The Geography of Metacommunities: Landscape Characteristics Drive Geographic Variation in the Assembly Process through Selecting Species Pool Attributes.

AbstractThe nonrandom association between landscape characteristics and the dominant life history strategies observed in species pools is a typical pattern in nature. Here, we argue that these associations determine predictable changes in the relative importance of assembly mechanisms along broadscale geographic gradients (i.e., the geographic context of metacommunity dynamics). To demonstrate that, we employed simulation models in which groups of species with the same initial distribution of niche breadths and dispersal abilities interacted across a wide range of landscapes with contrasting characteristics. By assessing the traits of dominant species in the species pool in each landscape type, we determined how different landscape characteristics select for different life history strategies at the metacommunity level. We analyzed the simulated data using the same analytical approaches used in the study of empirical metacommunities to derive predictions about the causal relationships between landscape characteristics and dominant life histories in species pools, as well as their reciprocal influence on empirical inferences regarding the assembly process. We provide empirical support for these predictions by contrasting the assembly of moth metacommunities in a tropical versus a temperate mountainous landscape. Together, our model framework and empirical analyses demonstrate how the geographic context of metacommunities influences our understanding of community assembly across broadscale ecological gradients.

Spatial versus spatio-temporal approaches for studying metacommunities: a multi-taxon analysis in Mediterranean and tropical temporary ponds.

Prior research on metacommunities has largely focused on snapshot surveys, often overlooking temporal dynamics. In this study, our aim was to compare the insights obtained from metacommunity analyses based on a spatial approach repeated over time, with a spatio-temporal approach that consolidates all data into a single model. We empirically assessed the influence of temporal variation in the environment and spatial connectivity on the structure of metacommunities in tropical and Mediterranean temporary ponds. Employing a standardized methodology across both regions, we surveyed multiple freshwater taxa in three time periods within the same hydrological year from multiple temporary ponds in each region. To evaluate how environmental, spatial and temporal influences vary between the two approaches, we used nonlinear variation partitioning analyses based on generalized additive models. Overall, this study underscores the importance of adopting spatio-temporal analytics to better understand the processes shaping metacommunities. While the spatial approach suggested that environmental factors had a greater influence, our spatio-temporal analysis revealed that spatial connectivity was the primary driver influencing metacommunity structure in both regions. Temporal effects were equally important as environmental effects, suggesting a significant role of ecological succession in metacommunity structure.

The spatial frequency of climatic conditions affects niche composition and functional diversity of species assemblages: the case of Angiosperms.

Climatic conditions vary in spatial frequency globally. Spatially rare climatic conditions provide fewer suitable environments than common ones and should impose constraints on the types of species present locally and regionally. We used data on 467 North American angiosperms to test the effects of the spatial frequency of climatic conditions on ecological niche specialisation and functional diversity. We predicted that rare climates should favour generalist species that are able to inhabit a broader range of climatic conditions. Our results show that climate frequency filters species that differ in niche breadths and rare environments host species combinations with greater functional diversity. The proposed analytical approaches and hypotheses can be adapted to investigate different aspects of ecological assemblies and their biodiversity. We discuss different mechanisms regarding how spatial frequency of environments can affect niche composition and functional diversity. These should be useful while developing theoretical frameworks for generating a deeper understanding of its underpinnings.

Beyond neutrality: disentangling the effects of species sorting and spurious correlations in community analysis.

The methods of direct gradient analysis and variation partitioning are the most widely used frameworks to evaluate the contributions of species sorting to metacommunity structure. In many cases, however, species are also driven by spatial processes that are independent of environmental heterogeneity (e.g., neutral dynamics). As such, spatial autocorrelation can occur independently in both species (due to limited dispersal) and the environmental data, leading to spurious correlations between species distributions and the spatialized (i.e., spatially autocorrelated) environment. In these cases, the method of variation partitioning may present high Type I error rates (i.e., reject the null hypothesis more often than the pre-established critical level) and inflated estimates regarding the environmental component that is used to estimate the importance of species sorting. In this paper, we (1) demonstrate that metacommunities driven by neutral dynamics (via limited dispersal) alone or in combination with species sorting leads to inflated estimates and Type I error rates when testing for the importance of species sorting; and (2) propose a general and flexible new variation partitioning procedure to adjust for spurious contributions due to spatial autocorrelation from the environmental fraction. We used simulated metacommunity data driven by pure neutral, pure species sorting, and mixed (i.e., neutral + species sorting dynamics) processes to evaluate the performances of our new methodological framework. We also demonstrate the utility of the proposed framework with an empirical plant dataset in which we show that half of the variation initially due to the environment by the standard variation partitioning framework was due to spurious correlations.

A quantitative framework to estimate the relative importance of environment, spatial variation and patch connectivity in driving community composition.

Perhaps the most widely used quantitative approach in metacommunity ecology is the estimation of the importance of local environment vs. spatial structuring using the variation partitioning framework. Contrary to metapopulation models, however, current empirical studies of metacommunity structure using variation partitioning assume a space-for-dispersal substitution due to the lack of analytical frameworks that incorporate patch connectivity predictors of dispersal dynamics. Here, a method is presented that allows estimating the relative importance of environment, spatial variation and patch connectivity in driving community composition variation within metacommunities. The proposed approach is illustrated by a study designed to understand the factors driving the structure of a soft-bottom marine polychaete metacommunity. Using a standard variation partitioning scheme (i.e. where only environmental and spatial predictors are used), only about 13% of the variation in metacommunity structure was explained. With the connectivity set of predictors, the total amount of explained variation increased up to 51% of the variation. These results highlight the importance of considering predictors of patch connectivity rather than just spatial predictors. Given that information on connectivity can be estimated by commonly available data on species distributions for a number of taxa, the framework presented here can be readily applied to past studies as well, facilitating a more robust evaluation of the factors contributing to metacommunity structure.

Deconstructing the relationships between phylogenetic diversity and ecology: a case study on ecosystem functioning.

Recent studies have supported a link between phylogenetic diversity and various ecological properties including ecosystem function. However, such studies typically assume that phylogenetic branches of equivalent length are more or less interchangeable. Here we suggest that there is a need to consider not only branch lengths but also their placement on the phylogeny. We demonstrate how two common indices of network centrality can be used to describe the evolutionary distinctiveness of network elements (nodes and branches) on a phylogeny. If phylogenetic diversity enhances ecosystem function via complementarity and the representation of functional diversity, we would predict a correlation between evolutionary distinctiveness of network elements and their contribution to ecosystem process. In contrast, if one or a few evolutionary innovations play key roles in ecosystem function, the relationship between evolutionary distinctiveness and functional contribution may be weak or absent. We illustrate how network elements associated with high functional contribution can be identified from regressions between phylogenetic diversity and productivity using a well-known empirical data set on plant productivity from the Cedar Creek Long-Term Ecological Research. We find no association between evolutionary distinctiveness and ecosystem functioning, but we are able to identify phylogenetic elements associated with species of known high functional contribution within the Fabaceae. Our perspective provides a useful guide in the search for ecological traits linking diversity and ecosystem function, and suggests a more nuanced consideration of phylogenetic diversity is required in the conservation and biodiversity-ecosystem-function literature.

On the evolution of dispersal via heterogeneity in spatial connectivity.

Dispersal has long been recognized as a mechanism that shapes many observed ecological and evolutionary processes. Thus, understanding the factors that promote its evolution remains a major goal in evolutionary ecology. Landscape connectivity may mediate the trade-off between the forces in favour of dispersal propensity (e.g. kin-competition, local extinction probability) and those against it (e.g. energetic or survival costs of dispersal). It remains, however, an open question how differing degrees of landscape connectivity may select for different dispersal strategies. We implemented an individual-based model to study the evolution of dispersal on landscapes that differed in the variance of connectivity across patches ranging from networks with all patches equally connected to highly heterogeneous networks. The parthenogenetic individuals dispersed based on a flexible logistic function of local abundance. Our results suggest, all else being equal, that landscapes differing in their connectivity patterns will select for different dispersal strategies and that these strategies confer a long-term fitness advantage to individuals at the regional scale. The strength of the selection will, however, vary across network types, being stronger on heterogeneous landscapes compared with the ones where all patches have equal connectivity. Our findings highlight how landscape connectivity can determine the evolution of dispersal strategies, which in turn affects how we think about important ecological dynamics such as metapopulation persistence and range expansion.

Habitat-based polymorphism is common in stream fishes.

Morphological differences (size and shape) across habitats are common in lake fish where differences relate to two dominant contrasting habitats: the pelagic and littoral habitat. Repeated occurrence of littoral and pelagic morphs across multiple populations of several lake fish species has been considered as important evidence that polymorphism is adaptive in these systems. It has been suggested that these habitat-based polymorphic differences are due to the temporal stability of the differences between littoral and pelagic habitats. Although streams are spatially heterogeneous, they are also more temporally dynamic than lakes and it is still an open question whether streams provide the environmental conditions that promote habitat-based polymorphism. We tested whether fish from riffle, run and pool habitats, respectively, differed consistently in their morphology. Our test compared patterns of morphological variation (size and shape) in 10 fish species from the three stream habitat types in 36 separate streams distributed across three watersheds. For most species, body size and shape (after controlling for body size) differed across riffle, run and pool habitats. Unlike many lake species, the nature of these differences was not consistent across species, possibly because these species use these habitat types in different ways. Our results suggest that habitat-based polymorphism is an important feature also in stream fishes despite the fact that streams are temporally variable in contrast to lake systems. Future research is required to assess whether the patterns of habitat-based polymorphism encountered in streams have a genetic basis or they are simply the result of within generation phenotypic plasticity.

The interaction between the spatial distribution of resource patches and population density: consequences for intraspecific growth and morphology.

How individuals within a population distribute themselves across resource patches of varying quality has been an important focus of ecological theory. The ideal free distribution predicts equal fitness amongst individuals in a 1 : 1 ratio with resources, whereas resource defence theory predicts different degrees of monopolization (fitness variance) as a function of temporal and spatial resource clumping and population density. One overlooked landscape characteristic is the spatial distribution of resource patches, altering the equitability of resource accessibility and thereby the effective number of competitors. While much work has investigated the influence of morphology on competitive ability for different resource types, less is known regarding the phenotypic characteristics conferring relative ability for a single resource type, particularly when exploitative competition predominates. Here we used young-of-the-year rainbow trout (Oncorhynchus mykiss) to test whether and how the spatial distribution of resource patches and population density interact to influence the level and variance of individual growth, as well as if functional morphology relates to competitive ability. Feeding trials were conducted within stream channels under three spatial distributions of nine resource patches (distributed, semi-clumped and clumped) at two density levels (9 and 27 individuals). Average trial growth was greater in high-density treatments with no effect of resource distribution. Within-trial growth variance had opposite patterns across resource distributions. Here, variance decreased at low-population, but increased at high-population densities as patches became increasingly clumped as the result of changes in the levels of interference vs. exploitative competition. Within-trial growth was related to both pre- and post-trial morphology where competitive individuals were those with traits associated with swimming capacity and efficiency: larger heads/bodies/caudal fins and less angled pectoral fins. The different degrees of within-population growth variance at the same density level found here, as a function of spatial resource distribution, provide an explanation for the inconsistencies in within-site growth variance and population regulation often noted with regard to density dependence in natural landscapes.

Early growth trajectories affect sexual responsiveness.

The trajectory of an animal's growth in early development has been shown to have long-term effects on a range of life-history traits. Although it is known that individual differences in behaviour may also be related to certain life-history traits, the linkage between early growth or development and individual variation in behaviour has received little attention. We used brief temperature manipulations, independent of food availability, to stimulate compensatory growth in juvenile three-spined sticklebacks Gasterosteus aculeatus. Here, we examine how these manipulated growth trajectories affected the sexual responsiveness of the male fish at the time of sexual maturation, explore associations between reproductive behaviour and investment and lifespan and test whether the perceived time stress (until the onset of the breeding season) influenced such trade-offs. We found a negative impact of growth rate on sexual responsiveness: fish induced (by temperature manipulation) to grow slowest prior to the breeding season were consistently quickest to respond to the presence of a gravid female. This speed of sexual responsiveness was also positively correlated with the rate of development of sexual ornaments and time taken to build a nest. However, after controlling for effects of growth rate, those males that had the greatest sexual responsiveness to females had the shortest lifespan. Moreover, the time available to compensate in size before the onset of the breeding season (time stress) affected the magnitude of these effects. Our results demonstrate that developmental perturbations in early life can influence mating behaviour, with long-term effects on longevity.

Combining the fourth-corner and the RLQ methods for assessing trait responses to environmental variation.

Assessing trait responses to environmental gradients requires the simultaneous analysis of the information contained in three tables: L (species distribution across samples), R (environmental characteristics of samples), and Q (species traits). Among the available methods, the so-called fourth-corner and RLQ methods are two appealing alternatives that provide a direct way to test and estimate trait-nvironment relationships. Both methods are based on the analysis of the fourth-corner matrix, which crosses traits and environmental variables weighted by species abundances. However, they differ greatly in their outputs: RLQ is a multivariate technique that provides ordination scores to summarize the joint structure among the three tables, whereas the fourth-corner method mainly tests for individual trait-environment relationships (i.e., one trait and one environmental variable at a time). Here, we illustrate how the complementarity between these two methods can be exploited to promote new ecological knowledge and to improve the study of trait-environment relationships. After a short description of each method, we apply them to real ecological data to present their different outputs and provide hints about the gain resulting from their combined use.

Heterogeneous dispersal networks to improve biodiversity science.

Dispersal has a key role in shaping spatial patterns of biodiversity, yet its spatial heterogeneity is often overlooked in biodiversity analyses and management strategies. Properly parameterised heterogeneous dispersal networks capture the complex interplay between landscape structure and species-specific dispersal capacities. However, this heterogeneity is recurrently neglected when studying the processes underlying biodiversity variation. To address this gap, we introduce a conceptual framework detailing the fundamental processes driving dispersal heterogeneity and its effects on biodiversity dynamics. We propose methods to parameterise heterogeneous dispersal networks, facilitating their integration into commonly used quantitative frameworks for biodiversity analyses. By considering the architecture of heterogeneous dispersal networks, we demonstrate their critical role in guiding biodiversity management strategies.

The biotic and abiotic contexts of ecological selection mediate the dominance of distinct dispersal strategies in competitive metacommunities.

While the influence of dispersal on ecological selection is the subject of intense research, we still lack a thorough understanding of how ecological selection operates to favour distinct dispersal strategies in metacommunities. To address this issue, we developed a model framework in which species with distinct quantitative dispersal traits that govern the three stages of dispersal-departure, movement and settlement-compete under different ecological contexts. The model identified three primary dispersal strategies (referred to as nomadic, homebody and habitat-sorting) that consistently dominated metacommunities owing to the interplay of spatiotemporal environmental variation and different types of competitive interactions. We outlined the key characteristics of each strategy and formulated theoretical predictions regarding the abiotic and biotic conditions under which each strategy is more likely to prevail in metacommunities. By presenting our results as relationships between dispersal traits and well-known ecological gradients (e.g. seasonality), we were able to contrast our theoretical findings with previous empirical research. Our model demonstrates how landscape environmental characteristics and competitive interactions at the intra- and interspecific levels can interact to favour distinct multivariate and context-dependent dispersal strategies in metacommunities. This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.

Inferring explicit weighted consensus networks to represent alternative evolutionary histories.

BACKGROUND: The advent of molecular biology techniques and constant increase in availability of genetic material have triggered the development of many phylogenetic tree inference methods. However, several reticulate evolution processes, such as horizontal gene transfer and hybridization, have been shown to blur the species evolutionary history by causing discordance among phylogenies inferred from different genes. METHODS: To tackle this problem, we hereby describe a new method for inferring and representing alternative (reticulate) evolutionary histories of species as an explicit weighted consensus network which can be constructed from a collection of gene trees with or without prior knowledge of the species phylogeny. RESULTS: We provide a way of building a weighted phylogenetic network for each of the following reticulation mechanisms: diploid hybridization, intragenic recombination and complete or partial horizontal gene transfer. We successfully tested our method on some synthetic and real datasets to infer the above-mentioned evolutionary events which may have influenced the evolution of many species. CONCLUSIONS: Our weighted consensus network inference method allows one to infer, visualize and validate statistically major conflicting signals induced by the mechanisms of reticulate evolution. The results provided by the new method can be used to represent the inferred conflicting signals by means of explicit and easy-to-interpret phylogenetic networks.

A community of metacommunities: exploring patterns in species distributions across large geographical areas.

Ecological communities show extremely complex patterns of variation in space, and quantifying the relative importance of spatial and environmental factors underpinning patterns of species distributions is one of the main goals of community ecology. Although we have accumulated good knowledge about the processes driving species distributions within metacommunities, we have few insights about whether (and how) environmental and spatial features can actually generate consistent species distributional patterns across multiple metacommunities. In this paper we applied the elements of metacommunity structure (EMS) framework to identify and classify metacommunities according to multiple but discrete patterns of species distributions. Given that each pattern has unique underlying structuring mechanisms, exploring and comparing such patterns across multiple metacommunities spanning large geographical areas provides a way to test the existence of general principles underlying species distributions within metacommunities. In this study, we applied the EMS framework into a data set containing about 9000 lakes distributed across 85 fish metacommunities across Ontario, Canada, and estimated the relative importance of local and spatial factors in explaining their distributional patterns. Nested and Clementsian gradients were the patterns that fitted most metacommunities; nested metacommunities were distributed throughout the province, while Clementsian gradient metacommunities were concentrated in the southeastern region. Sixty-five percent of nested metacommunities were located in low-energy watersheds (i.e., colder climate and shorter growing season), whereas metacommunities representing Clementsian gradients were present in high-energy watersheds (i.e., relatively warmer climate and longer growing season). Taken together, our results reveal that the environmental and spatial properties in which metacommunities are embedded are at least partially responsible for their species distributional patterns.

Inconsistent response of taxonomic groups to space and environment in mediterranean and tropical pond metacommunities.

The metacommunity concept provides a theoretical framework that aims at explaining organism distributions by a combination of environmental filtering, dispersal, and drift. However, few works have attempted a multitaxon approach and even fewer have compared two distant biogeographical regions using the same methodology. We tested the expectation that temperate (mediterranean-climate) pond metacommunities would be more influenced by environmental and spatial processes than tropical ones, because of stronger environmental gradients and a greater isolation of waterbodies. However, the pattern should be different among groups of organisms depending on their dispersal abilities. We surveyed 30 tropical and 32 mediterranean temporary ponds from Costa Rica and Spain, respectively, and obtained data on 49 environmental variables. We characterized the biological communities of bacteria and archaea (from the water column and the sediments), phytoplankton, zooplankton, benthic invertebrates, amphibians and birds, and estimated the relative role of space and environment on metacommunity organization for each group and region, by means of variation partitioning using generalized additive models. Purely environmental effects were important in both tropical and mediterranean ponds, but stronger in the latter, probably due to their larger limnological heterogeneity. Spatially correlated environment and pure spatial effects were greater in the tropics, related to higher climatic heterogeneity and dispersal processes (e.g., restriction, surplus) acting at different scales. The variability between taxonomic groups in the contribution of spatial and environmental factors to metacommunity variation was very wide, but higher in active, compared with passive, dispersers. Higher environmental effects were observed in mediterranean passive dispersers, and higher spatial effects in tropical passive dispersers. The unexplained variation was larger in the tropical setting, suggesting a higher role for stochastic processes, unmeasured environmental factors, or biotic interactions in the tropics, although this difference affected some actively dispersing groups (insects and birds) more than passive dispersers. These results, despite our limitations in comparing only two regions, provide support, for a wide variety of aquatic organisms, for the classic view of stronger abiotic niche constraints in temperate areas compared with the tropics. The heterogeneous response of taxonomic groups between regions also points to a stronger influence of regional context than organism adaptations on metacommunity organization.

Reproducibility in ecology and evolution: Minimum standards for data and code.

We call for journals to commit to requiring open data be archived in a format that will be simple and clear for readers to understand and use. If applied consistently, these requirements will allow contributors to be acknowledged for their work through citation of open data, and facilitate scientific progress.

Using directed phylogenetic networks to retrace species dispersal history.

Methods designed for inferring phylogenetic trees have been widely applied to reconstruct biogeographic history. Because traditional phylogenetic methods used in biogeographic reconstruction are based on trees rather than networks, they follow the strict assumption in which dispersal among geographical units have occurred on the basis of single dispersal routes across regions and are, therefore, incapable of modelling multiple alternative dispersal scenarios. The goal of this study is to describe a new method that allows for retracing species dispersal by means of directed phylogenetic networks obtained using a horizontal gene transfer (HGT) detection method as well as to draw parallels between the processes of HGT and biogeographic reconstruction. In our case study, we reconstructed the biogeographic history of the postglacial dispersal of freshwater fishes in the Ontario province of Canada. This case study demonstrated the utility and robustness of the new method, indicating that the most important events were south-to-north dispersal patterns, as one would expect, with secondary faunal interchange among regions. Finally, we showed how our method can be used to explore additional questions regarding the commonalities in dispersal history patterns and phylogenetic similarities among species.

Measuring protected-area isolation and correlations of isolation with land-use intensity and protection status.

Protected areas cover over 12% of the terrestrial surface of Earth, and yet many fail to protect species and ecological processes as originally envisioned. Results of recent studies suggest that a critical reason for this failure is an increasing contrast between the protected lands and the surrounding matrix of often highly altered land cover. We measured the isolation of 114 protected areas distributed worldwide by comparing vegetation-cover heterogeneity inside protected areas with heterogeneity outside the protected areas. We quantified heterogeneity as the contagion of greenness on the basis of NDVI (normalized difference vegetation index) values, for which a higher value of contagion indicates less heterogeneous land cover. We then measured isolation as the difference between mean contagion inside the protected area and mean contagion in 3 buffer areas of increasing distance from the protected-area border. The isolation of protected areas was significantly positive in 110 of the 114 areas, indicating that vegetation cover was consistently more heterogeneous 10-20 km outside protected areas than inside their borders. Unlike previous researchers, we found that protected areas in which low levels of human activity are allowed were more isolated than areas in which high levels are allowed. Our method is a novel way to assess the isolation of protected areas in different environmental contexts and regions.