Does local competition increase the coexistence of species in intransitive networks?

Competitive intransitivity, a situation in which species' competitive ranks cannot be listed in a strict hierarchy, promotes species coexistence through "enemy's enemy indirect facilitation." Theory suggests that intransitivity-mediated coexistence is enhanced when competitive interactions occur at local spatial scales, although this hypothesis has not been thoroughly tested. Here, we use a lattice model to investigate the effect of local vs. global competition on intransitivity-mediated coexistence across a range of species richness values and levels of intransitivity. Our simulations show that local competition can enhance intransitivity-mediated coexistence in the short-term, yet hinder it in the long-term, when compared to global competition. This occurs because local competition slows species disaggregation, allowing weaker competitors to persist longer in the shifting spatial refuges of intransitive networks, enhancing short-term coexistence. Conversely, our simulations show that, in the long-term, local competition traps disaggregated species in unfavorable areas of the competitive arena, where they are excluded by superior competitors. As a result, in the long-term, global intransitive competition allows a greater number of species to coexist than local intransitive competition.     

Spatially stochastic settlement and the coexistence of benthic marine animals

For sessile organisms, dispersal and recruitment are typically spatially stochastic, but there is little understanding of how this variability scales up to influence processes such as competitive coexistence. Here we argue that coexistence of benthic marine animals is enhanced by stochastic differences between species in the spatial distribution of larval settlement. Differentiation of settlement distributions among competitors results in intraspecifically aggregated settlement, which can reduce overall interspecific competition and increase overall intraspecific competition. We test for the components of this mechanism using a pair of subtidal invertebrates, and we find that the mean interspecific effect of the dominant competitor is substantially reduced by natural settlement variability. Using a simulation parameterized with experimental data, we find that variable settlement could play an important role in long-term coexistence between these species. This mechanism may apply broadly to benthic marine communities, which can be highly diverse and typically exhibit large settlement fluctuation over a range of scales.     

Role of competition in phytoplankton population for the occurrence and control of plankton bloom in the presence of environmental fluctuations

Termination of harmful algal blooms (HABs) and coexistence of phytoplankton–zooplankton populations are of great importance to human health, ecosystem, environment, tourism and fisheries. In this paper, we propose a three component model consisting of non-toxic phytoplankton (NTP), toxin producing phytoplankton (TPP) and zooplankton (Z). The growth of zooplankton species is assume to reduce due to toxic chemicals released by TPP population. We have extended the model proposed by Chattopadhyay et al. [Chattopadhyay, J., Sarkar, R.R., Pal, S., 2004. Mathematical modelling of harmful algal blooms supported by experimental findings. Ecol. Comp. 1, 225–235] by including competition terms between TPP and NTP. We observe the effect of competition factors both in the presence and absence of the environmental fluctuation. From our field as well as model analysis we observe that competition helps in the coexistence of the species, but if the effect of competition is very high on the TPP population, it results in planktonic bloom. It is shown that the coexistence equilibrium loses its stability when the competition coefficient crosses a critical value and resulting Hopf-bifurcation around the positive equilibrium depicting oscillations phenomena of the populations.     

Nonequilibrium coexistence in a competition model with nutrient storage

Resource competition theory predicts that, in equilibrium, the number of coexisting species cannot exceed the number of limiting resources. In some competition models, however, competitive interactions may result in nonequilibrium dynamics, allowing the coexistence of many species on few resources. The relevance of these findings is still unclear, since some assumptions of the underlying models are unrealistic. Most importantly, these models assume that individual growth directly reflects the availability of external resources, whereas real organisms can store resources, thereby decoupling their growth from external fluctuations. Here we study the effects of resource storage by extending the well-known Droop model to the context of multiple species and multiple resources. We demonstrate that the extended Droop model shows virtually the same complex dynamics as models without storage. Depending on the model parameters, one may obtain competitive exclusion, stable equilibrium coexistence, periodic and non-periodic oscillations, and chaos. Again, nonequilibrium dynamics allows for the coexistence of many species on few resources. We discuss our findings in the light of earlier work on resource competition, highlighting the role of luxury consumption, trade-offs in competitive abilities, and ecological stoichiometry.     

Niche and fitness differences relate the maintenance of diversity to ecosystem function

The frequently observed positive correlation between species diversity and community biomass is thought to depend on both the degree of resource partitioning and on competitive dominance between consumers, two properties that are also central to theories of species coexistence. To make an explicit link between theory on the causes and consequences of biodiversity, we define in a precise way two kinds of differences among species: niche differences, which promote coexistence, and relative fitness differences, which promote competitive exclusion. In a classic model of exploitative competition, promoting coexistence by increasing niche differences typically, although not universally, increases the "relative yield total", a measure of diversity's effect on the biomass of competitors. In addition, however, we show that promoting coexistence by decreasing relative fitness differences also increases the relative yield total. Thus, two fundamentally different mechanisms of species coexistence both strengthen the influence of diversity on biomass yield. The model and our analysis also yield insight on the interpretation of experimental diversity manipulations. Specifically, the frequently reported "complementarity effect" appears to give a largely skewed estimate of resource partitioning. Likewise, the "selection effect" does not seem to isolate biomass changes attributable to species composition rather than species richness, as is commonly presumed. We conclude that past inferences about the cause of observed diversity-function relationships may be unreliable, and that new empirical estimates of niche and relative fitness differences are necessary to uncover the ecological mechanisms responsible for diversity-function relationships.     

Environmental variability and allocation trade-offs maintain species diversity in a process-based model of succulent plant communities

Ecological theory suggests that environmental variability can promote coexistence, provided that species occupy differential niches. In this study, we focus on two questions: (1) Do allocation trade-offs provide a sufficient basis for niche differentiation in succulent plant communities? (2) What is the relative importance of different forms of environmental variability on species diversity and community composition? We approach these questions with a generic, individual-based simulation model. In our model, plants compete for water in a spatially explicit environment. Species differ in their size at maturity and in the allocation of carbon to roots, leaves and storage tissue. The model was fully specified with independent literature data. Model output was compared to characteristics of a species-rich community in the semi-arid Richtersveld (South Africa). The model reproduced the coexistence of plants with different sizes at maturity, the dominance of succulent shrubs, and the level of vegetation cover. We analyzed the effects of three forms of environmental variability: (a) temporal fluctuations in precipitation (rain and fog), (b) spatial heterogeneity of water supply due to run-on and run-off processes and (c) ‘rock pockets’ that limit root competition in space. The three types of variability had differential effects on diversity: diversity exhibited a strong hump-shaped response to temporal variation. Spatial variability increased diversity, with the strongest increase occurring at intermediate levels of temporal variability. Finally, rock pockets had the weakest effect, but contributed to diversity by providing refuges for small species, particularly at low temporal variability. The model thus shows that spatio-temporal variation of resource supply can maintain diversity over long time scales even in small systems, as is the case in the Richtersveld succulent communities. Trade-offs in allocation provide the basis for necessary niche differentiation. By describing resource competition between individual plants, our model provides a mechanistic basis for the link from species traits to community composition at given environmental conditions. It thereby contributes to an understanding of the forces shaping plant communities. Such an understanding is critical to reduce the threats environmental change poses to biodiversity and ecosystem services.     

Partitioning the factors of spatial variation in regeneration density of shade-tolerant tree species

Understanding coexistence of highly shade-tolerant tree species is a longstanding challenge for forest ecologists. A conceptual model for the coexistence of sugar maple (Acer saccharum) and American beech (Fagus grandibfolia) has been proposed, based on a low-light survival/high-light growth trade-off, which interacts with soil fertility and small-scale spatiotemporal variation in the environment. In this study, we first tested whether the spatial distribution of seedlings and saplings can be predicted by the spatiotemporal variability of light availability and soil fertility, and second, the manner in which the process of environmental filtering changes with regeneration size. We evaluate the support for this hypothesis relative to the one for a neutral model, i.e., for seed rain density predicted from the distribution of adult trees. To do so, we performed intensive sampling over 86 quadrats (5 x 5 m) in a 0.24-ha plot in a mature maple-beech community in Quebec, Canada. Maple and beech abundance, soil characteristics, light availability, and growth history (used as a proxy for spatiotemporal variation in light availability) were finely measured to model variation in sapling composition across different size classes. Results indicate that the variables selected to model species distribution do effectively change with size, but not as predicted by the conceptual model. Our results show that variability in the environment is not sufficient to differentiate these species' distributions in space. Although species differ in their spatial distribution in the small size classes, they tend to correlate at the larger size class in which recruitment occurs. Overall, the results are not supportive of a model of coexistence based on small-scale variations in the environment. We propose that, at the scale of a local stand, the lack of fit of the model could result from the high similarity of species in the range of environmental conditions encountered, and we suggest that coexistence would be stable only at larger spatial scales at which variability in the environment is greater.     

Interaction rules affect species coexistence in intransitive networks

Intransitive communities, those in which species' abilities cannot be ranked in a hierarchy, have been the focus of theoretical and empirical research, as intransitivity could help explain the maintenance of biodiversity. Here we show that models for intransitive competition embedding slightly different interaction rules can produce opposite patterns. In particular, we find that interactions in which an individual can be outcompeted by its neighbors, but cannot outcompete its neighbors, produce negative frequency dependence that, in turn, promotes coexistence. Whenever the interaction rule is modified toward symmetry (the individual and the neighbors can outcompete each other) the negative frequency dependence vanishes, producing different coexistence levels. Macroscopically, we find that asymmetric interactions yield highest biodiversity if species compete globally, while symmetric interactions favor highest biodiversity if competition takes place locally.     

Evolving metacommunities: toward an evolutionary perspective on metacommunities

The metacommunity framework predicts that local coexistence depends on the outcome of local species interactions and regional migration. In analogous fashion, spatial structure among populations can shape species interactions through evolutionary mechanisms. Yet, most metacommunity theories assume that populations do not evolve. Here, we evaluate how evolution shapes local species coexistence and exclusion within the multiscale and multispecies context embodied by the metacommunity framework. In general, coexistence in joint ecological-evolutionary models requires low to intermediate dispersal rates that can promote maintenance of both regional species and genetic diversity. These conditions support a set of key mechanisms that modify patterns of species coexistence including local adaptation, gene storage effects, genetic rescue effects, spatial genetic subsidies, and metacommunity evolution. Multispecies extensions indicate that correlated selection can further alter the outcome of interspecific interactions depending on the magnitude and direction of correlations and shape of fitness trade-offs. We suggest that an evolving metacommunity perspective has the potential to generate novel predictions about community structure and function by incorporating the genetic and species diversity that characterize natural communities. In adopting such a perspective, we seek to facilitate understanding about the interactions between evolutionary and metacommunity dynamics.     

From single fine roots to a black alder forest ecosystem: How system behaviour emerges from single component activities

Based on empirical findings in a natural black alder ecosystem in Northern Germany we developed an individual based model that integrates components of a black alder ecosystem interacting on different levels of organisation. The factors determining seasonal fine root biomass development of forest ecosystems are not yet fully understood.We used an object oriented model approach to investigate this complex matter for black alder trees. Processes like growth, storage, respiration, transport, nutrient mineralisation and uptake as well as interactions among these factors are described on the level of functionally differentiated plant organs (fine roots, coarse roots, stem, branches, leaves) and soil units. The object structure of the model is determined by spatial relations between plant modules as well as between plant modules and their local environment modules.As results of model application we found that (i) on the organ level, spatio-temporal plasticity of (root) growth allocation is related to spatio-temporal variation of resource availability, (ii) on the plant level, balanced root:shoot growth appears in response to variation of available resources light and nutrients, (iii) on the population level, tree stand development (population structure, self-thinning) resulted from coexistence and competition between plant individuals.For the understanding of the root compartment it seems relevant that the model implementation of local scale fine root dynamics is consistent with a self-organised large scale spatial heterogeneity of fine root activity pattern. On the other hand, fine-root dynamics cannot be explained as a result of autonomous dynamics. A reference to above-ground processes is a necessary condition and the overall plant seems to act as an integrator providing boundary conditions for local activity pattern. At the same time fine-root characteristics are of some importance for properties on hierarchically higher levels, e.g. co-existence in a tree population or element cycling in the ecosystem.As a conclusion, modelling of the spatio-temporal dynamics of tree root systems appears as a paradigmatic example of scale and organisation level integrating processes.     

The community effects of phenotypic and genetic variation within a predator population

Natural populations are heterogeneous mixtures of individuals differing in physiology, morphology, and behavior. Despite the ubiquity of phenotypic variation within natural populations, its effects on the dynamics of ecological communities are not well understood. Here, we use a quantitative genetics framework to examine how phenotypic variation in a predator affects the outcome of apparent competition between its two prey species. Classical apparent competition theory predicts that prey have reciprocally negative effects on each other. The addition of phenotypic trait variation in predation can marginalize these negative effects, mediate coexistence, or generate positive indirect effects between the prey species. Long-term coexistence or facilitation, however, can be preceded by long transients of extinction risk whenever the heritability of phenotypic variation is low. Greater heritability can circumvent these ecological transients but also can generate oscillatory and chaotic dynamics. These dramatic changes in ecological outcomes, in the sign of indirect effects, and in stability suggest that studies which ignore intraspecific trait variation may reach fundamentally incorrect conclusions regarding ecological dynamics.     

Exploring community assembly through an individual-based model for trophic interactions

Traditionally, the dynamics of community assembly has been analyzed by means of deterministic models of differential equations. Despite the theoretical advances provided by such models, they are restricted to questions about community-wide features. The individual-based modeling offers an opportunity to link bionomic features to patterns at the community scale, allowing us to understand how trait-based assembly rules can arise by dynamical processes. The present paper introduces an individual-based model of community assembly, and discusses some of the major advantages and drawbacks of this approach. The model was framed to deal with predation among size-structured populations, incorporating allometric constraints to energetic requirements, movement, life-history features and interaction relationships among individuals. A protocol of assembly procedure is proposed, in which a period of intense species introductions is followed by a period without introductions. The resultant communities did not present any pattern of trait over-dispersion, meaning that the multivariate distances of bionomic features among co-occurring species were neither larger nor more regular than expected in a random collection of species. It suggests a weak influence of interspecific interactions in the model environment and individualistic rules of coexistence, driven mainly by the spatial structure. This highlights that trait over-dispersion and resource partitioning should not be considered a necessary condition for coexistence, even in communities entirely structured by internal processes like predation and competition.     

Human–wildlife coexistence in a changing world

Human–wildlife conflict (HWC) is a key topic in conservation and agricultural research. Decision makers need evidence-based information to design sustainable management plans and policy instruments. However, providing objective decision support can be challenging because realities and perceptions of human–wildlife interactions vary widely between and within rural, urban, and peri-urban areas. Land users who incur costs through wildlife argue that wildlife-related losses should be compensated and that prevention should be subsidized. Supporters of human–wildlife coexistence policies, such as urban-dwelling people, may not face threats to their livelihoods from wildlife. Such spatial heterogeneity in the cost and benefits of living with wildlife is germane in most contemporary societies. This Special Section features contributions on wildlife-induced damages that range from human perspectives (land use, psychology, governance, local attitudes and perceptions, costs and benefits, and HWC and coexistence theory) to ecological perspectives (animal behavior). Building on current literature and articles in this section, we developed a conceptual model to help frame HWC and coexistence dimensions. The framework can be used to determine damage prevention implementation levels and approaches to HWC resolution. Our synthesis revealed that inter- and transdisciplinary approaches and multilevel governance approaches can help stakeholders and institutions implement sustainable management strategies that promote human–wildlife coexistence.     

What enables coexistence in plant communities? Weak versus strong species traits and the role of local processes

Explaining the coexistence of species that basically depend on the same resources has been a brainteaser for generations of ecologists. Different mechanisms have been proposed to facilitate coexistence in plant communities, where space is an important resource. Using a stochastic cellular automaton simulation model we analyze - separately and in combination - the influence of different species traits and processes which alter local competition on the coexistence of plant species over a fixed time horizon. We show that different species traits operate on different time scales in competition. We therefore suggest the concept of weak versus strong traits according to short- or long-term exclusion of species differing in these traits. As a consequence, highly non-linear trade-offs between weak and strong traits can result in communities. Furthermore, we found that trade-offs based on physiological species traits such as plant lifetime, dispersal range and plant growth, did not support broad and long-term coexistence—further processes such as density-dependent mortality and light-dependent colonization were necessary. This suggests that coexistence in plant communities requires (stabilizing) local processes to support the (equalizing) trade-offs in species traits.     

Appreciating Ecological Complexity: Habitat Contours as a Conceptual Landscape Model

Abstract:  Organisms respond to their surroundings at multiple spatial scales, and different organisms respond differently to the same environment. Existing landscape models, such as the "fragmentation model" (or patch-matrix-corridor model) and the "variegation model," can be limited in their ability to explain complex patterns for different species and across multiple scales. An alternative approach is to conceptualize landscapes as overlaid species-specific habitat contour maps. Key characteristics of this approach are that different species may respond differently to the same environmental conditions and at different spatial scales. Although similar approaches are being used in ecological modeling, there is much room for habitat contours as a useful conceptual tool. By providing an alternative view of landscapes, a contour model may stimulate more field investigations stratified on the basis of ecological variables other than human-defined patches and patch boundaries. A conceptual model of habitat contours may also help to communicate ecological complexity to land managers. Finally, by incorporating additional ecological complexity, a conceptual model based on habitat contours may help to bridge the perceived gap between pattern and process in landscape ecology. Habitat contours do not preclude the use of existing landscape models and should be seen as a complementary approach most suited to heterogeneous human-modified landscapes.     

Root plasticity buffers competition among plants: theory meets experimental data

Morphological plasticity is a striking characteristic of plants in natural communities. In the context of foraging behavior particularly, root plasticity has been documented for numerous species. Root plasticity is known to mitigate competitive interactions by reducing the overlap of the individuals' rhizospheres. But despite its obvious effect on resource acquisition, plasticity has been generally neglected in previous empirical and theoretical studies estimating interaction intensity among plants. In this study, we developed a semi-mechanistic model that addresses this shortcoming by introducing the idea of compensatory growth into the classical-zone-of influence (ZOI) and field-of-neighborhood (FON) approaches. The model parameters describing the belowground plastic sphere of influence (PSI) were parameterized using data from an accompanying field experiment. Measurements of the uptake of a stable nutrient analogue at distinct distances to the neighboring plants showed that the study species responded plastically to belowground competition by avoiding overlap of individuals' rhizospheres. An unexpected finding was that the sphere of influence of the study species Bromus hordeaceus could be best described by a unimodal function of distance to the plant's center and not with a continuously decreasing function as commonly assumed. We employed the parameterized model to investigate the interplay between plasticity and two other important factors determining the intensity of competitive interactions: overall plant density and the distribution of individuals in space. The simulation results confirm that the reduction of competition intensity due to morphological plasticity strongly depends on the spatial structure of the competitive environment. We advocate the use of semi-mechanistic simulations that explicitly consider morphological plasticity to improve our mechanistic understanding of plant interactions.     

Analyzing the spatial structure of a Sri Lankan tree species with multiple scales of clustering

Clustering at multiple critical scales may be common for plants since many different factors and processes may cause clustering. This is especially true for tropical rain forests for which theories explaining species coexistence and community structure rest heavily on spatial patterns. We used point pattern analysis to analyze the spatial structure of Shorea congestiflora, a dominant species in a 25-ha forest dynamics plot in a rain forest at Sinharaja World Heritage Site (Sri Lanka), which apparently shows clustering at several scales. We developed cluster processes incorporating two critical scales of clustering for exploring the spatial structure of S. congestiflora and interpret it in relation to factors such as competition, dispersal limitation, recruitment limitation, and Janzen-Connell effects. All size classes showed consistent large-scale clustering with a cluster radius of approximately 25 m. Inside the larger clusters, small-scale clusters with a radius of 8 m were evident for recruits and saplings, weak for intermediates, and disappeared for adults. The pattern of all trees could be divided into two independent patterns: a random pattern (nearest neighbor distance > 8 m) comprising approximately 12% of the trees and a nested double-cluster pattern. This finding suggests two independent recruitment and/or seed dispersal mechanisms. Saplings were several times as abundant as recruits and may accumulate several recruit generations. Recruits were only weakly associated with adults and occupied about half of the large-scale clusters, but saplings almost all. This is consistent with recruitment limitation. For approximately 70% (95%) of all juveniles the nearest adult was less than 26 m away (53 m), suggesting a dispersal limitation that may also be related to the critical large-scale clustering. Our example illustrates the manner in which the use of a specific and complex null hypothesis of spatial structure in point pattern analysis can help us better understand the biology of a species and generate specific hypotheses to be further investigated in the field.     

Coexistence of Fisheries with River Dolphin Conservation

Abstract: Freshwater biodiversity conservation is generally perceived to conflict with human use and extraction (e.g., fisheries). Overexploited fisheries upset the balance between local economic needs and endangered species’ conservation. We investigated resource competition between fisheries and Ganges river dolphins (Platanista gangetica gangetica) in a human‐dominated river system in India to assess the potential for their coexistence. We surveyed a 65‐km stretch of the lower Ganga River to assess habitat use by dolphins (encounter rates) and fishing activity (habitat preferences of fishers, intensity of net and boat use). Dolphin abundance in the main channel increased from 179 (SE 7) (mid dry season) to 270 (SE 8) (peak dry season), probably as a result of immigration from upstream tributaries. Dolphins preferred river channels with muddy, rocky substrates, and deep midchannel waters. These areas overlapped considerably with fishing areas. Sites with 2–6 boats/km (moderately fished) were more preferred by dolphins than sites with 8–55 boats/km (heavily fished). Estimated spatial (85%) and prey–resource overlap (75%) between fisheries and dolphins (chiefly predators of small fish) suggests a high level of competition between the two groups. A decrease in abundance of larger fish, indicated by the fact that small fish comprised 74% of the total caught, may have intensified the present competition. Dolphins seem resilient to changes in fish community structure and may persist in overfished rivers. Regulated fishing in dolphin hotspots and maintenance of adequate dry season flows can sustain dolphins in tributaries and reduce competition in the main river. Fish‐stock restoration and management, effective monitoring, curbing destructive fishing practices, secure tenure rights, and provision of alternative livelihoods for fishers may help reconcile conservation and local needs in overexploited river systems.     

Spatial arrangement of competitors influences coexistence of reef-building corals

Spatial aggregation among strong competitors has been identified as a putative mechanism promoting the coexistence of weak competitors in intensely competitive communities. With notable exceptions in plant communities, few investigators have tested this hypothesis experimentally. In this study, we manipulated the spatial arrangement of corals to test whether within-patch aggregation of a strong coral competitor enhances the success of a weaker coral competitor. Corals grown in simple aggregated arrangements, where the number and type of competitors were held constant, grew almost twice as much as those in non-aggregated arrangements. These growth results suggest that species coexistence is promoted by aggregation within competitive neighborhoods. Thus spatial aggregation may be one of several important mechanisms contributing to the persistence of weak competitors and species coexistence on coral reefs.