Spatial variation and density-dependent dispersal in competitive coexistence

It is well known that dispersal from localities favourable to a species' growth and reproduction (sources) can prevent competitive exclusion in unfavourable localities (sinks). What is perhaps less well known is that too much emigration can undermine the viability of sources and cause regional competitive exclusion. Here, I investigate two biological mechanisms that reduce the cost of dispersal to source communities. The first involves increasing the spatial variation in the strength of competition such that sources can withstand high rates of emigration; the second involves reducing emigration from sources via density-dependent dispersal. I compare how different forms of spatial variation and modes of dispersal influence source viability, and hence source-sink coexistence, under dominance and pre-emptive competition. A key finding is that, while spatial variation substantially reduces dispersal costs under both types of competition, density-dependent dispersal does so only under dominance competition. For instance, when spatial variation in the strength of competition is high, coexistence is possible (regardless of the type of competition) even when sources experience high emigration rates; when spatial variation is low, coexistence is restricted even under low emigration rates. Under dominance competition, density-dependent dispersal has a strong effect on coexistence. For instance, when the emigration rate increases with density at an accelerating rate (Type III density-dependent dispersal), coexistence is possible even when spatial variation is quite low; when the emigration rate increases with density at a decelerating rate (Type II density-dependent dispersal), coexistence is restricted even when spatial variation is quite high. Under pre-emptive competition, density-dependent dispersal has only a marginal effect on coexistence. Thus, the diversity-reducing effects of high dispersal rates persist under pre-emptive competition even when dispersal is density dependent, but can be significantly mitigated under dominance competition if density-dependent dispersal is Type III rather than Type II. These results lead to testable predictions about source-sink coexistence under different regimes of competition, spatial variation and dispersal. They identify situations in which density-independent dispersal provides a reasonable approximation to species' dispersal patterns, and those under which consideration of density-dependent dispersal is crucial to predicting long-term coexistence.     

Source-sink dynamics within a plant population: the impact of substrate and herbivory on palm demography

Site factors have frequently been shown to affect survival, growth, and reproduction in plant populations. The source-sink concept proposed by Pulliam is one way of integrating this spatial demographic variation into population models. Source-sink models describe a population where propagules from “source” habitats sustain less productive “sink” areas. We adapted this concept to model the population dynamics of the understory palm Chamaedorea radicalis on two substrates, rock outcrops and forest floor. In our model, sources and sinks correspond to fine-scale demographic structure within the population, rather than spatially discrete subpopulations as described in the Pulliam model. We constructed a stage-structured population matrix model that integrates the site-specific demography of individuals across two habitats types that are linked by migration. We then parameterized this model with field data from C. radicalis. To address whether observed differences in palm demography between rock outcrops and the forest floor were due to natural variation between microsites or due to differences in browsing intensity from free range livestock, we parameterized separate models based on the substrate-specific demography of protected, non-browsed palms and of palms exposed to burro browse. Results showed that herbivory reduced survival and fecundity on the forest floor, which in the absence of seed migration resulted in a projected decline of forest floor palms (sinks). However with seed dispersal, palms persisted and total population growth (both substrates) was projected to be positive, indicating that seed dispersal from non-browsed palms on rock outcrops (sources) was sufficient to sustain C. radicalis on the forest floor.     

Patterns of dispersal and dynamics among habitat patches varying in quality

Both source-sink theory and extensions of optimal foraging theory ("balanced dispersal" theory) address dispersal and population dynamics in landscapes where habitat patches vary in quality. However, studying dispersal mechanisms empirically has proven difficult, and dispersal is rarely tied back to long-term spatial dynamics. We used a manipulable laboratory system consisting of bacteria and protozoa to investigate the ability of source-sink and optimal foraging theories to explain both dispersal and emergent spatial dynamics. Consistent with source-sink models and contrary to balanced dispersal models, there was a consistent net flux of protist individuals from high to low resource patches. However, unlike the simplest source-sink models, intermediate rates of dispersal led to highest abundances in low resource patches. Side experiments found strong density dependence in local population dynamics and differences in average protist body size in high and low resource patches. Parameterization and analysis of a two-patch model showed that high migration from high to low resource patches could have depressed population density in low resource patches, creating pseudosinks. The movement of individuals and biomass from sources to sinks (a form of ecosystem subsidy) resulted in the convergence of body size and population densities in sources and sinks. Our results indicate a need to carefully consider movement patterns and interaction with local dynamics in potential source-sink systems.     

The role of density-dependent dispersal in source-sink dynamics

I investigate two aspects of source-sink theory that have hitherto received little attention: density-dependent dispersal and the cost of dispersal to sources. The cost arises because emigration reduces the per capita growth rate of sources, thus predisposing them to extinction. I show that source-sink persistence depends critically on the interplay between these two factors. When the emigration rate increases with abundance at an accelerating rate, dispersal costs to sources is the lowest and risk of source-sink extinction the least. When the emigration rate increases with abundance at a decelerating rate, dispersal costs to sources is the highest and the risk of source-sink extinction the greatest. Density-independent emigration has an intermediate effect. Thus, density-dependent dispersal per se does not increase or decrease source-sink persistence relative to density-independent dispersal. The exact mode of dispersal is crucial. A key point to appreciate is that these effects of dispersal on source-sink extinction arise from the temporal density-dependence that dispersal induces in the per capita growth rates of source and sink populations. Temporal density-dependence due to dispersal is beneficial at low abundances because it rescues sinks from extinction, and detrimental at high abundances because it drives otherwise viable sources to extinction. These results are robust to the nature of population dynamics in the sink, whether exponential or logistic. They provide a means of assessing the relative costs and benefits of preserving sink habitats given three biological parameters.     

Demographic stochasticity alters expected outcomes in experimental and simulated non‐neutral communities

Theory has shown that the effects of demographic stochasticity on communities may depend on the magnitude of fitness differences between species. In particular, it has been suggested that demographic stochasticity has the potential to significantly alter competitive outcomes when fitness differences are small (nearly neutral), but that it has negligible effects when fitness differences are large (highly non‐neutral). Here we test such theory experimentally and extend it to examine how demographic stochasticity affects exclusion frequency and mean densities of consumers in simple, but non‐neutral, consumer–resource communities. We used experimental microcosms of protists and rotifers feeding on a bacterial resource to test how varying absolute population sizes (a driver of demographic stochasticity) affected the probability of competitive exclusion of the weakest competitor. To explore whether demographic stochasticity could explain our experimental results, and to generalize beyond our experiment, we paired the experiment with a continuous‐time stochastic model of resource competition, which we simulated for 11 different fitness inequalities between competiting consumers. Consistent with theory, in both our experiments and our simulations we found that demographic stochasticity altered competitive outcomes in communities where fitness differences were small. However, we also found that demographic stochasticity alone could affect communities in other ways, even when fitness differences between competitors were large. Specifically, demographic stochasticity altered mean densities of both weak and strong competitors in experimental and simulated communities. These findings highlight how demographic stochasticity can change both competitive outcomes in non‐neutral communities and the processes underlying overall community dynamics.     

Active dispersal is differentially affected by inter‐ and intraspecific competition in closely related nematode species

Competition is one of the main drivers of dispersal, which can be an important mechanism to achieve permanent or temporal coexistence of multiple species. This coexistence can be achieved by a dispersal‐competition tradeoff, spatial store effects or neutral dynamics. Here we test the effect of inter‐ and intraspecific competition on dispersal of four species of the marine nematode species complex Litoditis marina. A previous study in closed microcosms without a possibility for dispersal had demonstrated pronounced interspecific competition, leading to the exclusion of one species. We now investigated whether 1) the dispersal is affected by interspecific interactions, by intraspecific competition (density) or by food availability, 2) the dispersal dynamics influence assemblage composition and can lead to co‐occurrence of the species, and 3) the abiotic environment (here salinity) can affect these dynamics. We show that density is the main driver for dispersal in two of the four species. Dispersal of a third species always started at the same time irrespective of density, whereas in the fourth species interspecific interactions accelerated dispersal. Remarkably, this fourth species was not a strong competitor, suggesting that a dispersal–competition tradeoff does not explain the observed coexistence. Salinity did not alter the timing of dispersal when interspecific interactions were present but did affect assemblage composition. Consequently, spatial store effects may influence coexistence. All four species co‐occurred in fairly stable abundances throughout the present experiment indicating the importance of species specific dispersal strategies for coexistence. Co‐occurrence can be facilitated because competition is postponed or avoided by dispersal. Neutral dynamics also played a role as intra‐ and interspecific competition were of similar importance in three of the four species. We conclude that dispersal is a driver of the coexistence of closely related nematode species, and that population density and interspecific interactions shape these dynamics.     

Dispersal-mediated coexistence of competing predators

Models of metapopulations have often ignored local community dynamics and spatial heterogeneity among patches. However, persistence of a community as a whole depends both on the local interactions and the rates of dispersal between patches. We study a mathematical model of a metacommunity with two consumers exploiting a resource in a habitat of two different patches. They are the exploitative competitors or the competing predators indirectly competing through depletion of the shared resource. We show that they can potentially coexist, even if one species is sufficiently inferior to be driven extinct in both patches in isolation, when these patches are connected through diffusive dispersal. Thus, dispersal can mediate coexistence of competitors, even if both patches are local sinks for one species because of the interactions with the other species. The spatial asynchrony and the competition-colonization trade-off are usual mechanisms to facilitate regional coexistence. However, in our case, two consumers can coexist either in synchronous oscillation between patches or in equilibrium. The higher dispersal rate of the superior prompts rather than suppresses the inferior. Since differences in the carrying capacity between two patches generate flows from the more productive patch to the less productive, loss of the superior by emigration relaxes competition in the former, and depletion of the resource by subsidized consumers decouples the local community in the latter.     

Subsidies mediate interactions between communities across space

Most spatial ecology focuses on how species dispersal affects community dynamics and coexistence. Ecosystems, however, are also commonly connected by flows of resources. We experimentally tested how neighbouring communities indirectly influence each other in absence of dispersal, via resource exchanges. Using two‐patch microcosm meta‐ecosystems, we manipulated community composition and dynamics, by varying separately species key functional traits (autotroph versus heterotroph species and size of consumer species) and trophic structure of aquatic communities (species growing alone or in presence of competitors or predators). We then analysed the effects of species functional traits and trophic structure on communities connected through spatial subsidies in the absence of actual dispersal. Both functional traits and trophic structure strongly affected dynamics across neighbouring communities. Heterotroph communities connected to autotroph neighbours developed better than with heterotroph neighbours, such that coexistence of competitors was determined by the functional traits of the neighbouring community. Densities in autotroph communities were also strikingly higher when receiving subsidies from heterotroph communities compared to their own subsidies when grown in isolated ecosystems. In contrast, communities connected to predator‐dominated ecosystems collapsed, without any direct contact with the predators. Our results demonstrate that because community composition and structure modify the distribution of biomass within a community, they may also affect communities connected through subsidies through quantitative and qualitative changes of detritus flows. This stresses that ecosystem management should account for such interdependencies mediated by spatial subsidies, given that local community alterations cascade across space onto other ecosystems even if species dispersal is completely absent.     

Dispersal behaviour of African wild dogs in Kenya

Dispersal behaviour plays a key role in social organisation, demography and population genetics. We describe dispersal behaviour in a population of African wild dogs (Lycaon pictus) in Kenya. Almost all individuals, of both sexes, left their natal packs, with 45 of 46 reproductively active “alpha” individuals acquiring their status through dispersal. Dispersal age, group size and distance did not differ between males and females. However, only females embarked on secondary dispersal, probably reflecting stronger reproductive competition among females than males. When dispersing, GPS-collared wild dogs travelled further than when resident, both in daylight and by night, following routes an order of magnitude longer than the straight-line distance covered. Dispersers experienced a daily mortality risk three times that experienced by adults in resident packs. The detailed movement data provided by GPS-collars helped to reconcile differences between dispersal patterns reported previously from other wild dog populations. However, the dispersal patterns observed at this and other sites contrast with those assumed in published demographic models for this endangered species. Given the central role of dispersal in demography, models of wild dog population dynamics need to be updated to account for improved understanding of dispersal processes.     

Spatial heterogeneity, source-sink dynamics, and the local coexistence of competing species

Patch occupancy theory predicts that a trade-off between competition and dispersal should lead to regional coexistence of competing species. Empirical investigations, however, find local coexistence of superior and inferior competitors, an outcome that cannot be explained within the patch occupancy framework because of the decoupling of local and spatial dynamics. We develop two-patch metapopulation models that explicitly consider the interaction between competition and dispersal. We show that a dispersal-competition trade-off can lead to local coexistence provided the inferior competitor is superior at colonizing empty patches as well as immigrating among occupied patches. Immigration from patches that the superior competitor cannot colonize rescues the inferior competitor from extinction in patches that both species colonize. Too much immigration, however, can be detrimental to coexistence. When competitive asymmetry between species is high, local coexistence is possible only if the dispersal rate of the inferior competitor occurs below a critical threshold. If competing species have comparable colonization abilities and the environment is otherwise spatially homogeneous, a superior ability to immigrate among occupied patches cannot prevent exclusion of the inferior competitor. If, however, biotic or abiotic factors create spatial heterogeneity in competitive rankings across the landscape, local coexistence can occur even in the absence of a dispersal-competition trade-off. In fact, coexistence requires that the dispersal rate of the overall inferior competitor not exceed a critical threshold. Explicit consideration of how dispersal modifies local competitive interactions shifts the focus from the patch occupancy approach with its emphasis on extinction-colonization dynamics to the realm of source-sink dynamics. The key to coexistence in this framework is spatial variance in fitness. Unlike in the patch occupancy framework, high rates of dispersal can undermine coexistence, and hence diversity, by reducing spatial variance in fitness.     

Intra- and interspecific density-dependent dispersal in an aquatic prey-predator system

1. Dispersal intensity is a key process for the persistence of prey-predator metacommunities. Consequently, knowledge of the ecological mechanisms of dispersal is fundamental to understanding the dynamics of these communities. Dispersal is often considered to occur at a constant per capita rate; however, some experiments demonstrated that dispersal may be a function of local species density. 2. Here we use aquatic experimental microcosms under controlled conditions to explore intra- and interspecific density-dependent dispersal in two protists, a prey Tetrahymena pyriformis and its predator Dileptus sp. 3. We observed intraspecific density-dependent dispersal for the prey and interspecific density-dependent dispersal for both the prey and the predator. Decreased prey density lead to an increase in predator dispersal, while prey dispersal increased with predator density. 4. Additional experiments suggest that the prey is able to detect its predator through chemical cues and to modify its dispersal behaviour accordingly. 5. Density-dependent dispersal suggests that regional processes depend on local community dynamics. We discuss the potential consequences of density-dependent dispersal on metacommunity dynamics and stability.     

The Dispersal System of a Butterfly: A Test of Source-Sink Theory Suggests the Intermediate-Scale Hypothesis

Theory predicts source-sink dynamics can occur in species with the ideal preemptive distribution but not with the ideal free distribution. Source-sink dynamics can also occur in species with passive dispersal, in which a fixed fraction of the population disperses each generation. However, in nature, dispersal often approximates random diffusion rather than ideal choices or fixed probabilities. Here, I ask which dispersal system occurred in a butterfly (Euphydryas editha) known to have source-sink dynamics. The study used 13 experimental sites, where vacant and occupied habitat patches were juxtaposed. I estimated movement during the flight season and tested hypotheses about the type of dispersal system. Ideal free and ideal preemptive models were rejected because per capita movement rates were density independent. Passive dispersal was rejected because per capita rates were related to patch area and habitat preference. The diffusion model best explained the data because it predicted both the area relationship and an odd feature of the habitat preference: immigration was not higher in preferred habitat; rather, emigration was lower. The diffusion model implied that source-sink dynamics were driven by diffusion from areas of high to low population density. Existing source-sink theory assumes fine-scale patchiness, in which animals have perfect knowledge and ease of mobility. The results from the butterfly suggest that source-sink dynamics arise at coarser spatial scales, where diffusion models apply.     

Balanced dispersal between spatially varying local populations: an alternative to the source-sink model

Analysis of long-term monitoring data on breeding collared flycatchers (Ficedula albicollis Temm.) has revealed equal numbers of immigrations and emigrations between neighboring populations of different sizes. Dispersal patterns were close to patterns simulated under a conditional dispersal and with populations near saturation level. Local growth rates of the 11 sites were computed and did not support the idea that the observed balanced exchanges could be the result of a source-sink system. This is the first empirical evidence for a system of discrete habitat patches with component populations that exist as simultaneous sources and sinks to their neighbors. Dispersal propensities were inversely related to population sizes, which showed little variation in time. These results are consistent with recent modeling of dispersal as an evolutionarily stable strategy, and they demonstrate that dispersal can be an active phenomenon requiring neither the dominance hierarchies nor the temporal instability generally invoked by ecological and population genetic models. We note a parallel to the concept of Ideal Free Distributions and discuss implications for the evolution of dispersal mechanisms in fragmented populations.     

Mechanisms of coexistence in competitive metacommunities

Although there is a large body of theory on spatial competitive coexistence, very little of it involves comparative analyses of alternative mechanisms. We thus have limited knowledge of the conditions under which multiple spatial mechanisms can operate or of emergent properties arising from interactions between mechanisms. Here we present a mathematical framework that allows for comparative analysis of spatial coexistence mechanisms. The basis for comparison is mechanisms operating in spatially homogeneous competitive environments (e.g., life-history trade-offs) versus mechanisms operating in spatially heterogeneous competitive environments (e.g., source-sink dynamics). Our comparative approach leads to several new insights about spatial coexistence. First, we show that spatial variation in the expression of a life-history trade-off leads to a unique regional pattern that cannot be predicted by considering trade-offs or source-sink dynamics alone. This result represents an instance where spatial heterogeneity constrains rather than promotes coexistence, and it illustrates the kind of counterintuitive emergent properties that arise due to interactions between different classes of mechanisms. Second, we clarify the role of dispersal mortality in spatial coexistence. Previous studies have shown that coexistence can be constrained or facilitated by dispersal mortality. Our broader analysis distinguishes situations where dispersal mortality is not necessary for coexistence from those where such mortality is essential for coexistence because it preserves spatial variation in the strength of competition. These results form the basis for two important future directions: evolution of life-history traits in spatially heterogeneous environments and elucidation of the cause and effect relationship(s) between biodiversity and ecosystem functioning.     

Local community size mediates ecological drift and competition in metacommunities

The outcome of competitive interactions is likely to be influenced by both competitive dominance (i.e. niche-based dynamics) and ecological drift (i.e. neutral dynamics governed by demographic stochasticity). However, spatial models of competition rarely consider the joint operation of these two processes. We develop a model based on the original competition-colonization trade-off model that incorporates niche and neutral processes and several realistic facets of ecological dynamics: it allows local competition (i.e. competition within a patch) to occur within communities of a finite size, it allows competitors to vary in the degree of competitive asymmetry, and it includes the role of local migration (i.e. propagule pressure). The model highlights the role of community size, i.e. the number of competitors in the local community, in mediating the relative importance of stochastic and deterministic forces. In metacommunities where local communities are small, ecological drift is substantial enough that strong competitors become effectively neutral, creating abrupt changes in the outcome of competition not predicted by the standard competition-colonization trade-off. Importantly, the model illustrates that, even when other aspects of species interactions (e.g. migration ability, competitive ability) are unchanged, local community size can alter the dynamics of metacommunity persistence. Our work demonstrates that activities which reduce the size of local communities, such as habitat destruction and degradation, effectively compound the extinction debt.     

Determinism in a transient assemblage: the roles of dispersal and local competition

Both dispersal and local competitive ability may determine the outcome of competition among species that cannot coexist locally. I develop a spatially implicit model of two-species competition at a small spatial scale. The model predicts the relative fitness of two competitors based on local reproductive rates and regional dispersal rates in the context of the number, size, and extinction probability of habitat patches in the landscape. I test the predictions of this model experimentally using two genotypes of the bacteriophagous soil nematode Caenorhabditis elegans in patchy microcosms. One genotype has higher fecundity while the other is a better disperser. With such a fecundity-dispersal trade-off between competitors, the model predicts that relative fitness will be affected most by local population size when patches do not go extinct and by the number of patches when there is a high probability of patch extinction. The microcosm experiments support the model predictions. Both approaches suggest that competitive dominance in a patchily distributed transient assemblage will depend upon the architecture and predictability of the environment. These mechanisms, operating at a small scale with high spatial admixture, may be embedded in a larger metacommunity process.     

Competition along a spatial gradient of resource supply: a microbial experimental model

In a set of laboratory experiments, we examined competition for phosphorus between algae and bacteria under various carbon:phosphorus (C:P) supply ratios in spatially homogeneous and heterogeneous microcosms. Experimental results were compared to those predicted by theoretical models of resource competition. In the spatially heterogeneous microcosm, algae that were inferior competitors for P persisted in vessels with high local C:P supply ratios that would cause exclusion in the spatially homogeneous microcosms. Resource competition theory, adapted to this system, provided a starting point for explaining these results. Spatial structure can enhance local diversity because locally inferior competitors are transported from source habitats into sink habitats where they would otherwise be excluded. Such local sources were determined by their resource supply ratios. These results verify the hypothesis that spatial processes enhance local diversity when a system of local habitats is divided into sources and sinks in such a way that each persisting species has at least one source within the system. However, existing theoretical models did not accurately predict distributions of competitor abundance within this experimental system.     

The effect of spatial pattern on community dynamics; a comparison of simulated and field data

The effect of the spatial limits of dispersal and competition on plant community dynamics was studied using Monte-Carlo simulation. The model generates community point patterns, using life-table data, dispersion parameters and radii of competitive effects. These data have been estimated in a field situation, for the 11 most abundant weed species growing on the refuse soil dumps of a strip coal mine. In a simulation experiment, the patterns produced by two versions of the model were compared. The first was based on the field situation as much as possible; the other used the same input parameters except for dispersal, which was randomized in this case. We found considerable differences regarding the temporal changes of species abundances, the realized competitive abilities and the spatial patterns generated by the two versions. An important conclusion of this comparison is that the realized competitive effect (both intra-and interspecific) of a species is dependent not only on constant competition parameters, but on the abundance relations and on the spatial patterns of the competing populations as well. It is concluded that the spatial limits of dispersal and competition may result in the increased persistence of weak competitors, moderate the realized competitive effects of strong species, and shape the spatial coalition structure of the community.     

Attractive sinks, or how individual behavioural decisions determine source–sink dynamics

Gundersen et al . (2001 , Source-sink dynamics: how sinks affect demography of sources. Ecol. Lett. , 4, 14–21.) suggested that sinks can severely affect the demography of populations in source habitats. We propose this is a common result when animals lack cues associated with reduced fitness inside sinks and consequently select habitat inappropriately. These attractive sinks can result either from undetected risks of mortality (as in the experiment of Gundersen et al . 2001 ) or from undetected poor breeding probabilities (due to bioaccumulation of pesticides, for instance). Thus, individual habitat choice is a key process underlying source–sink dynamics.