour inter‐specific niche differentiation and coexistence.

Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. Generalizing and predicting dispersal processes are challenging due to context dependence of seed dispersal, environmental heterogeneity and interdependent processes occurring over multiple spatial and temporal scales. Current population models often use simple phenomenological descriptions of dispersal processes, limiting their ability to examine the role of population persistence and spread, especially under global change. To move seed dispersal ecology forward, we need to evaluate the impact of any single seed dispersal event within the full spatial and temporal context of a plant’s life history and environmental variability that ultimately influences a population’s ability to persist and spread. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. We synthesize suitable theoretical frameworks for this work and discuss concepts, approaches and available data from diverse subdisciplines to help operationalize concepts, highlight recent breakthroughs across research areas and discuss ongoing challenges and open questions. We address knowledge gaps in the movement ecology of seeds and the integration of dispersal and demography that could benefit from such a synthesis. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity., Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity.

Current understanding of mutualistic networks is grounded largely in data on interaction frequency, yet mutualistic network dynamics are also shaped by interaction quality—the functional outcomes of individual interactions on reproduction and survival. The difficulty of obtaining data on functional outcomes has resulted in limited understanding of functional variation among a network's pairwise species interactions, of the study designs that are necessary to capture major sources of functional variation, and of predictors of functional variation that may allow generalization across networks. In this community‐scale study, we targeted a key functional outcome in plant–frugivore networks: the impact of frugivore gut passage on seed germination. We used captive frugivore feeding trials and germination experiments in an island ecosystem, attaining species‐level coverage across all extant native frugivores and the plants they consume to (a) assess sources of functional variation, (b) separate effects of pulp removal from those of scarification via gut passage, and (c) test trait‐based correlates of gut passage effect sizes. We found antagonistic seed predation effects of a frugivore previously assumed to be a seed disperser, highlighting the need to consider functional outcomes rather than interaction frequency alone. The other frugivores each exhibited similar impacts for individual plant species, with benefits primarily caused by pulp removal rather than scarification, supporting the use of animal functional groups in this context. In contrast, plant species varied widely in impacts of gut passage on germination. Species with smaller seeds and more frugivore partners had larger benefits of gut passage, showing promise for network metrics and functional traits to predict functional variation among plants. Synthesis. Combining network and demographic approaches, we assessed the degree and sources of variation in a key functional outcome of plant–frugivore interactions across an entire network. Using a detailed study design, our work shows how simpler study designs can capture primary sources of functional variation and that functional traits and network metrics may allow generalization across networks. Efficiently measuring and generalizing sources of functional variation within mutualistic networks will strengthen our ability to model network dynamics and predict mutualist responses to global change.

Following defaunation, the loss of interactions with mutualists such as pollinators or seed dispersers may be compensated through increased interactions with remaining mutualists, ameliorating the negative cascading impacts on biodiversity. Alternatively, remaining mutualists may respond to altered competition by reducing the breadth or intensity of their interactions, exacerbating negative impacts on biodiversity. Despite the importance of these responses for our understanding of the dynamics of mutualistic networks and their response to global change, the mechanism and magnitude of interaction compensation within real mutualistic networks remains largely unknown. We examined differences in mutualistic interactions between frugivores and fruiting plants in two island ecosystems possessing an intact or disrupted seed dispersal network. We determined how changes in the abundance and behavior of remaining seed dispersers either increased mutualistic interactions (contributing to “interaction compensation”) or decreased interactions (causing an “interaction deficit”) in the disrupted network. We found a “rich-get-richer” response in the disrupted network, where remaining frugivores favored the plant species with highest interaction frequency, a dynamic that worsened the interaction deficit among plant species with low interaction frequency. Only one of five plant species experienced compensation and the other four had significant interaction deficits, with interaction frequencies 56-95% lower in the disrupted network. These results do not provide support for the strong compensating mechanisms assumed in theoretical network models, suggesting that existing network models underestimate the prevalence of cascading mutualism disruption after defaunation. This work supports a mutualist biodiversity-ecosystem functioning relationship, highlighting the importance of mutualist diversity for sustaining diverse and resilient ecosystems. This article is protected by copyright. All rights reserved.

1. Polyphenols are one of the most common groups of secondary metabolites in plants and thought to play a key role in enhancing plant fitness by protecting plants against enemies. Although enemy-inflicted mortality at the seed stage can be an important regulator of plant populations and a key determinant of community structure, few studies have assessed community-level patterns of polyphenol content in seeds. 2. We describe the distribution of the main seed polyphenol groups across 196 tree and liana species on Barro Colorado Island (Panama) and community-level patterns in two aspects of their biological activity (protein precipitation and oxidative capacity). Taking advantage of substantial variation in morphological and ecological traits in the studied plant community, we test for correlations and trade-offs between seed polyphenols and non-chemical plant traits hypothesised to make plant species more or less likely to invest in polyphenol production. 3. The majority of species have polyphenols in their seeds. The incidence and concentrations of polyphenols were related to a set of non-chemical plant traits. Polyphenols were most likely to be present (and where present, to be expressed in high concentrations) in species with large seeds, short seed dormancy times, low investment in mechanical seed defences, high wood density, high leaf mass per area, tough leaves, and slow growth rates. 4. Synthesis: Our study reveals a potential trade-off between chemical and mechanical seed defences and shows that plant species that invest in physical defences at later life stages (high wood density and tough leaves) tend not to invest in physical defences of seeds but instead produce secondary metabolites likely to act as seed defences. Overall, our results conform to predictions from the resource availability hypothesis, which states that species in resource-limited environments (such as slow-growing shade-tolerant tree species) will invest more in defences than fast-growing pioneer species.

Introductions of species by humans are causing the homogenization of species composition across biogeographic barriers1–3. The ecological and evolutionary consequences of introduced species derive from their effects on networks of species interactions4,5, but we lack a quantitative understanding of the impacts of introduced species on ecological networks and their biogeographic patterns globally. Here we address this data gap by analysing mutualistic seed-dispersal interactions from 410 local networks, encompassing 24,455 unique pairwise interactions between 1,631 animal and 3,208 plant species. We show that species introductions reduce biogeographic compartmentalization of the global meta-network, in which nodes are species and links are interactions observed within any local network. This homogenizing effect extends across spatial scales, decreasing beta diversity among local networks and modularity within networks. The prevalence of introduced interactions is directly related to human environmental modifications and is accelerating, having increased sevenfold over the past 75 years. These dynamics alter the coevolutionary environments that mutualists experience6, and we find that introduced species disproportionately interact with other introduced species. These processes are likely to amplify biotic homogenization in future ecosystems7 and may reduce the resilience of ecosystems by allowing perturbations to propagate more quickly and exposing disparate ecosystems to similar drivers. Our results highlight the importance of managing the increasing homogenization of ecological complexity. A quantitative analysis of the impact of species introductions on mutualistic seed-dispersal networks indicates that introduced species are increasingly erasing natural patterns of network biodiversity.

There is growing realization that intraspecific variation in seed dispersal can have important ecological and evolutionary consequences. However, we do not have a good understanding of the drivers or causes of intraspecific variation in dispersal, how strong an effect these drivers have, and how widespread they are across dispersal modes. As a first step to developing a better understanding, we present a broad, but not exhaustive, review of what is known about the drivers of intraspecific variation in seed dispersal, and what remains uncertain. We start by decomposing ‘drivers of intraspecific variation in seed dispersal’ into intrinsic drivers (i.e. variation in traits of individual plants) and extrinsic drivers (i.e. variation in ecological context). For intrinsic traits, we further decompose intraspecific variation into variation among individuals and variation of trait values within individuals. We then review our understanding of the major intrinsic and extrinsic drivers of intraspecific variation in seed dispersal, with an emphasis on variation among individuals. Crop size is the best-supported and best-understood intrinsic driver of variation across dispersal modes; overall, more seeds are dispersed as more seeds are produced, even in cases where per seed dispersal rates decline. Fruit/seed size is the second most widely studied intrinsic driver, and is also relevant to a broad range of seed dispersal modes. Remaining intrinsic drivers are poorly understood, and range from effects that are probably widespread, such as plant height, to drivers that are most likely sporadic, such as fruit or seed colour polymorphism. Primary extrinsic drivers of variation in seed dispersal include local environmental conditions and habitat structure. Finally, we present a selection of outstanding questions as a starting point to advance our understanding of individual variation in seed dispersal., Intraspecific variation in the quantity and quality of seed dispersal has important ecological and evolutionary consequences, yet it is generally ignored in favour of simply using population mean values. In a broad but not exhaustive review we show that drivers of intraspecific variation in seed dispersal are diverse and pervasive. These include both intrinsic (i.e. variation in traits of individual plants) and extrinsic (i.e. variation in ecological context) drivers. Further, these drivers appear to interact frequently in complex ways. Current theory does not include or account for these complex and interacting drivers.

How can hundreds of tree species coexist in a single hectare of tropical forest when the environmental conditions, as well as the species’ basic requirements, appear so similar? A leading explanation, particularly in tropical forests, is called the Janzen–Connell hypothesis after the two ecologists who proposed it in the early 1970s (1, 2). In this mechanism, specialized predators and pathogens concentrated near adult plants differentially reduce survival and growth of conspecific offspring relative to seedlings of other species. These natural enemies limit regeneration of common species and confer a relative advantage to rare species because they have more enemy-free regeneration sites. This has driven decades of empirical work to understand whether enemies are specialized enough in nature to cause this pattern and allow coexistence by this mechanism, and a recent metaanalysis supports the prediction that seed or seedling survival is greater away from conspecific adults (3). In PNAS, Eck et al. (4) show that natural enemies can be even more specialized, specializing on individual genotypes within a wild population. Through simulation models exploring the theoretical implications of this field result, Eck et al. (4) show that this greater specialization may weaken the stabilizing effects of natural enemies on species coexistence but also may select for greater dispersal over evolutionary time. The body of theory developed since the original formulation of the Janzen–Connell hypothesis has primarily asked ( i ) How specialized must the natural enemies be? and ( ii ) At what scale must these patterns operate? With regard to natural enemies, a long-standing conclusion is that species specificity of natural enemies allows coexistence. If a rare plant species shares a generalist enemy with an abundant plant species, then this abundant generalist enemy could extirpate the rare species from the community. With regard to patterns, the spatial scale matters—both in terms of enemy … [↵][1]1To whom correspondence should be addressed. Email: haldre{at}iastate.edu. [1]: #xref-corresp-1-1

Invasive vertebrate predators are directly responsible for the extinction or decline of many vertebrate species, but their indirect impacts often go unmeasured, potentially leading to an underestimation of their full impact. When invasives extirpate functionally important mutualists, dependent species are likely to be affected as well. Here, we show that the invasive brown treesnake, directly responsible for the extirpation of forest birds from the island of Guam, is also indirectly responsible for a severe decline in plant recruitment as a result of disrupting the fruit-frugivore mutualism. To assess the impact of frugivore loss on plants, we compare seed dispersal and recruitment of two fleshy-fruited tree species on Guam and three nearby islands with intact disperser communities. We conservatively estimate that the loss of frugivorous birds caused by the brown treesnake may have caused a 61–92% decline in seedling recruitment. This case study highlights the potential for predator invasions to cause indirect, pervasive and easily overlooked interaction cascades., Invasive brown treesnakes decimated the forest bird community on the island of Guam. Now, Rogers and colleagues document the indirect effects of the snake on trees, linking snake-initiated bird loss to reduced seed dispersal and plant recruitment on Guam compared to nearby uninvaded islands.

Seed size and toughness affect seed predators, and size-dependent investment in mechanical defence could affect relationships between seed size and predation. We tested how seed toughness and mechanical defence traits (tissue density and protective tissue content) are related to seed size among tropical forest species. Absolute toughness increased with seed size. However, smaller seeds had higher specific toughness both within and among species, with the smallest seeds requiring over 2000 times more energy per gram to break than the largest seeds. Investment in mechanical defence traits varied widely but independently of the toughness-mass allometry. Instead, a physical scaling relationship confers a toughness advantage on small seeds independent of selection on defence traits and without a direct cost. This scaling relationship may contribute to seed size diversity by decreasing fitness differences among large and small seeds. Allometric scaling of toughness reconciles predictions and conflicting empirical relationships between seed size and predation.

The Mariana islands have a species-poor but functionally diverse and largely endemic bird assemblage that varies due to biogeographic legacy and anthropogenic impacts. The largest island in the chain, Guam, is the setting for one of the most extreme examples of recent avian population declines, indicating the capacity for avifaunal collapse and loss of function in neighboring islands. We performed a systematic survey of resident land birds in remnant karst forest on the Mariana Islands' 3 largest islands following Guam to assess the status of the avifauna in this habitat, characterize inter-island heterogeneity in bird communities, and consider the resulting differences in the functional roles of birds across the archipelago's native forests. We identified significantly greater functional diversity on Rota than either Saipan or Tinian, but lower bird population densities, species richness, and Shannon diversity. We recommend continued monitoring of avian population trends across the archipelago and assessments of ecosystem functions like pollination, seed dispersal, and food web dynamics.

Seed dispersal enables plants to reach hospitable germination sites and escape natural enemies. Understanding when and how much seed dispersal matters to plant fitness is critical for understanding plant population and community dynamics. At the same time, the complexity of factors that determine if a seed will be successfully dispersed and subsequently develop into a reproductive plant is daunting. Quantifying all factors that may influence seed dispersal effectiveness for any potential seed-vector relationship would require an unrealistically large amount of time, materials and financial resources. On the other hand, being able to make dispersal predictions is critical for predicting whether single species and entire ecosystems will be resilient to global change. Building on current frameworks, we here posit that seed dispersal ecology should adopt plant functional groups as analytical units to reduce this complexity to manageable levels. Functional groups can be used to distinguish, for their constituent species, whether it matters (i) if seeds are dispersed, (ii) into what context they are dispersed and (iii) what vectors disperse them. To avoid overgeneralization, we propose that the utility of these functional groups may be assessed by generating predictions based on the groups and then testing those predictions against species-specific data. We suggest that data collection and analysis can then be guided by robust functional group definitions. Generalizing across similar species in this way could help us to better understand the population and community dynamics of plants and tackle the complexity of seed dispersal as well as its disruption., Seed dispersal is critical to plant fitness and plant community dynamics. However, measuring and tracking all factors that influence seed dispersal effectiveness for any potential seed-disperser relationship would require an unrealistically large amount of time, materials and financial resources. Building on current frameworks, we suggest that seed dispersal ecology quantify seed dispersal at the scale of plant functional groups, in order to reduce complexity to manageable levels. Based on functional group classifications, plant species can be distinguished by their level of dependence on seed dispersal and its mechanisms.

As the single opportunity for plants to move, seed dispersal has an important impact on plant fitness, species distributions and patterns of biodiversity. However, models that predict dynamics such as risk of extinction, range shifts and biodiversity loss tend to rely on the mean value of parameters and rarely incorporate realistic dispersal mechanisms. By focusing on the mean population value, variation among individuals or variability caused by complex spatial and temporal dynamics is ignored. This calls for increased efforts to understand individual variation in dispersal and integrate it more explicitly into population and community models involving dispersal. However, the sources, magnitude and outcomes of intraspecific variation in dispersal are poorly characterized, limiting our understanding of the role of dispersal in mediating the dynamics of communities and their response to global change. In this manuscript, we synthesize recent research that examines the sources of individual variation in dispersal and emphasize its implications for plant fitness, populations and communities. We argue that this intraspecific variation in seed dispersal does not simply add noise to systems, but, in fact, alters dispersal processes and patterns with consequences for demography, communities, evolution and response to anthropogenic changes. We conclude with recommendations for moving this field of research forward., Seed dispersal—the single opportunity for plants to move—is important for plant fitness, species distributions and patterns of biodiversity. Models that predict extinction risk of species, range shifts and biodiversity tend to rely on average dispersal distances. However, we know that seed dispersal is highly variable even within a single species (e.g. some seeds go very far and some barely move away from their parent plant, some seeds end up in great quality habitats and some end up on roads). This paper looks at the consequences of this variation in seed dispersal for plants and their ability to respond to future global change.

Patterns of nest site selection exhibited at the scale of a population should result from initial preferences of individuals occupying nest sites as well as preferences exhibited by individuals moving between nest sites. We tested whether nest-site preferences measured at the population scale were predictive of patterns of burrow switching by Leach's Storm-Petrels (Oceanodroma leucorhoa), a long-lived seabird that nests in underground burrows. Breeding pairs generally choose from the pool of available existing burrows rather than constructing new burrows, and a portion of the burrows in a colony remains unused in any breeding season. We quantified burrow preference at a colony on Kent Island, New Brunswick, over four breeding seasons. We used a classification and regression tree analysis to build a predictive model of nest-site selection. Preferentially occupied burrows were drier, longer, had larger nest chambers, and were in areas of higher burrow density. To measure preferences during burrow switching, we tracked individuals that switched burrows, comparing characteristics of the burrows in which these birds were originally found to those they inhabited at the end of the study period. Characteristics preferred by switching individuals were a subset of those observed at the scale of the population; individuals moved to burrows that were drier, longer, and had larger nest chambers. Our results show how preferences of individuals that move between nest sites contribute to nest site preferences exhibited at the population scales commonly tested. RESUMEN Modelo de la seleccion de cavidades permite predecir el patron de cambio de estas en Oceanodroma leucorhoa El patron de seleccion del lugar en donde anidar, exhibido a escala de una poblacion, debe ser el resultado de preferencias individuales de organismos que ocupan lugares para anidar, e igualmente de preferencias exhibidas por individuos que cambian de lugar para anidar. Pusimos apruebas si las preferencias, medidas a escala de poblacion, podian ser utilizadas para predecir el patron de cambio de cavidades de parte de Oceanodroma leucorhoa, que es un ave marina que anida en cavidades bajo tierra. Las parejas usualmente seleccionan entre un grupo de cavidades disponibles, en vez de estos construir la propia, y una porcion de las cavidades permanecen sin ser utilizadas a lo largo de la epoca de reproduccion. Cuantificamos la preferencia de uso de cavidades en una colonia localizada en la Isla Kent, New Brunswick, durante cuatro temporadas reproductivas. Utilizamos una clasificacion y un analisis de regresion para construir un modelo de prediccion para la seleccion de lugar de anidar. Los lugares ocupados, con preferencia, se mantenian secos, eran largos, de mayor tamano y se encontraban en lugares con alta densidad de cavidades. Para determinar las preferencias en el cambio de uso de cavidades, seguimos a individuos que cambiaron de cavidad, y compararnos las caracteristicas del lugar utilizado originalmente, con aquellas cavidades siendo utilizadas al final del estudio. Las caracteristica preferidas, por individuos que cambiaron de cavidad, eran un subconjunto de aquellas observadas a nivel poblacional. Las aves se movieron a cavidades mas secas, mas largas y con una camara de anidamiento mas amplia. Nuestros resultados muestran como las preferencias individuales, de aves que se mueven a otras cavidades para anidar, contribuyen a las preferencias del lugar de anidamiento exhibido a escala poblacional.

[Aim] Biogeographical comparisons of interaction networks help to elucidate differences in ecological communities and ecosystem functioning at large scales. Neotropical ecosystems have higher diversity and a different composition of frugivores and fleshy-fruited plants compared with Afrotropical systems, but a lack of intercontinental comparisons limits understanding of (a) whether plant–frugivore networks are structured in a similar manner, and (b) whether the same species traits define the roles of animals across continents., [Location] Afrotropics and Neotropics., [Time period] 1977-2015., [Taxa] Fleshy-fruited plants and frugivorous vertebrates., [Methods] We compiled a dataset comprising 17 Afrotropical and 48 Neotropical weighted seed-dispersal networks quantifying frugivory interactions between 1,091 fleshy-fruited plant and 665 animal species, comprising in total 8,251 interaction links between plants and animals. In addition, we compiled information on the body mass of animals and their degree of frugivory. We compared four standard network-level metrics related to interaction diversity and specialization, accounting for differences related to sampling effort and network location. Furthermore, we tested whether animal traits (body mass, degree of frugivory) differed between continents, whether these traits were related to the network roles of species and whether these relationships varied between continents., [Results] We found significant structural differences in networks between continents. Overall, Neotropical networks were less nested and more specialized than Afrotropical networks. At the species level, a higher body mass and degree of frugivory were associated with an increasing diversity of plant partners. Specialization of frugivores increased with the degree of frugivory, but only in the Neotropics., [Main conclusions] Our findings show that Afrotropical networks have a greater overlap in plant partners among vertebrate frugivores than the more diverse networks in the Neotropics that are characterized by a greater niche partitioning. Hence, the loss of frugivore species could have stronger impacts on ecosystem functioning in the more specialized Neotropical communities compared with the more generalized Afrotropical communities., We thank Beth A. Kaplin and Norbert J. Cordeiro for their guidance and support for P.J.D., who received a travel grant by The Center for Tropical Studies and Conservation (CTEC). L.C. and I.G. were supported by the Robert Bosch Foundation. D.M.D. (DE 2754/1‐1), F.S. (HE 3041/20‐1), M.Q., V.S., E.L.N. (Research Unit 823‐825), and K.B.G., M.S. and M.G.R.V. (FOR 1246) thank the German Research Foundation (DFG) for funding. F.A.F.J. acknowledges funding by a CAPES scholarship, N.F. and D.G.S. by the Robert Bosch Foundation, M.G., C.E., A.P. and M.A.P. by Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP 2010/52315‐7; 2015/15172‐7; 2016/18355‐8) and Conselho Nacional de Desenvolvimento Científico (CNPq), M.C.M. by Doctoral Fellowships from COLCIENCIAS and Rufford, M.S.S. by Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) and FONCyT (PICT2013‐2759 and PICT2016‐0608), P.G.B. by CONICET (PIP 2014‐592) and FONCyT (PICT 2013‐1280), R.A.R. by a Doctoral Fellowship from CONICET, R.H. and S.T. (IF/00441/2013) and M.C. (SFRH/BD/96050/2013) by Fundação para a Ciência e Tecnologia, Portugal, and A.T. (CGL2013‐44386‐P) and D.G. (CGL2015‐68963‐C2‐2‐R) by the Spanish government. T.

Species interactions, both mutualistic and antagonistic, are widely recognized as providing important ecosystem services. Fruit‐eating animals influence plant recruitment by increasing germination during gut passage and moving seeds away from conspecifics. However, relative to studies focused on the importance of frugivores for plant population maintenance, few studies target frugivores as ecosystem service providers, and frugivores are underappreciated as ecosystem service providers relative to other mutualists such as pollinators. Here, we use an accidental experiment to elucidate the role of seed dispersal by frugivores for maintaining a culturally and economically important plant, the donne’ sali chili (Capsicum frutescens) in the Mariana Islands. One of the islands (Guam) has lost nearly all of its native forest birds due to an invasive snake (Boiga irregularis), whereas nearby islands have relatively intact bird populations. We hypothesized that frugivore loss would influence chili recruitment and abundance, which could have economic and cultural impacts. By using video cameras, we confirmed that birds were the primary seed dispersers. We used captive bird feeding trials to obtain gut‐passed seeds to use in a seedling emergence experiment. The experiment showed that gut‐passed seeds emerged sooner and at a higher proportion than seeds from whole fruits. Consistent with our findings that birds benefit chilies, we observed lower chili abundance on Guam than on islands with birds. In a survey questionnaire of island residents, the majority of residents reported an association between the wild chili and local cultural values and traditions. In addition, we identified a thriving market for chili products, suggesting benefits of wild chilies to people in the Marianas both as consumers and producers. Our study therefore documents seed dispersal as both a cultural and a supporting ecosystem service. We provide a comprehensive case study on how seed‐dispersed plants decline in the absence of their disperser, and how to apply mixed‐methods in ecosystem service assessments. Furthermore, we suggest that scientists and resource managers may utilize fruit–frugivore mutualisms concerning socially valuable plants to gather support for frugivore and forest conservation efforts.

Seed ingestion by frugivorous vertebrates commonly benefits plants by moving seeds to locations with fewer predators and pathogens than under the parent. For plants with high local population densities, however, movement from the parent plant is unlikely to result in ‘escape’ from predators and pathogens. Changes to seed condition caused by gut passage may also provide benefits, yet are rarely evaluated as an alternative. Here, we use a common bird-dispersed chilli pepper (Capsicum chacoense) to conduct the first experimental comparison of escape-related benefits to condition-related benefits of animal-mediated seed dispersal. Within chilli populations, seeds dispersed far from parent plants gained no advantage from escape alone, but seed consumption by birds increased seed survival by 370% – regardless of dispersal distance – due to removal during gut passage of fungal pathogens and chemical attractants to granivores. These results call into question the pre-eminence of escape as the primary advantage of dispersal within populations and document two overlooked mechanisms by which frugivores can benefit fruiting plants.

The global decline of mutualists such as pollinators and seed dispersers may cause negative direct and indirect impacts on biodiversity. Mutualistic network models used to understand the stability of mutualistic systems indicate that species with low partner diversity are most vulnerable to coextinction following mutualism disruption. However, existing models have not considered how species vary in their dependence on mutualistic interactions for reproduction or survival, overlooking the potential influence of this variation on species' coextinction vulnerability and on network stability. Using global databases and field experiments focused on the seed dispersal mutualism, we found that plants and animals that depend heavily on mutualistic interactions have higher partner diversity. Under simulated network disruption, this empirical relationship strongly reduced coextinction because the species most likely to lose mutualists depend least on their mutualists. The pattern also reduced the importance of network structure for stability; nested network structure had little effect on coextinction after simulations incorporated the empirically derived relationship between partner diversity and mutualistic dependence. Our results highlight a previously unknown source of stability in mutualistic networks and suggest that differences among species in their mutualistic strategy, rather than network structure, primarily accounts for stability in mutualistic communities.

Rare plant species often suffer stronger conspecific negative density dependence (CNDD) in studies that assess the impact of local conspecific density on individual survival. All else equal, this causes a relative disadvantage among rare species that appears inconsistent with the role of CNDD in coexistence. The resolution to this apparent paradox is for lower species abundance to decrease the frequency of conspecific interactions sufficiently to outweigh the disadvantage of stronger CNDD. Whether this occurs in natural systems is untested because existing metrics do not isolate demographic impacts of CNDD, and it is also uncertain for tropical forest trees because the greater spatial aggregation observed in rare species could cause higher frequency of conspecific interactions despite lower abundance on the landscape. We develop a new metric, effective density-dependent mortality (EDDM), to quantify the proportion of individuals that are killed by density-dependent effects. We apply EDDM to a long-term study of seed fall and recruitment at Barro Colorado Island, Panama. Rare species had stronger CNDD but lower conspecific densities, and EDDM increased with abundance. Lower abundance, thus, reduces the frequency of conspecific interaction and, consequently, mortality associated with CNDD. This mechanism allows rare species to avoid a disadvantage-when-rare that would, all else equal, result from stronger CNDD in rare species. Our work provides empirical support for a resolution to the apparently paradoxical findings that rare species experience stronger CNDD and may help reconcile contrasting findings for the relationship between the CNDD strength and abundance.

Frugivores play differing roles in shaping dispersal patterns yet seed dispersal distance is rarely quantified across entire communities. We model seed dispersal distance using gut passage times and bird movement for the majority (39 interactions) of known bird–tree interactions on the island of Saipan to highlight differences in seed dispersal distances provided by the five avian frugivores. One bird species was found to be a seed predator rather than a disperser. The remaining four avian species dispersed seeds but differences in seed dispersal distance were largely driven by interspecific variation in bird movement rather than intraspecific variation in gut passage times. The median dispersal distance was at least 56 m for all species-specific combinations, indicating all species play a role in reducing high seed mortality under the parent tree. However, one species—the Micronesian Starling—performed 94% of dispersal events greater than 500 m, suggesting this species could be a key driver of long-distance dispersal services (e.g. linking populations, colonizing new areas). Assessing variation in dispersal patterns across this network highlights key sources of variation in seed dispersal distances and suggests which empirical approaches are sufficient for modelling how seed dispersal mutualisms affect populations and communities.

Plants can influence the source and severity of seed predation through various mechanisms; the use of secondary metabolites for chemical defense, for example, is well documented. Gut passage by frugivores can also reduce mortality of animal-dispersed seeds, although this mechanism has gained far less attention than secondary metabolites. Apart from influencing the severity of seed predation, gut passage may also influence the source of seed predation. In Bolivia, we compared impacts of these two mechanisms, gut passage and secondary metabolites, on the source of seed predation in Capsicum chacoense, a wild chili species that is polymorphic for pungency (individual plants either produce fruits and seeds containing or lacking capsaicinoids). Using physical exclosures, we isolated seed removal by insects, mammals, and birds; seeds in the trials were from either pungent or non-pungent fruits and were either passed or not passed by seed-dispersing birds. Pungency had little influence on total short-term seed removal by animals, although prior work on this species indicates that capsaicin reduces mortality caused by fungi at longer time scales. Gut passage strongly reduced removal by insects, altering the relative impact of the three predator types. The weak impact of pungency on short-term predation contrasts with previous studies, highlighting the context dependence of secondary metabolites. The strong impact of gut passage demonstrates that this mechanism alone can influence which seed predators consume seeds, and that impacts of gut passage can be larger than those of secondary metabolites, which are more commonly acknowledged as a defense mechanism.

Specialised natural enemies maintain forest diversity by reducing tree survival in a density- or distance-dependent manner. Fungal pathogens, insects and mammals are the enemy types most commonly hypothesised to cause this phenomenon. Still, their relative importance remains largely unknown, as robust manipulative experiments have generally targeted a single enemy type and life history stage. Here, we use fungicide, insecticide and physical exclosure treatments to isolate the impacts of each enemy type on two life history stages (germination and early seedling survival) in three tropical tree species. Distance dependence was evident for five of six species-stage combinations, with each enemy type causing distance dependence for at least one species stage and their importance varying widely between species and stages. Rather than implicating one enemy type as the primary agent of this phenomenon, our field experiments suggest that multiple agents acting at different life stages collectively contribute to this diversity-promoting mechanism.