Habitat generalists drive nestedness in a tropical mountaintop insect metacommunity
Frederico S. NevesPedro Giovâni da SilvaRicardo SolarCássio Alencar NunesMarina do Vale BeirãoHumberto BrantFlávio Siqueira de CastroWesley DáttiloRoger GuevaraGeraldo Wilson Fernandes
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Abstract Nestedness is widely observed in natural metacommunities, but its underlying mechanisms are still poorly understood. The distribution of habitats in the landscape and differences in dispersal rates of distinct insect taxa can determine the nestedness of the metacommunity. Here, we evaluated how species habitat specialization contributes to metacommunity nestedness in insect groups with different dispersal capacities in a mountaintop landscape in south-eastern Brazil. We sampled ants, butterflies and dung beetles in two main habitats, naturally fragmented forest islands and a grassland matrix (campo rupestre), during both dry and rainy seasons. We classified species according to their degree of habitat specialization (generalists or specialists) based on the relative frequencies and abundances between these two contrasting habitats. Forty of 211 species were classified as habitat specialists, seven as habitat generalists. It was not possible to classify the remaining species. The metacommunity was nested in structure, with habitat generalist species contributing more to nestedness than habitat specialists. Nonetheless, habitat distribution in the landscape did not affect the nestedness of the metacommunity. Our findings reveal that species sorting (for habitat specialists) and mass effects (for habitat generalists) are concurrent processes in the mountaintop forest–grassland mosaic. Our study helps to advance our understanding of the differences in the distribution of generalist and specialist species in a tropical mountaintop landscape and improves our ability to predict and manage the increasingly adverse effects of changes in land use and climate on metacommunities and ecosystem functions.Keywords:
Nestedness
Metacommunity
Linyphiidae
Although there has been growing interest in the effect of dispersal on species diversity, much remains unknown about how dispersal occurring at multiple scales influences diversity. We used an experimental microbial landscape to determine whether dispersal occurring at two different scales - among local communities and among metacommunities - affects diversity differently. At the local scale, dispersal initially had a positive effect and subsequently a neutral effect on diversity, whereas at the metacommunity and landscape scales, dispersal showed a consistently negative effect. The timing in which dispersal affected beta diversity also differed sharply between local communities and metacommunities. These patterns were explained by scale- and time-dependent effects of dispersal in allowing spread of species and in removing spatial refuges from predators. Our results suggest that the relative contribution of opposing mechanisms by which dispersal affects diversity changes considerably over time and space in hierarchical landscapes in which dispersal occurs at multiple scales.
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A recent challenge in community ecology is to understand under what conditions local and regional processes may be more important in shaping community structure. We investigated the role of dispersal mode and generation time for communities of macroinvertebrates in two sets of connected ponds during three consecutive years. We found no evidence that generation time affected metacommunity structure, possibly because statistical power was limited because the range of generation times present was small. In contrast, we found that the spatial structure of the macroinvertebrate metacommunities differed with dispersal mode in one of two sets of ponds. Passive dispersers showed positive distance-dissimilarity correlations, suggesting mass effects via the pond connections. These correlations did not stretch beyond the first pond downstream suggesting that, in this chain of connected ponds, intervening ponds effectively buffered dispersal. Active dispersers did not show any consistent spatial pattern, possibly because intensive over-land dispersal homogenized the metacommunity. Thus, although pond connections probably equally promoted dispersal of actively and passively dispersing macroinvertebrates, the implications for the structure of their metacommunities may depend on their dispersal mode. We conclude that dispersal mode has the potential to affect the mechanisms that are integral to metacommunity structure.
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Metacommunity nestedness can be affected by both idiosyncratic species and species turnover, and diversity partitioning allows one to separate turnover and nested components within β-diversity. Thus, complimentary analysis of metacommunity nestedness and diversity partitioning allows for the identification of the underlying changes at both local and regional scales. We examined changes of fish assemblages in metacommunity nestedness and α-, β-, and γ-diversities resulting from the intense loss of native species and the invasion of nonnative species in Chinese highland lakes over the past 60 years. We found metacommunity nestedness rose markedly over time, following the loss of both β- and γ-diversity resulting from the loss of native species, and the increase of α diversity by the addition of nonnative species. This pattern is contradictory to the selective extinction leading to larger nestedness in natural ecosystems and indicates the human-induced negative effects on the metacommunity. However, β-diversity partitioning showed that the turnover component due to species replacement among lakes still contributes more than the nested component, suggesting the importance of avoiding setting conservation priorities based exclusively on metacommunity nestedness theory, but taking a more holistic metacommunity-approach to conservation instead.
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Abstract The structure of interactions between species within a community plays a key role in maintaining biodiversity. Previous studies found that the effects of these structures might vary substantially depending on interaction type, for example, a highly connected and nested architecture stabilizes mutualistic communities, while the stability of antagonistic communities is enhanced in modular and weakly connected structures. Here we show that, when network dynamics are modeled using a patch‐dynamic metacommunity framework, the qualitative differences between antagonistic and mutualistic systems disappear, with nestedness and modularity interacting to promote metacommunity persistence. However, the interactive effects are significantly weaker in antagonistic metacommunities. Our model also predicts an increase in connectance, nestedness, and modularity over time in both types of interaction, except in antagonistic networks, where nestedness declines. At steady state, we find a strong negative correlation between nestedness and modularity in both mutualistic and antagonistic metacommunities. These predictions are consistent with the structural trends found in a large data set of real‐world antagonistic and mutualistic communities.
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Difference in dispersal ability is a key driver of species coexistence in metacommunities. However, the available frameworks for interpreting species diversity patterns in natura often overlook trade-offs and evolutionary constraints associated with dispersal. Here, we build a metacommunity model accounting for dispersal evolution and a competition–dispersal trade-off. Depending on the distribution of carrying capacities among communities, species dispersal values are distributed either around a single strategy (evolutionarily stable strategy, ESS), or around distinct strategies (evolutionary branching, EB). We show that limited dispersal generates spatial aggregation of dispersal traits in ESS and EB scenarios, and that the competition–dispersal trade-off strengthens the pattern in the EB scenario. Importantly, individuals in larger (respectively (resp.) smaller) communities tend to harbour lower (resp. higher) dispersal, especially under the EB scenario. We explore how dispersal evolution affects species diversity patterns by comparing those from our model to the predictions of a neutral metacommunity model. The most marked difference is detected under EB, with distinctive values of both α- and β-diversity (e.g. the dissimilarity in species composition between small and large communities was significantly larger than neutral predictions). We conclude that, from an empirical perspective, jointly assessing community carrying capacity with species dispersal strategies should improve our understanding of diversity patterns in metacommunities.
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Although it is well‐known that dispersal of organisms within a metacommunity will influence patterns of coexistence and richness, theoretical and experimental studies generally assume that dispersal rates are constant through time. However, dispersal is often a highly variable process that can vary seasonally and/or when stochastic events (e.g. wind storms, droughts, floods) occur. Using a well‐known source–sink metacommunity model, we present novel predictions for local and regional species richness when stochasticity in dispersal is expressly considered. We demonstrate that dispersal stochasticity alters some of the predictions obtained with constant dispersal; the peak of the predicted hump‐shaped relationship between dispersal and local species richness is diminished and shifted towards higher values of dispersal. Dispersal stochasticity increases extinction probabilities of inferior competitor species particularly in metacommunities subjected to severe isolation events (i.e. decreases of dispersal) or homogenization events (i.e. sudden increases of dispersal). Our results emphasize how incorporating dispersal stochasticity into theoretical predictions will broaden our understanding of metacommunities dynamics and their responses to natural and human‐related disturbances.
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Traditionally metacommunity studies have quantified the relative importance of dispersal and environmental processes on observed β-diversity. Separating β-diversity into its replacement and nestedness components and linking such patterns to metacommunity drivers can provide richer insights into biodiversity organization across spatial scales. It is often very difficult to measure actual dispersal rates in the field and to define the boundaries of natural metacommunities. To overcome those limitations, we revisited an experimental metacommunity dataset to test the independent and interacting effects of environmental heterogeneity and dispersal on each component of β-diversity. We show that the balance between the replacement and nestedness components of β-diversity resulting from eutrophication changes completely depending on dispersal rates. Nutrient enrichment negatively affected local zooplankton diversity and generated a pattern of β-diversity derived from nestedness in unconnected, environmentally heterogeneous landscapes. Increasing dispersal erased the pattern of nestedness, whereas the replacement component gained importance. In environmentally homogeneous metacommunities, dispersal limitation created community dissimilarity via species replacement whereas the nestedness component remained low and unchanged across dispersal levels. Our study provides novel insights into how environmental heterogeneity and dispersal interact and shape metacommunity structure.
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Climate change is increasingly affecting the structure and dynamics of ecological communities both at local and at regional scales, and this can be expected to have important consequences for their robustness and long-term persistence. The aim of the present work is to analyse how the spatial structure of the landscape and dispersal patterns of species (dispersal rate and average dispersal distance) affects metacommunity response to two disturbances: (i) increased mortality during dispersal and (ii) local species extinction. We analyse the disturbances both in isolation and in combination. Using a spatially and dynamically explicit metacommunity model, we find that the effect of dispersal on metacommunity persistence is two-sided: on the one hand, high dispersal significantly reduces the risk of bottom-up extinction cascades following the local removal of a species; on the other hand, when dispersal imposes a risk to the dispersing individuals, high dispersal increases extinction risks, especially when dispersal is global. Large-bodied species with long generation times at the highest trophic level are particularly vulnerable to extinction when dispersal involves a risk. This suggests that decreasing the mortality risk of dispersing individuals by improving the quality of the habitat matrix may greatly increase the robustness of metacommunities.
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