Scatter-hoarding birds disperse seeds to sites unfavorable for plant regeneration.

Scatter-hoarding birds provide effective long-distance seed dispersal for plants. Transporting seeds far promotes population spread, colonization of new areas, and connectivity between populations. However, whether seeds transported over long distances are deposited in habitats favorable to plant regeneration has rarely been investigated, mainly due to methodological constraints. To investigate dispersal patterns and distances of Swiss stone pine (Pinus cembra) seeds we utilized advances in tracking technology to track the movements of their sole disperser, the spotted nutcracker (Nucifraga caryocatactes). We found routine individual movements between single seed harvesting and seed caching site. Harvesting sites of individual birds overlapped, whereas seed caching sites were separated and located on average 5.3 km away from the harvesting site. Interestingly, most distant caching sites were located at low elevations and in spruce forest, where Swiss stone pine does not naturally occur. This suggests that nutcrackers disperse seeds over long distances but that a large portion of these seeds are cached outside the known pine habitat. Therefore, we conclude that the implications of such long-distance seed dispersal movements for plant populations should be carefully considered in combination with the effects of habitat quality on plant recruitment.

Tricky partners: native plants show stronger interaction preferences than their exotic counterparts.

In ecological networks, neutral predictions suggest that species' interaction frequencies are proportional to their relative abundances. Deviations from neutral predictions thus correspond to interaction preferences (when positive) or avoidances (when negative), driven by nonneutral (e.g., niche-based) processes. Exotic species interact with many partners with which they have not coevolved, and it remains unclear whether this systematically influences the strength of neutral processes on interactions, and how these interaction-level differences scale up to entire networks. To fill this gap, we compared interactions between plants and frugivorous birds at nine forest sites in New Zealand varying in the relative abundance and composition of native and exotic species, with independently sampled data on bird and plant abundances from the same sites. We tested if the strength and direction of interaction preferences differed between native and exotic species. We further evaluated whether the performance of neutral predictions at the site level was predicted by the proportion of exotic interactions in each network from both bird and plant perspectives, and the species composition in each site. We found that interactions involving native plants deviated more strongly from neutral predictions than did interactions involving exotics. This "pickiness" of native plants could be detrimental in a context of global biotic homogenization where they could be increasingly exposed to novel interactions with neutrally interacting mutualists. However, the realization of only a subset of interactions in different sites compensated for the neutrality of interactions involving exotics, so that neutral predictions for whole networks did not change systematically with the proportion of exotic species or species composition. Therefore, the neutral and niche processes that underpin individual interactions may not scale up to entire networks. This shows that seemingly simplistic neutral assumptions entail complex processes and can provide valuable understanding of community assembly or invasion dynamics.

Frugivore species maintain their structural role in the trophic and spatial networks of seed dispersal interactions.

Trophic relationships have inherent spatial dimensions associated with the sites where species interactions, or their delayed effects, occur. Trophic networks among interacting species may thus be coupled with spatial networks linking species and habitats whereby animals connect patches across the landscape thanks to their high mobility. This trophic and spatial duality is especially inherent in processes like seed dispersal by animals, where frugivores consume fruit species and deposit seeds across habitats. We analysed the frugivore-plant interactions and seed deposition patterns of a diverse assemblage of frugivores in a heterogeneous landscape in order to determine whether the roles of frugivores in network topology are correlated across trophic and spatial networks of seed dispersal. We recorded fruit consumption and seed deposition by birds and mammals during 2 years in the Cantabrian Range (N Spain). We then constructed two networks of trophic (i.e. frugivore-plant) and spatial (i.e. frugivore-seed deposition habitat) interactions and estimated the contributions of each frugivore species to the network structure in terms of nestedness, modularity and complementary specialization. We tested whether the structural role of frugivore species was correlated across the trophic and spatial networks, and evaluated the influence of each frugivore abundance and body mass in that relationship. Both the trophic and the spatial networks were modular and specialized. Trophic modules matched medium-sized birds with fleshy-fruited trees, and small bird and mammals with small-fruit trees and shrubs. Spatial modules associated birds with woody canopies, and mammals with open habitats. Frugivore species maintained their structural role across the trophic and spatial networks of seed dispersal, even after accounting for frugivore abundance and body mass. The modularity found in our system points to complementarity between birds and mammals in the seed dispersal process, a fact that may trigger landscape-scale secondary succession. Our results open up the possibility of predicting the consumption pattern of a diverse frugivore community, and its ecological consequences, from the uneven distribution of fleshy-fruit resources in the landscape.

RESUMEN Las relaciones tróficas tienen una dimensión espacial asociada a los sitios donde ocurren o tienen lugar sus efectos. Las redes tróficas entre especies pueden, por tanto, emparejarse con las redes espaciales que vinculan a las especies con sus hábitats, y donde los animales conectan parches gracias a su alta movilidad. Esta dualidad trófica y espacial es especialmente inherente a procesos como la dispersión de semillas, donde los animales frugívoros consumen distintas especies de plantas con fruto carnoso y depositan sus semillas en distintos hábitats. Aquí analizamos las interacciones frugívoro-planta y los patrones de deposición de semillas de un conjunto diverso de frugívoros en un paisaje heterogéneo, con el fin de determinar si el papel topológico de los frugívoros en la red trófica de dispersión de semillas se correlaciona con dicho papel en la red espacial de deposición. Para ello, registramos el consumo de frutos y la deposición de semillas por aves y mamíferos durante dos años en la Cordillera Cantábrica (N España). Construimos dos redes de interacciones, una trófica (frugívoro-planta) y otra espacial (frugívoro-hábitat de deposición de semillas), y estimamos la contribución de las especies animales a la estructura de cada red, en términos de anidamiento, modularidad y especialización. Analizamos si el papel estructural de los distintos frugívoros se correlacionaba a través de ambas redes, y evaluamos la influencia de la abundancia relativa y la masa corporal de los frugívoros en dicha relación. Encontramos un patrón general de modularidad y especialización tanto en la red trófica como en la espacial. Los módulos tróficos agruparon principalmente a aves de mediano tamaño con árboles con frutos carnosos, y a aves de pequeño tamaño y mamíferos con arbustos de fruto carnoso. Los módulos espaciales asociaron en general a las aves con hábitats arbolados y a los mamíferos con hábitats abiertos. Las especies de frugívoros mantuvieron su papel estructural tanto en la red trófica como en la espacial, incluso después de tener en cuenta la abundancia relativa y la masa corporal de los frugívoros. El patrón de modularidad detectado en el sistema apunta a una importante complementariedad funcional entre aves y mamíferos en el proceso de dispersión de semillas, de modo que puede favorecer la sucesión secundaria de la vegetación a escala de paisaje. Nuestros resultados abren la posibilidad de predecir el patrón de consumo de una comunidad diversa de frugívoros así como sus consecuencias ecológicas a partir de la distribución desigual de los recursos tróficos en el paisaje.

Downsizing of animal communities triggers stronger functional than structural decay in seed-dispersal networks.

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.

Defaunation effects on plant recruitment depend on size matching and size trade-offs in seed-dispersal networks.

Defaunation by humans causes a loss of large animals in many ecosystems globally. Recent work has emphasized the consequences of downsizing in animal communities for ecosystem functioning. However, no study so far has integrated network theory and life-history trade-offs to mechanistically evaluate the functional consequences of defaunation in plant-animal networks. Here, we simulated an avian seed-dispersal network and its derived ecosystem function seedling recruitment to assess the relative importance of different size-related mechanisms. Specifically, we considered size matching (between bird size and seed size) and size trade-offs, which are driven by differences in plant or animal species abundance (negative size-quantity relationship) as well as in recruitment probability and disperser quality (positive size-quality relationship). Defaunation led to impoverished seedling communities in terms of diversity and seed size, but only if models accounted for size matching. In addition, size trade-off in plants, in concert with size matching, provoked rapid decays in seedling abundance in response to defaunation. These results underscore a disproportional importance of large animals for ecosystem functions. Downsizing in ecological networks will have severe consequences for ecosystem functioning, especially in interaction networks that are structured by size matching between plants and animals.