Movement‐mediated community assembly and coexistence

Organismal movement is ubiquitous and facilitates important ecological mechanisms that drive community and metacommunity composition and hence biodiversity. In most existing ecological theories and models in biodiversity research, movement is represented simplistically, ignoring the behavioural basis of movement and consequently the variation in behaviour at species and individual levels. However, as human endeavours modify climate and land use, the behavioural processes of organisms in response to these changes, including movement, become critical to understanding the resulting biodiversity loss. Here, we draw together research from different subdisciplines in ecology to understand the impact of individual‐level movement processes on community‐level patterns in species composition and coexistence. We join the movement ecology framework with the key concepts from metacommunity theory, community assembly and modern coexistence theory using the idea of micro–macro links, where various aspects of emergent movement behaviour scale up to local and regional patterns in species mobility and mobile‐link‐generated patterns in abiotic and biotic environmental conditions. These in turn influence both individual movement and, at ecological timescales, mechanisms such as dispersal limitation, environmental filtering, and niche partitioning. We conclude by highlighting challenges to and promising future avenues for data generation, data analysis and complementary modelling approaches and provide a brief outlook on how a new behaviour‐based view on movement becomes important in understanding the responses of communities under ongoing environmental change.     

Predicting food‐web structure with metacommunity models

Synthesis Metacommunity theory aims to elucidate the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity research has focused largely on assemblages of competing organisms within a single trophic level. Here, we test the ability of metacommunity models to predict the network structure of the aquatic food web found in the leaves of the northern pitcher plant Sarracenia purpurea. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions play an important role in structuring Sarracenia food webs. Our approach can be applied to any well‐resolved food web for which data are available from multiple locations. The metacommunity framework explores the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity models and empirical studies have focused mostly on assemblages of competing organisms within a single trophic level. Studies of multi‐trophic metacommunities are predominantly restricted to simplified trophic motifs and rarely consider entire food webs. We tested the ability of the patch‐dynamics, species‐sorting, mass‐effects, and neutral metacommunity models, as well as three hybrid models, to reproduce empirical patterns of food web structure and composition in the complex aquatic food web found in the northern pitcher plant Sarracenia purpurea. We used empirical data to determine regional species pools and estimate dispersal probabilities, simulated local food‐web dynamics, dispersed species from regional pools into local food webs at rates based on the assumptions of each metacommunity model, and tested their relative fits to empirical data on food‐web structure. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions were important in structuring Sarracenia food webs. However, differences in dispersal abilities were also important in models that accurately reproduced empirical food web properties. Although the models were tested using pitcher‐plant food webs, the approach we have developed can be applied to any well‐resolved food web for which data are available from multiple locations.     

A simulation‐based approach to understand how metacommunity characteristics influence emergent biodiversity patterns

To understand controls over biodiversity, it is necessary to take a multi‐scale approach to understand how local and regional factors affect the community assembly processes that drive emergent patterns. This need is reflected in the growing use of the metacommunity concept to interpret multi‐scale measures of biodiversity, including metrics derived from diversity partitioning (e.g. α, β and γ diversity) and variation partitioning (e.g. spatial and environmental components of compositional turnover) techniques. However, studies have shown limited success using these metrics to characterize underlying community assembly dynamics. Here we demonstrate how a metacommunity simulation package (MCSim) can be used to evaluate when and how biodiversity metrics can be used to make inferences about metacommunity characteristics. We examined a wide range of parameter settings representing ecologically relevant scenarios. We used artificial neural networks (ANNs) to assess the sensitivity of diversity and variation partitioning metrics (calculated from simulation outcomes) to metacommunity parameter settings. In the scenarios examined in this study, the niche‐neutral gradient strongly influenced most biodiversity metrics, metacommunity size exhibited a marginal influence over some metrics, and dispersal dynamics only affected a subset of variation partitioning outcomes. Variation partitioning response curves along the niche‐neutral gradient were not monotonic; however, simulation outcomes suggest other biodiversity metrics (e.g. dissimilarity saturation) can be used in combination with variation partitioning metrics to make inferences about metacommunity properties. With the growing availability of archived ecological data, we expect future work will apply simulation‐based techniques to better understand links between biodiversity and the metacommunity characteristics that are presumed to control the underlying community assembly processes.     

On the integration of biotic interaction and environmental constraints at the biogeographical scale

Biogeography is primarily concerned with the spatial distribution of biodiversity, including performing scenarios in a changing environment. The efforts deployed to develop species distribution models have resulted in predictive tools, but have mostly remained correlative and have largely ignored biotic interactions. Here we build upon the theory of island biogeography as a first approximation to the assembly dynamics of local communities embedded within a metacommunity context. We include all types of interactions and introduce environmental constraints on colonization and extinction dynamics. We develop a probabilistic framework based on Markov chains and derive probabilities for the realization of species assemblages, rather than single species occurrences. We consider the expected distribution of species richness under different types of ecological interactions. We also illustrate the potential of our framework by studying the interplay between different ecological requirements, interactions and the distribution of biodiversity along an environmental gradient. Our framework supports the idea that the future research in biogeography requires a coherent integration of several ecological concepts into a single theory in order to perform conceptual and methodological innovations, such as the switch from single‐species distribution to community distribution.     

A process‐based metacommunity framework linking local and regional scale community ecology

The metacommunity concept has the potential to integrate local and regional dynamics within a general community ecology framework. To this end, the concept must move beyond the discrete archetypes that have largely defined it (e.g. neutral vs. species sorting) and better incorporate local scale species interactions and coexistence mechanisms. Here, we present a fundamental reconception of the framework that explicitly links local coexistence theory to the spatial processes inherent to metacommunity theory, allowing for a continuous range of competitive community dynamics. These dynamics emerge from the three underlying processes that shape ecological communities: (1) density‐independent responses to abiotic conditions, (2) density‐dependent biotic interactions and (3) dispersal. Stochasticity is incorporated in the demographic realisation of each of these processes. We formalise this framework using a simulation model that explores a wide range of competitive metacommunity dynamics by varying the strength of the underlying processes. Using this model and framework, we show how existing theories, including the traditional metacommunity archetypes, are linked by this common set of processes. We then use the model to generate new hypotheses about how the three processes combine to interactively shape diversity, functioning and stability within metacommunities.     

Towards a multi‐trophic extension of metacommunity ecology

Metacommunity theory provides an understanding of how spatial processes determine the structure and function of communities at local and regional scales. Although metacommunity theory has considered trophic dynamics in the past, it has been performed idiosyncratically with a wide selection of possible dynamics. Trophic metacommunity theory needs a synthesis of a few influential axis to simplify future predictions and tests. We propose an extension of metacommunity ecology that addresses these shortcomings by incorporating variability among trophic levels in ‘spatial use properties’. We define ‘spatial use properties’ as a set of traits (dispersal, migration, foraging and spatial information processing) that set the spatial and temporal scales of organismal movement, and thus scales of interspecific interactions. Progress towards a synthetic predictive framework can be made by (1) documenting patterns of spatial use properties in natural food webs and (2) using theory and experiments to test how trophic structure in spatial use properties affects metacommunity dynamics.     

Extinction debt in source-sink metacommunities

In an increasingly modified world, understanding and predicting the consequences of landscape alteration on biodiversity is a challenge for ecologists. To this end, metacommunity theory has developed to better understand the complexity of local and regional interactions that occur across larger landscapes. While metacommunity ecology has now provided several alternative models of species coexistence at different spatial scales, predictions regarding the consequences of landscape alteration have been done exclusively for the competition-colonization trade off model (CC). In this paper we investigate the effects of landscape perturbation on source-sink metacommunities. We show that habitat destruction perturbs the equilibria among species competitive effects within the metacommunity, driving both direct extinctions and an indirect extinction debt. As in CC models, we found a time lag for extinction following habitat destruction that varied in length depending upon the relative importance of direct and indirect effects. However, in contrast to CC models, we found that the less competitive species are more affected by habitat destruction. The best competitors can sometimes even be positively affected by habitat destruction, which corresponds well with the results of field studies. Our results are complementary to those results found in CC models of metacommunity dynamics. From a conservation perspective, our results illustrate that landscape alteration jeopardizes species coexistence in patchy landscapes through complex indirect effects and delayed extinctions patterns.     

Biodiversity conservation in metacommunity networks: linking pattern and persistence

A central goal of conservation science is to identify the most important habitat patches for maintaining biodiversity on a landscape. Spatial biodiversity patterns are often used for such assessments, and patches that harbor unique diversity are generally prioritized over those with high community similarity to other areas. This places an emphasis on biodiversity representation, but removing a patch can have cascading effects on biodiversity persistence in the remaining ecological communities. Metacommunity theory provides a mechanistic route to the linking of biodiversity patterns on a landscape with the subsequent dynamics of diversity loss after habitat is degraded. Using spatially explicit neutral theory, I focus on the situation where spatial patterns of diversity and similarity are generated by the structure of dispersal networks and not environmental gradients. I find that gains in biodiversity representation are nullified by losses in persistence, and as a result the effects of removing a patch on metacommunity diversity are essentially independent of complementarity or other biodiversity patterns. In this scenario, maximizing protected area and not biodiversity representation is the key to maintaining diversity in the long term. These results highlight the need for a broader understanding of how conservation paradigms perform under different models of metacommunity dynamics.     

Elements of metacommunity structure and community‐environment relationships in stream organisms

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The ecological dynamics of clade diversification and community assembly

Clades diversify in an ecological context, but most macroevolutionary models do not directly encapsulate ecological mechanisms that influence speciation and extinction. A data set of 245 chordate, arthropod, mollusk, and magnoliophyte phylogenies had a majority of clades that showed rapid lineage accumulation early with a slowing more recently, whereas a small but significant minority showed accelerated lineage accumulation in their recent histories. Previous analyses have demonstrated that macroevolutionary birth-death models can replicate the pattern of slowing lineage accumulation only by a strong decrease in speciation rate with increasing species richness and extinction rate held extremely low or absent. In contrast, the metacommunity model presented here could generate the full range of patterns seen in the real phylogenies by simply manipulating the degree of ecological differentiation of new species at the time of speciation. Specifically, the metacommunity model predicts that clades showing decelerating lineage accumulation rates are those that have diversified by ecological modes of speciation, whereas clades showing accelerating lineage accumulation rates are those that have diversified primarily by modes of speciation that generate little or no ecological diversification. A number of testable predictions that integrate data from molecular systematics, community ecology, and biogeography are also discussed.     

Species diversity in neutral metacommunities: a network approach

Biologists seek an understanding of the processes underlying spatial biodiversity patterns. Neutral theory links those patterns to dispersal, speciation and community drift. Here, we advance the spatially explicit neutral model by representing the metacommunity as a network of smaller communities. Analytic theory is presented for a set of equilibrium diversity patterns in networks of communities, facilitating the exploration of parameter space not accessible by simulation. We use this theory to evaluate how the basic properties of a metacommunity – connectivity, size, and speciation rate – determine overall metacommunity γ -diversity, and how that is partitioned into α - and β -components. We find spatial structure can increase γ -diversity relative to a well-mixed model, even when θ is held constant. The magnitude of deviations from the well-mixed model and the partitioning into α - and β -diversity is related to the ratio of migration and speciation rates. γ -diversity scales linearly with metacommunity size even as α - and β -diversity scale nonlinearly with size.     

Unifying ecology and macroevolution with individual‐based theory

A contemporary goal in both ecology and evolutionary biology is to develop theory that transcends the boundary between the two disciplines, to understand phenomena that cannot be explained by either field in isolation. This is challenging because macroevolution typically uses lineage‐based models, whereas ecology often focuses on individual organisms. Here, we develop a new parsimonious individual‐based theory by adding mild selection to the neutral theory of biodiversity. We show that this model generates realistic phylogenies showing a slowdown in diversification and also improves on the ecological predictions of neutral theory by explaining the occurrence of very common species. Moreover, we find the distribution of individual fitness changes over time, with average fitness increasing at a pace that depends positively on community size. Consequently, large communities tend to produce fitter species than smaller communities. These findings have broad implications beyond biodiversity theory, potentially impacting, for example, invasion biology and paleontology.     

Community assembly is a race between immigration and adaptation: eco‐evolutionary interactions across spatial scales

Both ecological and evolutionary mechanisms have been proposed to describe how natural communities become assembled at both regional and biogeographical scales. Yet, these theories have largely been developed in isolation. Here, we unite these separate views and develop an integrated eco‐evolutionary framework of community assembly. We use a simulation approach to explore the factors determining the interplay between ecological and evolutionary mechanisms systematically across spatial scales. Our results suggest that the same set of ecological and evolutionary processes can determine community assembly at both regional and biogeographical scales. We find that the importance of evolution and community monopolization effects, defined as the eco‐evolutionary dynamics that occur when local adaptation of early established immigrants is fast enough to prevent the later immigration of better pre‐adapted species, are not restricted to adaptive radiations on remote islands. They occur at dispersal rates of up to ten individuals per generation, typical for many species at the scale of regional metacommunities. Dispersal capacity largely determines whether ecological species sorting or evolutionary monopolization structure metacommunity diversity and distribution patterns. However, other factors related to the spatial scale at which community assembly processes are acting, such as metacommunity size and the proportion of empty patches, also affect the relative importance of ecology versus evolution. We show that evolution often determines community assembly, and this conclusion is robust to a wide range of assumptions about spatial scale, mode of reproduction, and environmental structure. Moreover, we found that community monopolization effects occur even though species fully pre‐adapted to each habitat are abundant in the metacommunity, a scenario expected a priori to prevent any meaningful effect of evolution. Our results strongly support the idea that the same eco‐evolutionary processes underlie community assembly at regional and biogeographical scales.     

Emerging concepts in temporary‐river ecology

1. Temporary rivers and streams are among the most common and most hydrologically dynamic freshwater ecosystems. The number of temporary rivers and the severity of flow intermittence may be increasing in regions affected by climatic drying trends or water abstraction. Despite their abundance, temporary rivers have been historically neglected by ecologists. A recent increase in temporary‐river research needs to be supported by new models that generate hypotheses and stimulate further research. In this article, we present three conceptual models that address spatial and temporal patterns in temporary‐river biodiversity and biogeochemistry. 2. Temporary rivers are characterised by the repeated onset and cessation of flow, and by complex hydrological dynamics in the longitudinal dimension. Longitudinal dynamics, such as advancing and retreating wetted fronts, hydrological connections and disconnections, and gradients in flow permanence, influence biotic communities and nutrient and organic matter processing. 3. The first conceptual model concerns connectivity between habitat patches. Variable connectivity suggests that the metacommunity and metapopulation concepts are applicable in temporary rivers. We predict that aggregations of local communities in the isolated water bodies of temporary rivers function as metacommunities. These metacommunities may become longitudinally nested due to interspecific differences in dispersal and mortality. The metapopulation concept applies to some temporary river species, but not all. In stable metapopulations, rates of local extinction are balanced by recolonisation. However, extinction and recolonisation in many temporary‐river species are decoupled by frequent disturbances, and populations of these species are usually expanding or contracting. 4. The second conceptual model predicts that large‐scale biodiversity varies as a function of aquatic and terrestrial patch dynamics and water‐level fluctuations. Habitat mosaics in temporary rivers change in composition and configuration in response to inundation and drying, and these changes elicit a range of biotic responses. In the model, aquatic biodiversity initially increases directly with water level due to increasing abundance of aquatic patches. When most of the channel is inundated and most aquatic patches are connected, further increases in aquatic habitat and connectivity cause aquatic biodiversity to decline due to community homogenisation and reduced habitat diversity. The predicted responses of terrestrial biodiversity to changes in water level are the inverse of aquatic biodiversity responses. 5. The third conceptual model represents temporary rivers as longitudinal, punctuated biogeochemical reactors. Advancing fronts carry water, solutes and particulate organic matter downstream; subsequent flow recessions and drying result in deposition of transported material in reserves such as pools and bar tops. Material processing is rapid during inundated periods and slower during dry periods. The efficiency of material processing is predicted to increase with the number of cycles of transport, deposition and processing that occur down the length of a temporary river. 6. We end with a call for conservation and resource management that addresses the unique properties of temporary rivers. Primary objectives for effective temporary river management are preservation or restoration of aquatic‐terrestrial habitat mosaics, preservation or restoration of natural flow intermittence, and identification of flow requirements for highly valued species and processes.     

Time-dependent solutions of the spatially implicit neutral model of biodiversity

Previous research into the neutral theory of biodiversity has focused mainly on equilibrium solutions rather than time-dependent solutions. Understanding the time-dependent solutions is essential for applying neutral theory to ecosystems in which time-dependent processes, such as succession and invasion, are driving the dynamics. Time-dependent solutions also facilitate tests against data that are stronger than those based on static equilibrium patterns. Here I investigate the time-dependent solutions of the classic spatially implicit neutral model, in which a small local community is coupled to a much larger metacommunity through immigration. I present explicit general formulas for the eigenvalues, left eigenvectors and right eigenvectors of the models’s transition matrix. The time-dependent solutions can then be expressed in terms of these eigenvalues and eigenvectors. Some of these results are translated directly from existing results for the classic Moran model of population genetics (the Moran model is equivalent to the spatially implicit neutral model after a reparameterization); others of the results are new. I demonstrate that the asymptotic time-dependent solution corresponding to just these first two eigenvectors can be a good approximation to the full time-dependent solution. I also demonstrate the feasibility of a partial eigendecomposition of the transition matrix, which facilitates direct application of the results to a biologically relevant example in which a newly invading species is initially present in the metacommunity but absent from the local community.     

gen3sis: A general engine for eco-evolutionary simulations of the processes that shape Earth’s biodiversity

Understanding the origins of biodiversity has been an aspiration since the days of early naturalists. The immense complexity of ecological, evolutionary, and spatial processes, however, has made this goal elusive to this day. Computer models serve progress in many scientific fields, but in the fields of macroecology and macroevolution, eco-evolutionary models are comparatively less developed. We present a general, spatially explicit, eco-evolutionary engine with a modular implementation that enables the modeling of multiple macroecological and macroevolutionary processes and feedbacks across representative spatiotemporally dynamic landscapes. Modeled processes can include species’ abiotic tolerances, biotic interactions, dispersal, speciation, and evolution of ecological traits. Commonly observed biodiversity patterns, such as α, β, and γ diversity, species ranges, ecological traits, and phylogenies, emerge as simulations proceed. As an illustration, we examine alternative hypotheses expected to have shaped the latitudinal diversity gradient (LDG) during the Earth’s Cenozoic era. Our exploratory simulations simultaneously produce multiple realistic biodiversity patterns, such as the LDG, current species richness, and range size frequencies, as well as phylogenetic metrics. The model engine is open source and available as an R package, enabling future exploration of various landscapes and biological processes, while outputs can be linked with a variety of empirical biodiversity patterns. This work represents a key toward a numeric, interdisciplinary, and mechanistic understanding of the physical and biological processes that shape Earth’s biodiversity.

This study describes a novel mechanistic engine that predicts a realistic global latitudinal diversity gradient, species richness distribution and phylogenies. This approach is a step towards the interdisciplinary numeric understanding of the physical and biological processes that have shaped Earth’s biodiversity.     

The importance of landscape and habitat properties in explaining instantaneous and long‐term distributions of large branchiopods in subtropical temporary pans

1. The notion that the spatial configuration of habitat patches has to be taken into account to understand the structure and dynamics of ecological communities is the starting point of metacommunity ecology. One way to assess metacommunity structure is to investigate the relative importance of environmental heterogeneity and spatial structure in explaining community patterns over different spatial and temporal scales. 2. We studied metacommunity structure of large branchiopod assemblages characteristic of subtropical temporary pans in SE Zimbabwe using two community data sets: a community snapshot and a long‐term data set covering 4 years. We assessed the relative importance of environmental heterogeneity and dispersal (inferred from patch occupancy patterns) as drivers of community structure. Furthermore, we contrasted metacommunity patterns in pans that occasionally connect to the river (floodplain pans) and pans that lack such connections altogether (endorheic pans) using redundancy models. 3. Echoes of species sorting and dispersal limitation emerge from our data set, suggesting that both local and regional processes contribute to explaining branchiopod assemblages in this system. Relative importance of local and regional factors depended on the type of data set considered. Overall, habitat characteristics that vary in time, such as conductivity, hydroperiod and vegetation cover, best explained the instantaneous species composition observed during a snapshot sampling while long‐term species composition appeared to be linked to more constant intrinsic habitat properties such as river connectivity and spatial location.     

Network isolation and local diversity in neutral metacommunities

Biologists seek an understanding of the biological and environmental factors determining local community diversity. Recent advances in metacommunity ecology, and neutral theory in particular, highlight the importance of dispersal processes interacting with the spatial structure of a landscape for generating spatial patterns and maintaining biodiversity. The relative spatial isolation of a community is traditionally thought to have a large influence on local diversity. However, isolation remains an elusive concept to quantify, particularly in metacommunities with complex spatial structure. We represent the metacommunity as a network of local communities, and use network centrality measures to quantify the isolation of a local community. Using spatially explicit neutral theory, we examine how node position predicts variation in alpha diversity across a metacommunity. We find that diversity increases with node centrality in the network, but only when centrality is measured on a given scale in the network that widens with increasing dispersal rates and narrows with increasing evolutionary rates. More generally, complex biodiversity patterns form only when the underlying geography has structure on this critical scale. This provides a framework for understanding the influence of spatial geographic structure on global biodiversity patterns.     

The community ecology of pathogens: coinfection,coexistence and community composition

Disease and community ecology share conceptual and theoretical lineages, and there has been a resurgence of interest in strengthening links between these fields. Building on recent syntheses focused on the effects of host community composition on single pathogen systems, we examine pathogen (microparasite) communities using a stochastic metacommunity model as a starting point to bridge community and disease ecology perspectives. Such models incorporate the effects of core community processes, such as ecological drift, selection and dispersal, but have not been extended to incorporate host–pathogen interactions, such as immunosuppression or synergistic mortality, that are central to disease ecology. We use a two‐pathogen susceptible‐infected (SI) model to fill these gaps in the metacommunity approach; however, SI models can be intractable for examining species‐diverse, spatially structured systems. By placing disease into a framework developed for community ecology, our synthesis highlights areas ripe for progress, including a theoretical framework that incorporates host dynamics, spatial structuring and evolutionary processes, as well as the data needed to test the predictions of such a model. Our synthesis points the way for this framework and demonstrates that a deeper understanding of pathogen community dynamics will emerge from approaches working at the interface of disease and community ecology.     

Metacommunity‐scale biodiversity regulation and the self‐organised emergence of macroecological patterns

There exist a number of key macroecological patterns whose ubiquity suggests that the spatio‐temporal structure of ecological communities is governed by some universal mechanisms. The nature of these mechanisms, however, remains poorly understood. Here, we probe spatio‐temporal patterns in species richness and community composition using a simple metacommunity assembly model. Despite making no a priori assumptions regarding biotic spatial structure or the distribution of biomass across species, model metacommunities self‐organise to reproduce well‐documented patterns including characteristic species abundance distributions, range size distributions and species area relations. Also in agreement with observations, species richness in our model attains an equilibrium despite continuous species turnover. Crucially, it is in the neighbourhood of the equilibrium that we observe the emergence of these key macroecological patterns. Biodiversity equilibria in models occur due to the onset of ecological structural instability, a population‐dynamical mechanism. This strongly suggests a causal link between local community processes and macroecological phenomena.