Pathogens are hypothesized to play an important role in the maintenance of tropical forest plant species richness. Notably, species richness may be promoted by incomplete filling of niche space due interactions of host populations with their pathogens. A potentially important group of pathogens are endophytic fungi, which asymptomatically colonize plants and are diverse and abundant in tropical ecosystems. Endophytes may alter competitive abilities of host individuals and improve host fitness under stress, but may also become pathogenic. Little is known of the impacts of endophytes on niche-space filling of their hosts.Here we evaluate how a widespread fungal endophyte infecting a common tropical palm influences its recruitment and survival in natural ecosystems, and whether this impact is modulated by the abiotic environment, potentially constraining host niche-space filling. Iriartea deltoidea dominates many wet lowland Neotropical forests. Diplodia mutila is a common asymptomatic endophyte in mature plants; however, it causes disease in some seedlings. We investigated the effects of light availability on D. mutila disease expression.We found I. deltoidea seedlings to preferentially occur under shady conditions. Correspondingly, we also found that high light triggers endophyte pathogenicity, while low light favors endosymbiotic development, constraining recruitment of endophyte-infested seedlings to shaded understory by reducing seedling survival in direct light. Pathogenicity of D. mutila under high light is proposed to result from light-induced production of H2O2 by the fungus, triggering hypersensitivity, cell death, and tissue necrosis in the palm. This is the first study to demonstrate that endophytes respond to abiotic factors to influence plant distributions in natural ecosystems; and the first to identify light as a factor influencing where an endophyte is placed on the endosymbiont–pathogen continuum. Our findings show that pathogens can indeed constrain niche-space filling of otherwise successful tropical plant species, providing unoccupied niche space for other species.
To advance our understanding of the processes that govern the assembly of palm communities and the local coexistence of numerous palm species, we here synthesize available information in the literature on species diversity and growth-form composition in palm communities across the Americas. American palm communities surveyed had 4-48 (median 16) species in study plots covering 0.09-7.2 ha. Climate, soils, hydrology, and topography are the main factors determining palm community species richness. Tropical lowland terra firme rain forests are the most species-rich whereas forests that are inundated or grow on sandy soils or in areas with seasonal climate have much fewer species. Palm communities in the central-western Amazon and in Central America are significantly richer than the average region and those in the Caribbean significantly poorer in species. As for branching, the 789 species of tropical American palms belong to Corner's model (solitary, 268 species, 33%), Tomlinsons model (cespitose, 521 species, 66%) and Schoute's model (dichotomous branching, three species, <1%). We assigned the species to eight different growth forms: (i) Large tall-stemmed Palms (102 spp), (ii) Large-leaved medium-short-stemmed Palms (31 spp), (iii) Medium-sized Palms (95 spp), (iv) Medium/Small Palms with Stout Stem (42 spp), (v) Small Palms (423 spp), (vi) Large acaulescent Palms (28 spp), (vii) Small acaulescent Palms (56 spp), and (viii) Climbing Palms (12 spp). The eight growth forms are differently represented in the palm communities, and the categories Small Palms and Large tall-stemmed Palms dominate the communities both in terms of species richness and in number of individuals.
Aim It is a central issue in ecology and biogeography to understand what governs community assembly and the maintenance of biodiversity in tropical rain forest ecosystems. A key question is the relative importance of environmental species sorting (niche assembly) and dispersal limitation (dispersal assembly), which we investigate using a large dataset from diverse palm communities. Location Lowland rain forest, western Amazon River Basin, Peru. Methods We inventoried palm communities, registering all palm individuals and recording environmental conditions in 149 transects of 5 m × 500 m. We used ordination, Mantel tests and indicator species analysis (ISA) to assess compositional patterns, species responses to geographical location and environmental factors. Mantel tests were used to assess the relative importance of geographical distance (as a proxy for dispersal limitation) and environmental differences as possible drivers of dissimilarity in palm species composition. We repeated the Mantel tests for subsets of species that differ in traits of likely importance for habitat specialization and dispersal (height and range size). Results We found a strong relationship between compositional dissimilarity and environmental distance and a weaker but also significant relationship between compositional dissimilarity and geographical distance. Consistent with expectations, relationships with environmental and geographical distance were stronger for understorey species than for canopy species. Geographical distance had a higher correlation with compositional dissimilarity for small‐ranged species compared with large‐ranged species, whereas the opposite was true for environmental distance. The main environmental correlates were inundation and soil nutrient levels. Main conclusions The assembly of palm communities in the western Amazon appears to be driven primarily by species sorting according to hydrology and soil, but with dispersal limitation also playing an important role. The importance of environmental characteristics and geographical distance varies depending on plant height and geographical range size in agreement with functional predictions, increasing our confidence in the inferred assembly mechanisms.
Phylogenetic information can provide important insight into the assembly of hyper‐diverse tropical rainforest communities. The hierarchical nature and scale dependency of phylogenetic community structure can be used to disentangle complex networks of assembly mechanisms. Here, we outline how particular assembly mechanisms can be identified by analysing phylogenetic community structure on different scales related to space and environment. We present a case study on a large set of species‐rich palm communities from the western Amazon basin. We aimed to elucidate the assembly of these communities by separating the roles of broad‐scale biogeographic processes, habitat filtering and intra‐habitat processes. Overall, we found phylogenetic clustering, which could be caused by a variety of different processes. When we reduced the species pool according to regions, habitats or soil sand content, we found that the structure was consistent with a combination of regional biogeographic differentiation, habitat filtering and within‐habitat processes. The habitat filtering effect was strongest, although it varied in strength among communities in different habitats. Some phylogenetic structure could not be accounted for by biogeography or habitat, suggesting that intra‐habitat processes such as micro‐environmental filtering or biotic interactions also play a role. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••.
Summary1. Local and regional patterns of plant species richness in tropical rain forests, as well as their possible drivers, remain largely unexplored. The main hypotheses for local species richness (alpha diversity) are (i) local environmental determinism with species-saturated communities, and (ii) regional control, in which the immigration of species from the regional species pool (gamma diversity) determines how many species coexist locally. The species pool hypothesis suggests a combined influence of local and regional drivers on alpha diversity. Differences in gamma diversity may arise from divergent environmental conditions or biogeographic histories. 2. We investigated the cross-scale determinants of palm alpha and gamma diversity across the western Amazon using a large field-based data set: a census of all palm individuals in 312 transects, totalling 98 species. We used regression-based variation partitioning to understand how habitat, topography and region influence alpha diversity, and correlations to assess the importance of the present environment (climate, soil, regional topography) and history (long-term habitat stability) for average regional alpha diversity and gamma diversity, including the link between these two diversity measures (species pool effect). 3. Variation in alpha diversity was primarily explained by region (36%) and habitat (18%), whereas the effect of topography was negligible (1%). Within habitats, region was even more important (up to 69% explained variation). Within regions, habitat and topography covaried and had a variable but an important influence. The pure effect of topography remained of minor importance (up to 13%). 4. Average regional alpha diversity was related to gamma diversity, precipitation seasonality and possibly long-term habitat stability. Gamma diversity was related to long-term habitat stability, and possibly current climate. 5. Synthesis. Gamma diversity strongly influenced alpha diversity, although a clear influence of local environment was also evident, notably habitat type, with a minor, more geographically variable effect of small-scale topography. Apart from gamma diversity, the factor most strongly related to regional alpha diversity was precipitation seasonality, while gamma diversity itself was strongly linked to long-term habitat stability. These results imply that plant species richness is contingent on both contemporary and historical factors with a strong link between local species richness and the regional species pool.
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