Seed-size variation determines interspecific differential predation by mammals in a neotropical rain forest (original) (raw)
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Size-Related Differential Seed Predation in a Heavily Defaunated Neotropical Rain Forest
Biotropica, 2007
Hunting and habitat loss represent an increasingly common anthropogenic impact. Available evidence suggests that defaunation is typically differential: medium/large mammals are most affected while small rodents are less affected, or even favored. In heavily impacted areas, such as Los Tuxtlas, the residual mammalian fauna is dominated by small rodents. We tested the expectation that if small rodents will preferentially attack small-seeded species, large-seeded species may escape predation in the absence of medium/large seed predators and that this may affect germination and, possibly, recruitment. Experiments with caged rodents (Heteromys desmarestianus) under laboratory conditions showed a preference for smaller seeds. A field experiment involving seeds of contrasting size showed that small, unprotected seeds experienced a predation risk 30-times larger than protected seeds, while the effect of protection was not significant for large seeds. These patterns of predation led to significant differences in germination: protected small-seeded species had a fourfold greater germination than unprotected small-seeded species, while germination was not significantly different between exposed and protected large seeds. The observed contrasts in germination suggest that under heavy defaunation, small-seeded species are likely to be penalized by the overabundance of small rodents, while large-seeded species escape predation. Our results are consistent with the observation of dense seedling carpets dominated by large-seeded species in the understory of Los Tuxtlas. We speculate that the patterns we uncovered may explain, at least partly, the impoverished diversity of the understory vegetation that characterizes heavily defaunated understories and that this has the potential to affect forest regeneration.La cacería y la pérdida del hábitat representan un impacto antropogénico de creciente intensidad. La evidencia disponible sugiere que la defaunación es típicamente diferencial: los mamíferos medianos/grandes son los más afectados, mientras que los pequeños roedores son poco afectados o, incluso son favorecidos. En sitios fuertemente impactados, como Los Tuxtlas, la fauna residual dominante consiste de pequeños roedores. Aquí evaluamos la expectativa de que los pequeños roedores depredan preferentemente las especies de semillas pequeñas mientras que, en ausencia de depredadores medianos/grandes, las especies de semillas grandes escapan a la depredación; esto podría afectar la germinación y, posiblemente, el reclutamiento. En experimentos con el roedor Heteromys desmarestianus en condiciones de cautiverio, detectamos una mayor preferencia por las especies de semilla pequeña. En experimentos de exclusión (semillas protegidas y expuestas) en el campo, involucrando especies de semillas de tamaño contrastante encontramos que, en comparación con las semillas expuestas, las protegidas tuvieron una depredación 30 veces menor, mientras que el efecto de la exclusión no fue significativo en las especies de semilla grande. Estos patrones de depredación condujeron a diferencias significativas en la germinación: las semillas pequeñas y excluidas tuvieron una germinación cuatro veces mayor que las expuestas; la germinación de semillas grandes no difirió significativamente entre tratamientos. Dichos contrastes en la germinación sugieren que bajo situaciones de defaunación extrema, las especies de semilla pequeña tienden a ser mas castigadas por la sobre-abundancia de pequeños roedores, mientras que las grandes escapan a la depredación. Estos resultados son consistentes con la existencia de tapetes densamente dominados por plántulas de especies de semilla grande en el sotobosque de Los Tuxtlas. A partir de esto especulamos que los patrones encontrados explican, por lo menos en parte, el empobrecimiento florístico de la vegetación del sotobosque que caracteriza sitios fuertemente defaunados, y que esto puede afectar el potencial de regeneración de la selva.
Seed predation by Neotropical rainforest mammals
Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing ,1 kg), mediumsized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small-and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.
"Tropical tree species vary widely in their pattern of spatial dispersion. We focus on how seed predation may modify seed deposition patterns and affect the abundance and dispersion of adult trees in a tropical forest in India. Using plots across a range of seed densities, we examined whether seed predation levels by terrestrial rodents varied across six large-seeded, bird-dispersed tree species. Since inter-specific variation in density-dependent seed mortality may have downstream effects on recruitment and adult tree stages, we determined recruitment patterns close to and away from parent trees, along with adult tree abundance and dispersion patterns. Four species (Canarium resiniferum, Dysoxylum binectariferum, Horsfieldia kingii, and Prunus ceylanica) showed high predation levels (78.5–98.7%) and increased mortality with increasing seed density, while two species, Chisocheton cumingianus and Polyalthia simiarum, showed significantly lower seed predation levels and weak density-dependent mortality. The latter two species also had the highest recruitment near parent trees, with most abundant and aggregated adults. The four species that had high seed mortality had low recruitment under parent trees, were rare, and had more spaced adult tree dispersion. Biotic dispersal may be vital for species that suffer density-dependent mortality factors under parent trees. In tropical forests where large vertebrate seed dispersers but not seed predators are hunted, differences in seed vulnerability to rodent seed predation and density-dependent mortality can affect forest structure and composition."
Biotropica, 2007
Many of the mammals undergoing drastic declines in tropical forests worldwide are important seed dispersers and seed predators, and thus changes in mammal communities due to hunting will affect plant recruitment. It has been hypothesized that larger-seeded species will suffer greater reductions in seed removal and thus greater increases in predispersal seed predation than smaller-seeded species. We compared primary and secondary seed removal and predispersal seed predation of two tree species between hunted and nonhunted sites in Central Panama. Seeds of Oenocarpus mapora (Arecaceae) are 16-times greater in size than those of Cordia bicolor (Boraginaceae). We quantified primary seed removal and predispersal seed predation using seed traps, and we assessed secondary seed removal using seed removal plots. Primary removal of C. bicolor was 43 percent lower in the hunted sites, while primary removal of O. mapora was not significantly different. Secondary removal of unprotected O. mapora seeds on the ground was 59 percent lower in hunted sites, while secondary removal of C. bicolor was not significantly different. Predispersal seed predation of O. mapora by mammals was significantly lower in hunted sites, while predispersal seed predation by insects was not significantly different in either species. In combination with other studies, our results suggest that seed size is not a reliable predictor of the impacts of hunting. Mammal defaunation differentially affects stages and modes of seed dispersal and seed predation of different plant species, suggesting that these influences are complex and related to multiple plant traits.
Defaunation of large mammals leads to an increase in seed predation in the Atlantic forests
Defaunation can trigger cascading events in natural communities and may have strong consequences for plant recruitment in tropical forests. Several species of large seed predators, such as deer and peccaries, are facing dramatic population collapse in tropical forests yet we do not have information about the consequences of these extinctions for seed predation. Using remote camera traps we tested if defaunated forests have a lower seed predation rate of a keystone palm (Euterpe edulis) than pristine areas. Contrary to our expectation, we found that seed predation rates were 2.5 higher in defaunated forests and small rodents were responsible for most of the seeds eaten. Our results found that defaunation leads to changes in the seed predator communities with potential consequences for plant-animal interactions.
Oecologia, 1996
The effect of seed aggregation and distance from conspecific trees on seed predation was experimentally examined for two neotropical tree species, Macoubea guianensis (Apocynaceae) and Pouteria sp. (Sapotaceae) in a lowland tropical rain forest in northeastern Peru. Results of these experiments are discussed in the context of the Janzen-Connell model (Janzen 1970; Connell 1971), which predicts decreased seed survival near parent trees due to either density-or distance-responsive mortality, and Howe's model (Howe 1989) which predicts that trees with seeds dispersed in clumps (aggregated) will not suffer density-dependent predation, and will have higher survival of seeds near the parent tree than other trees. We also examined whether predation on seeds of these species was affected by seed placement in or near 30-m-wide strips regenerating after clear-cutting. Both species appeared to be mammal-dispersed but differed in how frugivores handled seeds, seed size, overall fruit crop size, and gemination time. Neither of the two species studied appeared to suffer seed predation in a manner predicted by the Janzen-Connell model, and patterns of seed predation for only one of the species was similar to predictions of Howe's model. For neither species did seed predation along the edge of, or in the center of, regenerating clear cuts differ from predation 15 m into the primary forest. For Pouteria, seed predation in and near regnerating strips was significantly greater than around forest trees, but the opposite pattern held for Macoubea. Overall, seed predation was much greater on Macoubea. The difference in seed predation for these
Oecologia, 2005
We investigated the role of seed predation by rodents in the recruitment of the fleshy-fruited trees Taxus baccata, Ilex aquifolium and Crataegus monogyna in temperate secondary forests in NW Spain. We measured the densities of dispersed seeds, early emerged seedlings, established recruits and adults, at four sites over a period of 2 years. Seed predation among species was compared by seed removal experiments and analysis of rodent larder-hoards. The three species differed markedly in local regeneration patterns. The rank order in the seed rain following decreasing seed density was Ilex, Taxus and Crataegus. However, Crataegus established 3.3 times more seedlings than Taxus. For all species, there was a positive linear relationship between the density of emerged seedlings and seed density, suggesting that recruitment was seed-rather than micrositelimited. A consistent pattern of seed selection among species was exerted by rodents, which preferred Taxus and, secondarily, Ilex seeds to Crataegus seeds. Predation ranking was the inverse of that of seed protection against predators, measured as the mass of woody coat per mass unit of the edible fraction. Recruitment potential, evaluated as the ratio of seedlings to seeds, was negatively related to seed predation, with the rank order Crataegus > Ilex > Taxus. The selective early recruitment limitation exerted by predation may have a demographic effect in the long term, as judged by the positive relationship between early seedling emergence and the density of established recruits. By modulating the pre-emptive competition for seed safe sites, rodents may preclude the progressive exclusion of species that produce low numbers of seeds (i.e. Crataegus) by those dominant in seed number (i.e. Ilex, Taxus), or at least foster the evenness for site occupation among seedlings of different species.
Oikos, 2006
. Trophic and non-trophic pathways mediate apparent competition through post-dispersal seed predation in a Patagonian mixed forest. Á Oikos 113: 469 Á480. Post-dispersal predation can be a major source of seed loss in temperate forests. Little is known, however, about how predator-mediated indirect interactions such as apparent competition alter survival patterns of canopy tree seeds. Understorey plants may enhance tree seed predation by providing sheltered habitat to granivores (non-trophic pathway). In addition, occurrence of different tree seeds in mixed patches may lead to short-term apparent competition between seed types, because of the granivores' foraging response to changes in food patch quality (trophic pathway). We hypothesised that understorey bamboo cover and mixing of seed species in food patches would both increase tree seed predation in a Nothofagus dombeyi(/Austrocedrus chilensis forest in northern Patagonia, Argentina. Seed removal experiments were conducted for three consecutive years (2000 Á2002) differing in overall granivory rates. Seed patch encounter and seed removal rates were consistently higher for the larger and more nutritious Austrocedrus seeds than for the smaller Nothofagus seeds. Seed removal was greater beneath bamboo than in open areas. This apparent competition pathway was stronger in a low-predation year (2000) than in high-predation years (2001 Á2002), suggesting a shift in microhabitat use by rodents. Patch composition had a significant, though weaker, impact on seed survival across study years, whereas seed density per patch enhanced encounter rates but did not influence seed removal. Removal of the less-preferred Nothofagus seeds increased in the presence of Austrocedrus seeds, but the reciprocal indirect effect was not observed. However, this non-reciprocal apparent competition between seed species was only significant in the high-predation years. Our study shows that granivore-mediated indirect effects can arise through different interaction pathways, affecting seed survival patterns according to the predator's preference for alternative seed types. Moreover, results indicate that the occurrence and relative strength of trophic vs non-trophic pathways of apparent competition may change under contrasting predation scenarios.