Intransitive competition and its effects on community functional diversity

Can competitive interactions be inferred from the analysis of community functional diversity patterns? Originally, at the scale of a community, competitive interactions were supposed to generate trait overdispersion patterns due to limiting similarity process. More recently, by highlighting the importance of competitive hierarchies, it has been shown that when only one resource limits species coexistence, competition can also lead to patterns of trait clustering. However, these two expectations (overdispersion and clustering) ignore potential multi‐species indirect competitive interactions, and especially intransitive competition. Indeed, little is yet known about intransitive competition and its influence on community's functional diversity. Here I propose a brief appraisal of empirical evidence for intransitive competition in nature, and an overview of the current understanding of this mechanism and its properties. I then demonstrate with a theoretical model that intransitive competitive interactions can actually generate random‐like functional diversity patterns. The variety of diversity patterns (overdispersion, clustering, randomness) that can emerge from diverse types of competitive interactions makes it difficult to identify the presence of competition in nature, potentially leading to an underestimation of its importance as a structuring force. New methodologies able to capture both simple and complex competition mechanisms are thus urgently needed.     

From richer to poorer: successful invasion by freshwater fishes depends on species richness of donor and recipient basins

Evidence for the theory of biotic resistance is equivocal, with experiments often finding a negative relationship between invasion success and native species richness, and large‐scale comparative studies finding a positive relationship. Biotic resistance derives from local species interactions, yet global and regional studies often analyze data at coarse spatial grains. In addition, differences in competitive environments across regions may confound tests of biotic resistance based solely on native species richness of the invaded community. Using global and regional data sets for fishes in river and stream reaches, we ask two questions: (1) does a negative relationship exist between native and non‐native species richness and (2) do non‐native species originate from higher diversity systems. A negative relationship between native and non‐native species richness in local assemblages was found at the global scale, while regional patterns revealed the opposite trend. At both spatial scales, however, nearly all non‐native species originated from river basins with higher native species richness than the basin of the invaded community. Together, these findings imply that coevolved ecological interactions in species‐rich systems inhibit establishment of generalist non‐native species from less diverse communities. Consideration of both the ecological and evolutionary aspects of community assembly is critical to understanding invasion patterns. Distinct evolutionary histories in different regions strongly influence invasion of intact communities that are relatively unimpacted by human actions, and may explain the conflicting relationship between native and non‐native species richness found at different spatial scales.     

How competitive intransitivity and niche overlap affect spatial coexistence

Competitive intransitivity is mostly considered outside the main body of coexistence theories that rely primarily on the role of niche overlap and differentiation. How the interplay of competitive intransitivity and niche overlap jointly affects species coexistence has received little attention. Here, we consider a rock–paper–scissors competition system where interactions between species can represent the full spectra of transitive–intransitive continuum and niche overlap/differentiation under different levels of competition asymmetry. By comparing results from pair approximation that only considers interference competition between neighbouring cells in spatial lattices, with those under the mean-field assumption, we show that 1) species coexistence under transitive competition is only possible at high niche differentiation; 2) in communities with partial or pure intransitive interactions, high levels of niche overlap are not necessary to beget species extinction; and 3) strong spatial clustering can widen the condition for intransitive loops to facilitate species coexistence. The two mechanisms, competitive intransitivity and niche differentiation, can support species persistence and coexistence, either separately or in combination. Finally, the contribution of intransitive loops to species coexistence can be enhanced by strong local spatial correlations, modulated and maximised by moderate competition asymmetry. Our study, therefore, provides a bridge to link intransitive competition to other generic ecological theories of species coexistence.     

A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

Disturbance, interspecific interaction and diversity in metapopulations

Metapopulation diversity patterns depend on the relations among the timescales of local biological interactions (predation, competition), the rates of dispersal among local populations and the patterns of disturbance. We investigate these relationships using a family of simple non-linear Markov chain models. We consider three models for interspecific competition; if the species are identified with early and late successional species, the models describe the facilitation, inhibition and tolerance models of ecological succession. By adding a third competing species we also compare transitive competitive hierarchies and intransitive competitive networks. Finally, we examine the effects of predation in mediating coexistence among competing prey species. In each model we find circumstances in which biotic or abiotic disturbance can increase both local and regional diversity, but those circumstances depend on the various timescales in the model in ways that arc neither obvious nor trivial.     

Competitive interactions change the pattern of species co‐occurrences under neutral dispersal

Non‐random patterns of species segregation and aggregation within ecological communities are often interpreted as evidence for interspecific interactions. However, it is unclear whether theoretical models can predict such patterns and how environmental factors may modify the effects of species interactions on species co‐occurrence. Here we extend a spatially explicit neutral model by including competitive effects on birth and death probabilities to assess whether competition alone is able to produce non‐random patterns of species co‐occurrence. We show that transitive and intransitive competitive hierarchies alone (in the absence of environmental heterogeneity) are indeed able to generate non‐random patterns with commonly used metrics and null models. Moreover, even weak levels of intransitive competition can increase local species richness. However, there is no simple rule or consistent directional change towards aggregation or segregation caused by competitive interactions. Instead, the spatial pattern depends on both the type of species interaction and the strength of dispersal. We conclude that co‐occurrence analysis alone may not able to identify the underlying processes that generate the patterns.     

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of Apeiba membranacea

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.     

Spatial coexistence of phytoplankton species in ecological timescale

The species diversity of phytoplankton is usually very high in wild aquatic systems, as seen in the paradox of plankton. Coexistence of many competitive phytoplankton species is extremely common in nature. However, experiments and mathematical theories show that interspecific competition often leads to the extinction of most inferior species. Here, we present a lattice version of a multi-species Lotka–Volterra competition model to demonstrate the importance of local interaction. Its mathematical equilibrium is the exclusion of all but one superior species. However, temporal coexistence of many competitive species is possible in an ecological time scale if interactions are local instead of global. This implies that the time scale is elongated many orders when interactions are local. Extremely high species diversity of phytoplankton in aquatic systems may be maintained by spatial coexistence in an ecological time scale.Tatsuo Miyazaki, Kei-ichi Tainaka, and Jin Yoshimura contributed equally to this study.     

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of <Emphasis Type="Italic">Apeiba membranacea</Emphasis>

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.     

Facilitative plant interactions and climate simultaneously drive alpine plant diversity

Interactions among species determine local‐scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse‐plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a ‘safety net’ sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.     

The geographic scaling of biotic interactions

A central tenet of ecology and biogeography is that the broad outlines of species ranges are determined by climate, whereas the effects of biotic interactions are manifested at local scales. While the first proposition is supported by ample evidence, the second is still a matter of controversy. To address this question, we develop a mathematical model that predicts the spatial overlap, i.e. co‐occurrence, between pairs of species subject to all possible types of interactions. We then identify the scale of resolution in which predicted range overlaps are lost. We found that co‐occurrence arising from positive interactions, such as mutualism (+/+) and commensalism (+/0), are manifested across scales. Negative interactions, such as competition (?/?) and amensalism (?/0), generate checkerboard patterns of co‐occurrence that are discernible at finer resolutions but that are lost and increasing scales of resolution. Scale dependence in consumer–resource interactions (+/?) depends on the strength of positive dependencies between species. If the net positive effect is greater than the net negative effect, then interactions scale up similarly to positive interactions. Our results challenge the widely held view that climate alone is sufficient to characterize species distributions at broad scales, but also demonstrate that the spatial signature of competition is unlikely to be discernible beyond local and regional scales.     

Recruitment facilitation can promote coexistence and buffer population growth in metacommunities

Although positive species interactions are ubiquitous in nature, theory has generally focused on the role of negative interactions to explain patterns of species diversity. Here, we incorporate recruitment facilitation, a positive interaction prevalent in marine and terrestrial systems, into a metacommunity framework to assess how the interplay between colonisation, competition and facilitation mediates coexistence. We show that when subordinate species facilitate the recruitment of dominant species, multi-species metacommunities can persist stably even if the colonisation rate of the dominant species is greater than that of the subordinate species. In addition, recruitment facilitation can buffer population growth from changes in colonisation rates, and thus explain the paradoxical mismatch between patterns of abundance and recruitment in marine systems. Overall, our results demonstrate that recruitment facilitation can have profound effects on the assembly, dissolution and regulation of metacommunities by mediating the relative influence of local and regional processes on population abundance and species diversity.     

From competition to facilitation: how tree species respond to neighbourhood diversity

Studies on tree communities have demonstrated that species diversity can enhance forest productivity, but the driving mechanisms at the local neighbourhood level remain poorly understood. Here, we use data from a large‐scale biodiversity experiment with 24 subtropical tree species to show that neighbourhood tree species richness generally promotes individual tree productivity. We found that the underlying mechanisms depend on a focal tree's functional traits: For species with a conservative resource‐use strategy diversity effects were brought about by facilitation, and for species with acquisitive traits by competitive reduction. Moreover, positive diversity effects were strongest under low competition intensity (quantified as the total basal area of neighbours) for acquisitive species, and under high competition intensity for conservative species. Our findings demonstrate that net biodiversity effects in tree communities can vary over small spatial scales, emphasising the need to consider variation in local neighbourhood interactions to better understand effects at the community level.     

Filtering across Spatial Scales: Phylogeny,Biogeography and Community Structure in Bumble Bees

Despite the expansion of phylogenetic community analysis to understand community assembly, few studies have used these methods on mobile organisms and it has been suggested the local scales that are typically considered may be too small to represent the community as perceived by organisms with high mobility. Mobility is believed to allow species to mediate competitive interactions quickly and thus highly mobile species may appear randomly assembled in local communities. At larger scales, however, biogeographical processes could cause communities to be either phylogenetically clustered or even. Using phylogenetic community analysis we examined patterns of relatedness and trait similarity in communities of bumble bees (Bombus) across spatial scales comparing: local communities to regional pools, regional communities to continental pools and the continental community to a global species pool. Species composition and data on tongue lengths, a key foraging trait, were used to test patterns of relatedness and trait similarity across scales. Although expected to exhibit limiting similarity, local communities were clustered both phenotypically and phylogenetically. Larger spatial scales were also found to have more phylogenetic clustering but less trait clustering. While patterns of relatedness in mobile species have previously been suggested to exhibit less structure in local communities and to be less clustered than immobile species, we suggest that mobility may actually allow communities to have more similar species that can simply limit direct competition through mobility.     

Multi-proxy evidence for competition between savanna woody species

Coexistence of trees and grasses in savannas should be possible if competition between the woody and the grassy components is less intense than the competition within each component. Although several studies have investigated competition between trees and grasses, little is known about tree–tree interactions. We used a multi-proxy approach to examine the spatial pattern of Acacia mellifera and other savanna woody species in a semi-arid savanna in South Africa. Spatial analysis of the point patterns of young and reproductively mature shrubs detected decreasing aggregation with size/age over all spatial scales. This indicated the prevalence of competition although the overall spatial shrub pattern was aggregated. In contrast to point pattern statistics that detect changes only when competition has led to the death of the inferior competitor, we also applied methods identifying the competitive effect on sizes of individual trees. Competition should lead to a negative spatial autocorrelation in size, which we observed in half of the studied cases. Quantile regressions show that nearest-neighbour distance increased steeply with combined size of the target shrub and its neighbours indicating strong competitive effects. The medians of the distributions of maximum root lengths of A. mellifera, of the scale of regular patterns, and of negative autocorrelations were not significantly different, suggesting that overlapping root systems mediate competitive interactions. A competitor removal experiment did not lead to increased shrub sizes, which may be due to the limited duration of the experiment. From the nearest neighbour and autocorrelation analyses, we conclude that competition had a strong impact on growth rates of savanna woody species. Competition-induced mortality only becomes obvious when analysing the shift towards less aggregated spatial patterns when shrubs become reproductively mature. As the overall clustered spatial pattern masks the perceptible effect of competition, a time component should always be included in spatial pattern-based inference of competition.     

Inferring macro‐ecological patterns from local presence/absence data

Biodiversity provides support for life, vital provisions, regulating services and has positive cultural impacts. It is therefore important to have accurate methods to measure biodiversity, in order to safeguard it when we discover it to be threatened. For practical reasons, biodiversity is usually measured at fine scales whereas diversity issues (e.g. conservation) interest regional or global scales. Moreover, biodiversity may change across spatial scales. It is therefore a key challenge to be able to translate local information on biodiversity into global patterns. Many databases give no information about the abundances of a species within an area, but only its occurrence in each of the surveyed plots. In this paper, we introduce an analytical framework (implemented in a ready‐to‐use R code) to infer species richness and abundances at large spatial scales in biodiversity‐rich ecosystems when species presence/absence information is available on various scattered samples (i.e. upscaling). This framework is based on the scale‐invariance property of the negative binomial. Our approach allows to infer and link within a unique framework important and well‐known biodiversity patterns of ecological theory, such as the species accumulation curve (SAC) and the relative species abundance (RSA) as well as a new emergent pattern, which is the relative species occupancy (RSO). Our estimates are robust and accurate, as confirmed by tests performed on both in silico‐generated and real forests. We demonstrate the accuracy of our predictions using data from two well‐studied forest stands. Moreover, we compared our results with other popular methods proposed in the literature to infer species richness from presence to absence data and we showed that our framework gives better estimates. It has thus important applications to biodiversity research and conservation practice.     

Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales

It is widely accepted that species diversity is contingent upon the spatial scale used to analyze patterns and processes. Recent studies using coarse sampling grains over large extents have contributed much to our understanding of factors driving global diversity patterns. This advance is largely unmatched on the level of local to landscape scales despite being critical for our understanding of functional relationships across spatial scales. In our study on West African bat assemblages we employed a spatially explicit and nested design covering local to regional scales. Specifically, we analyzed diversity patterns in two contrasting, largely undisturbed landscapes, comprising a rainforest area and a forest‐savanna mosaic in Ivory Coast, West Africa. We employed additive partitioning, rarefaction, and species richness estimation to show that bat diversity increased significantly with habitat heterogeneity on the landscape scale through the effects of beta diversity. Within the extent of our study areas, habitat type rather than geographic distance explained assemblage composition across spatial scales. Null models showed structure of functional groups to be partly filtered on local scales through the effects of vegetation density while on the landscape scale both assemblages represented random draws from regional species pools. We present a mixture model that combines the effects of habitat heterogeneity and complexity on species richness along a biome transect, predicting a unimodal rather than a monotonic relationship with environmental variables related to water. The bat assemblages of our study by far exceed previous figures of species richness in Africa, and refute the notion of low species richness of Afrotropical bat assemblages, which appears to be based largely on sampling biases. Biome transitions should receive increased attention in conservation strategies aiming at the maintenance of ecological and evolutionary processes.     

Forcing by rare species and intransitive loops creates distinct bouts of extinction events conditioned by spatial pattern in competition communities

Using a spatially explicit multiple species competition framework, the pattern of extinction for an effect/response competition cellular automata model is presented. It is found that, for a particular matrix with a negative row/column sum covariance, (1) there are two phases of transient behavior, presented in the framework of graph partitioning, and (2) the ultimate behavior of the system in space can be understood as mainly the operation of three distinct intransitive competitive loops. Of particular interest is the effect that a set of initially excluded species have on the final outcome of spatial pattern formation. Even though initial exclusion happens quite rapidly, and it would seem that species excluded early could not have an effect on the eventual outcomes of competition, there is a clear statistical effect, contributing to the extant literature suggesting that rare or seemingly inconsequential species can effect the behavior of the common and obvious ones.     

Ecology drives the worldwide distribution of human diseases

Identifying the factors underlying the origin and maintenance of the latitudinal diversity gradient is a central problem in ecology, but no consensus has emerged on which processes might generate this broad pattern. Interestingly, the vast majority of studies exploring the gradient have focused on free-living organisms, ignoring parasitic and infectious disease (PID) species. Here, we address the influence of environmental factors on the biological diversity of human pathogens and their global spatial organization. Using generalized linear multivariate models and Monte Carlo simulations, we conducted a series of comparative analyses to test the hypothesis that human PIDs exhibit the same global patterns of distribution as other taxonomic groups. We found a significant negative relationship between latitude and PID species richness, and a nested spatial organization, i.e., the accumulation of PID species with latitude, over large spatial scales. Additionally, our results show that climatic factors are of primary importance in explaining the link between latitude and the spatial pattern of human pathogens. Based on our findings, we propose that the global latitudinal species diversity gradient might be generated in large part by biotic interactions, providing strong support for the idea that current estimates of species diversity are substantially underestimated. When parasites and pathogens are included, estimates of total species diversity may increase by more than an order of magnitude.     

How specialised must natural enemies be to facilitate coexistence among plants?

The Janzen‐Connell hypothesis proposes that plant interactions with host‐specific antagonists can impair the fitness of locally abundant species and thereby facilitate coexistence. However, insects and pathogens that associate with multiple hosts may mediate exclusion rather than coexistence. We employ a simulation model to examine the effect of enemy host breadth on plant species richness and defence community structure, and to assess expected diversity maintenance in example systems. Only models in which plant enemy similarity declines rapidly with defence similarity support greater species richness than models of neutral drift. In contrast, a wide range of enemy host breadths result in spatial dispersion of defence traits, at both landscape and local scales, indicating that enemy‐mediated competition may increase defence‐trait diversity without enhancing species richness. Nevertheless, insect and pathogen host associations in Panama and Papua New Guinea demonstrate a potential to enhance plant species richness and defence‐trait diversity comparable to strictly specialised enemies.