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Bird-mediated seed dispersal is crucial for the regeneration and viability of ecosystems, often resulting in complex mutualistic species networks. Yet, how this mutualism drives the evolution of seed dispersing birds is still poorly understood. In the present study we combine whole genome re-sequencing analyses and morphometric data to assess the evolutionary processes that shaped the diversification of the Eurasian nutcracker (Nucifraga), a seed disperser known for its mutualism with pines (Pinus). Our results show that the divergence and phylogeographic patterns of nutcrackers resemble those of other non-mutualistic passerine birds and suggest that their early diversification was shaped by similar biogeographic and climatic processes. The limited variation in foraging traits indicates that local adaptation to pines likely played a minor role. Our study shows that close mutualistic relationships between bird and plant species might not necessarily act as a primary driver of evolution and diversification in resource-specialized birds., (© 2022. The Author(s).)

Plant recruitment is a multi-stage process determining population dynamics and species distributions. Still, we have limited understanding of how the successive demographic processes depend on the environmental context across species' distributional ranges. We conducted a large-scale transplant experiment to study recruitment of Pinus cembra over six years. We quantified the effects of environmental conditions on four demographic processes and identified the most limiting across and beyond the pines' elevational range over several years. Realized transition probabilities of the demographic processes varied substantially across the species' distributional range. Seed deposition decreased from the lower to the upper elevational range margin by 90%, but this reduction was offset by increased seed germination and seedling survival. Dispersal limitation at the upper range margin potentially stems from unsuitable seed caching conditions for the animal seed disperser, whereas increased seed germination might result from enemy escape from fungal pathogens and favourable abiotic conditions at the upper range margin. Our multi-year experiment demonstrates that environmental context is decisive for the local relevance of particular demographic processes. We conclude that experimental studies identifying the limiting demographic processes controlling species distributions are key for projecting future range dynamics of plants.

The species composition of local communities varies in space, and its similarity generally decreases with increasing geographic distance between communities, a phenomenon known as distance decay of similarity. It is, however, not known how changes in local species composition affect ecological processes, that is, whether they lead to differences in the local composition of species' functional roles. We studied eight seed-dispersal networks along the South American Andes and compared them with regard to their species composition and their composition of functional roles. We tested (1) if changes in bird species composition lead to changes in the composition of bird functional roles, and (2) if the similarity in species composition and functional-role composition decreased with increasing geographic distance between the networks. We also used cluster analysis to (3) identify bird species with similar roles across all networks based on the similarity in the plants they consume, (i) considering only the species identity of the plants and (ii) considering the functional traits of the plants. Despite strong changes in species composition, the networks along the Andes showed similar composition of functional roles. (1) Changes in species composition generally did not lead to changes in the composition of functional roles. (2) Similarity in species composition, but not functional-role composition, decreased with increasing geographic distance between the networks. (3) The cluster analysis considering the functional traits of plants identified bird species with similar functional roles across all networks. The similarity in functional roles despite the high species turnover suggests that the ecological process of seed dispersal is organized similarly along the Andes, with similar functional roles fulfilled locally by different sets of species. The high species turnover, relative to functional turnover, also indicates that a large number of bird species are needed to maintain the seed-dispersal process along the Andes., (© 2020 The Authors. Ecology published byWiley Periodicals, Inc. on behalf of Ecological Society of America.)

Downsizing of animal communities due to defaunation is prevalent in many ecosystems. Yet, we know little about its consequences for ecosystem functions such as seed dispersal. Here, we use eight seed-dispersal networks sampled across the Andes and simulate how downsizing of avian frugivores impacts structural network robustness and seed dispersal. We use a trait-based modeling framework to quantify the consequences of downsizing-relative to random extinctions-for the number of interactions and secondary plant extinctions (as measures of structural robustness) and for long-distance seed dispersal (as a measure of ecosystem function). We find that downsizing leads to stronger functional than structural losses. For instance, 10% size-structured loss of bird species results in almost 40% decline of long-distance seed dispersal, but in less than 10% of structural loss. Our simulations reveal that measures of the structural robustness of ecological networks underestimate the consequences of animal extinction and downsizing for ecosystem functioning.

Seed dispersal is an important ecosystem function, but it is contentious how structural and functional diversity of plant and bird communities are associated with seed-dispersal functions. We used structural equation models to test how structural (i.e., abundance, species richness) and functional diversity (i.e., functional dispersion and community-weighted means of functional traits) of fruiting plants and frugivorous birds directly and indirectly influence the respective components of fruit removal. We recorded plant and bird diversity in point counts and observed plant-frugivore interactions in a tropical mountain forest in Ecuador. We also recorded plant and bird morphological traits to calculate measures of functional diversity. We found that fruit abundance had a positive direct effect on bird abundance, which directly and indirectly mediated the abundance of removed fruits. Plant and bird species richness were only directly related to the richness of the removed fruits. Functional dispersion of the plant community was positively associated to that of the bird community and to that of the removed fruits. Consistently, we found positive associations between community-weighted means of plant and bird traits and between community-weighted means of plant traits and that of plants with removed fruits. In contrast, community-weighted means of the bird community were unrelated to that of the removed fruits. Overall, our results suggest that plant abundance directly and indirectly influences fruit removal, likely because of avian fruit tracking. However, we did not find strong links between the functional diversity of the frugivore community and removed fruits, suggesting that other factors in addition to plant-animal trait matching might be important for the functional diversity of removed fruits. Our findings highlight the importance of frugivore abundance for maintaining seed dispersal by animals in tropical forests.

Plant regeneration is essential for maintaining forest biodiversity and ecosystem functioning, which are globally threatened by human disturbance. Here we present the first integrative meta-analysis on how forest disturbance affects multiple ecological processes of plant regeneration including pollination, seed dispersal, seed predation, recruitment and herbivory. We analysed 408 pairwise comparisons of these processes between near-natural and disturbed forests. Human impacts overall reduced plant regeneration. Importantly, only processes early in the regeneration cycle that often depend on plant-animal interactions, i.e. pollination and seed dispersal, were negatively affected. Later processes, i.e. seed predation, recruitment and herbivory, showed overall no significant response to human disturbance. Conserving pollination and seed dispersal, including the animals that provide these services to plants, should become a priority in forest conservation efforts globally.

Seed dispersal is a key component in a plant's life cycle, allowing the colonization of new habitat and eventually enabling range shifts. For animal‐dispersed plants, the behaviour of seed‐dispersing animals has direct consequences for the fate of a seed. However, there is little empirical evidence on how individual behaviour of animals affects seed dispersal.Here we investigated the effects of individual behaviour on seed dispersal distance and location by analysing movement data of spotted nutcrackers (Nucifraga caryocatactes) during harvest and scatter‐hoarding of Swiss stone pine (Pinus cembra) seeds. To assess individual variation in seed dispersal patterns, we compared daytime hourly displacement patterns across individuals. We further identified seed caching locations of individual birds and analysed their potential for pine regeneration.We found that nutcrackers showed one of two distinct behaviours: short‐distance seed dispersal and long‐distance seed dispersal. The caching locations resulting from the two behaviours differed markedly in the regeneration potential of Swiss stone pine: seeds transported over short distances were cached at high elevations with high regeneration potential within the pine's current habitat, while seeds transported over long distances were cached at low elevations with low regeneration potential outside of the pine's habitat. The behaviours were not associated with bird body mass, relative wing length or sex.Our results emphasize that intraspecific individual behaviours of seed‐dispersing animals can have far‐reaching consequences for ecosystem functions. As a major determinant of seed dispersal dynamics, individual behaviour should be taken into account for a better understanding of ecosystem functions under current and future environmental conditions. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Questions: How do seed rain biomass and richness change from old‐growth tropical forests to pastures at different elevations? How do seed mass and seed dispersal mode change from forests to pastures across these elevations? What implications do these changes have for the recovery of deforested areas in tropical mountains? Location: Old‐growth montane forests and livestock pastures along an elevational gradient (1,000–3,000 m a.s.l.), located at Podocarpus National Park, San Francisco Reserve and surrounding agricultural lands, Andes of southern Ecuador. Methods: We collected seed rain for a 3‐month period using 324 traps installed at eighteen 1‐ha plots across elevations. Half of the traps were installed in nine 1‐ha forest plots, and the other half in nine 1‐ha pasture plots. For each trap, we identified the seeds and measured seed rain biomass and richness. We also recorded seed traits and calculated community‐weighted means of seed mass and seed dispersal mode (proportion of endozoochory). Results: Forests received a higher seed rain biomass than pastures, but only at the lowest elevation. Seed rain richness did not differ between habitat types at all elevations. Community‐weighted means of seed mass and the proportion of endozoochorous species declined from forests to pastures, especially at the lower elevations. Conclusions: Although seed rain biomass and richness were overall similar between forests and pastures, large‐seeded and endozoochorous species were generally poorly represented in the seed rain of pastures compared with that of forests. These findings show that biomass and richness of seed rain may be insufficient to quantify the restoration potential of natural seed rain in deforested areas. Information on seed traits, such as seed mass and seed dispersal mode, is important to optimize restoration efforts towards the regeneration of diverse old‐growth forests along elevational gradients in tropical mountains. [ABSTRACT FROM AUTHOR]

Aim: Climate change causes shifts in species ranges globally. Terrestrial plant species often lag behind temperature shifts, and it is unclear to what extent animal‐dispersed plants can track climate change. Here, we estimate the ability of bird‐dispersed plant species to track future temperature change on a tropical mountain. Location: Tropical elevational gradient (500–3500 m.a.s.l.) in the Manú biosphere reserve, Peru. Time period: From 1960–1990 to 2061–2080. Taxa: Fleshy‐fruited plants and avian frugivores. Methods: Using simulations based on the functional traits of avian frugivores and fruiting plants, we quantified the number of long‐distance dispersal (LDD) events that woody plant species would require to track projected temperature shifts on a tropical mountain by the year 2070 under different greenhouse gas emission scenarios [representative concentration pathway (RCP) 2.6, 4.5 and 8.5]. We applied this approach to 343 bird‐dispersed woody plant species. Results: Our simulations revealed that bird‐dispersed plants differed in their climate‐tracking ability, with large‐fruited and canopy plants exhibiting a higher climate‐tracking ability. Our simulations also suggested that even under scenarios of strong and intermediate mitigation of greenhouse gas emissions (RCP 2.6 and 4.5), sufficient upslope dispersal would require several LDD events by 2070, which is unlikely for the majority of woody plant species. Furthermore, the ability of plant species to track future changes in temperature increased in simulations with a low degree of trait matching between plants and birds, suggesting that plants in generalized seed‐dispersal systems might be more resilient to climate change. Main conclusion: Our study illustrates how the functional traits of plants and animals can inform predictive models of species dispersal and range shifts under climate change and suggests that the biodiversity of tropical mountain ecosystems is highly vulnerable to future warming. The increasing availability of functional trait data for plants and animals globally will allow parameterization of similar models for many other seed‐dispersal systems. [ABSTRACT FROM AUTHOR]

Network approaches provide insight into the complex web of interspecific interactions that structure ecological communities. However, because data on the functional outcomes of ecological networks are very rarely available, the effect of network structure on ecosystem functions, such as seed dispersal, is largely unknown. Here, we develop a new approach that is able to link interaction networks to a trait‐based seed–dispersal model to estimate community‐wide seed dispersal distances. We simulated networks, using a niche model based on size‐matching between plants and birds, that varied in the degree of niche partitioning, i.e. the overlap in interaction partners between coexisting species. We found that community‐wide dispersal distances were longest when networks had low degrees of niche partitioning. We further found that dispersal distances of plant species with small fruits peaked in models without niche partitioning, whereas dispersal distances of medium and large‐fruited plants peaked at low degrees of niche partitioning. Our simulations demonstrate that the degree of niche partitioning between species is an important determinant of the ecological functions derived from ecological networks and that simulation approaches can provide new insights into the relationship between the structural and functional components of ecological networks. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract: Abiotic factors, biotic interactions and dispersal ability determine the spatial distribution of species. Theory predicts that abiotic constraints set range limits under harsh climatic conditions and biotic interactions set range limits under benign climatic conditions, whereas dispersal ability should limit both ends of the distribution. However, empirical studies exploring how these three components jointly affect species across environmental gradients are scarce. Here, we present a study that jointly examines these factors to investigate the constraints of the recruitment of Swiss stone pine (Pinus cembra) at and beyond its upper and lower elevational range limits in the Swiss Alps. We investigated the natural recruitment of pines and additionally conducted seed transplant experiments to test how much abiotic factors (mean summer and winter temperatures, soil moisture), biotic interactions (understorey vegetation cover, canopy cover, seed predation) and/or seed deposition by the spotted nutcracker (Nucifraga caryocatactes) affect pine establishment. We found significant effects of biotic interactions and seed deposition by spotted nutcrackers on the recruitment of Swiss stone pine at both the upper and lower elevational range, but could not detect significant effects of abiotic factors. Importantly, dispersal limitation rather than temperature and soil moisture restricted the recruitment of pines at the upper elevational range. Synthesis. Our study highlights the importance of biotic interactions and dispersal ability in setting the upper range limits of species that have been regarded as mainly controlled by climate. This suggests that potential range shifts of plants in response to climate warming may strongly depend on seed dispersal and biotic interactions and not only on climatic factors. [ABSTRACT FROM AUTHOR]

Many animals hoard seeds for later consumption and establish seed caches that are often located at sites with specific environmental characteristics. One explanation for the selection of non-random caching locations is the avoidance of pilferage by other animals. Another possible hypothesis is that animals choose locations that hamper the perishability of stored food, allowing the consumption of unspoiled food items over long time periods., We examined seed perishability and pilferage avoidance as potential drivers for caching behaviour of spotted nutcrackers ( Nucifraga caryocatactes) in the Swiss Alps where the birds are specialized on caching seeds of Swiss stone pine ( Pinus cembra)., We used seedling establishment as an inverse measure of seed perishability, as established seedlings cannot longer be consumed by nutcrackers. We recorded the environmental conditions (i.e. canopy openness and soil moisture) of seed caching, seedling establishment and pilferage sites., Our results show that sites of seed caching and seedling establishment had opposed microenvironmental conditions. Canopy openness and soil moisture were negatively related to seed caching but positively related to seedling establishment, i.e. nutcrackers cached seeds preferentially at sites where seed perishability was low. We found no effects of environmental factors on cache pilferage, i.e. neither canopy openness nor soil moisture had significant effects on pilferage rates. We thus could not relate caching behaviour to pilferage avoidance., Our study highlights the importance of seed perishability as a mechanism for seed-caching behaviour, which should be considered in future studies. Our findings could have important implications for the regeneration of plants whose seeds are dispersed by seed-caching animals, as the potential of seedlings to establish may strongly decrease if animals cache seeds at sites that favour seed perishability rather than seedling establishment. [ABSTRACT FROM AUTHOR]

Spatial patterns of seed dispersal and recruitment of fleshy-fruited plants in tropical forests are supposed to be driven by the activity of animal seed dispersers, but the spatial patterns of seed dispersal, seedlings and saplings have rarely been analyzed simultaneously. We studied seed deposition and recruitment patterns of three Clusia species in a tropical montane forest of the Bolivian Andes and tested whether these patterns changed between habitat types (forest edge vs. forest interior), distance to the fruiting tree and consecutive recruitment stages of the seedlings. We recorded the number of seeds deposited in seed traps to assess the local seed-deposition pattern and the abundance and distribution of seedlings and saplings to evaluate the spatial pattern of recruitment. More seeds were removed and deposited at the forest edge than in the interior. The number of deposited seeds decreased with distance from the fruiting tree and was spatially clustered in both habitat types. The density of 1-yr-old seedlings and saplings was higher at forest edges, whereas the density of 2-yr-old seedlings was similar in both habitat types. While seedlings were almost randomly distributed, seeds and saplings were spatially clustered in both habitat types. Our findings demonstrate systematic changes in spatial patterns of recruits across the plant regeneration cycle and suggest that the differential effects of biotic and abiotic factors determine plant recruitment at the edges and in the interior of tropical montane forests. These differences in the spatial distribution of individuals across recruitment stages may have strong effects on plant community dynamics and influence plant species coexistence in disturbed tropical forests. [ABSTRACT FROM AUTHOR]