Evolution of dispersal in explicitly spatial metacommunities

We apply an evolutionary game theoretic approach to the evolution of dispersal in explicitly spatial metacommunities, using a flexible parametric class of dispersal kernels, namely 2Dt kernels, and study the resulting evolutionary dynamics and outcomes. We observe strong selective pressure on mean dispersal distance (i.e., the first moment), and weaker, but significant, one on the shape of dispersal kernel (i.e., higher moments). We investigate the effects of landscape topology and spatial heterogeneity on the resulting ‘optimal’ dispersal kernels. The shape—importantly the tail structure—and stability of evolutionarily optimal dispersal strategies are strongly affected by landscape topology or connectivity. Specifically, the results suggest that the optimal dispersal kernels in the river network topology have heavier tails and are stable, while those in the direct topology, where organisms are allowed to travel directly from one location to another, have relatively thin tails and may be unstable. We also find that habitat spatial heterogeneity enables coexistence and controls spatial distribution of distinct groups of dispersal strategies and that alteration in topology alone may not be sufficient to change such coexistence. This work provides a tool to translate environmental changes such as global climate change and human intervention into changes in dispersal behavior, which in turn may lead to important alterations of biodiversity and biological invasion patterns.     

Hierarchical zooplankton metacommunities: distinguishing between high and limiting dispersal mechanisms

Theoretical development in the field of community ecology needs ground proofing with empirical tests. In addition, these tests need to be continuously updated. Cottenie (2005) linked observed metacommunities to theoretical models based on whether environmental and/or spatial effects in the observed metacommunity significantly explain community structure. However, a species-sorting metacommunity with high dispersal and one with limited dispersal cannot be distinguished in this manner; both produce significant environmental and spatial effects. In the present study, we demonstrate a solution to this problem using a zooplankton rock pool metacommunity in Churchill, Manitoba, sampled in August 2006. We established a hierarchy of metacommunities in the Churchill rock bluff system—a large, across-bluff metacommunity, and small, within-bluff metacommunities. Using this spatial hierarchy, it is possible to determine the zooplankton dispersal capability in the rock bluff system and hence to link the metacommunity to its corresponding model. We found the zooplankton rock bluff system to exhibit limited dispersal, meaning that spatial effects were significant at the across-bluff scale, but depending on the bluff, were significant or insignificant at the within-bluff scale. Environmental effects were significant at both scales. This study demonstrates a novel way to determine dispersal capabilities in species that are cryptic dispersers, and to successfully link observed metacommunities with theoretical models. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: S. I. Dodson     

Synchrony of spatial populations induced by colored environmental noise and dispersal

Spatial synchrony of oscillating populations has been observed in many ecological systems, and its influences and causes have attracted the interest of ecologists. Spatially correlated environmental noises, dispersal, and trophic interactions have been considered as the causes of spatial synchrony. In this study, we develop a spatially structured population model, which is described by coupled-map lattices and incorporates both dispersal and colored environmental noise. A method for generating time series with desired spatial correlation and color is introduced. Then, we use these generated time series to analyze the influence of noise color on synchrony in population dynamics. The noise color refers to the temporal correlation in the time series data of the noise, and is expressed as the degree of (first-order) autocorrelation for autoregressive noise. Patterns of spatial synchrony are considered for stable, periodic and chaotic population dynamics. Numerical simulations verify that environmental noise color has a major influence on the level of synchrony, which depends strongly on how noise is introduced into the model. Furthermore, the influence of noise color also depends on patterns of dispersal between local populations. In addition, the desynchronizing effect of reddened noise is always weaker than that of white noise. From our results, we notice that the role of reddened environmental noise on spatial synchrony should be treated carefully and cautiously, especially for the spatially structured populations linked by dispersal.     

Top-down control in a patchy environment: revisiting the stabilizing role of food-dependent predator dispersal

In this paper, we revisit the stabilizing role that predator dispersal and aggregation have in the top-down regulation of predator-prey systems in a heterogeneous environment. We consider an environment consisting of sites interconnected by dispersal, and propose a novel mechanism of stabilization for the case with a non-sigmoid functional response of predators. We assume that the carrying capacity of the prey is infinitely large in each site, and show that successful top-down regulation of this otherwise globally unstable system is made possible through an interplay between the unevenness of prey fitness across the sites and the rapid food-dependent migration of predators. We argue that this mechanism of stabilization is different from those previously reported in the literature: in particular, it requires a high degree of synchronicity in local oscillations of species densities across the sites. Prey outbreaks take place synchronously, but the unevenness of prey growth rates across the sites results in a pronounced difference in the species densities, and so the predator quickly disperses to the sites with the highest prey abundances. For this reason, the consumption of prey mostly takes place in the sites with high densities of prey, which assures an efficient suppression of outbreaks. Furthermore, when the total size of prey population is low, the distribution of both species among the sites becomes more even, and this prevents overconsumption of the prey by the predator. Finally, we put forward the hypothesis that this mechanism, when considered in a tri-trophic plankton community in the water column, can explain the stability of the nutrient-rich low-chlorophyll open ocean regions.     

The effects of between-habitat dispersal rate on protist communities and metacommunities in microcosms at two spatial scales

The effect of manipulation of between-habitat dispersal rates in multiple patch systems was examined experimentally using protist communities in laboratory microcosms. Replicate landscapes of eight microcosms (patches) at two spatial scales (patch sizes) were inoculated with 13 species of protists. Dispersal was carried out by transferring a small random sample of medium and protists from one randomly selected microcosm to another within a landscape. Four dispersal rates (24, 6, 2 and 0 transfers very 3 days) were used, and the microcosms were sampled after 6 and 12 weeks. Patch size had a consistent effect on within-path (community) and within-landscape (metacommunity) diversity, both being lower in small patch systems. Higher dispersal rates had a slight effect on community and metacommunity diversity after 12 weeks, with a tendency for higher dispersal to slightly offset the rate of loss of species. Both dispersal and patch size had effects on the abundance of many individual species, though in a variety of ways. The individual species results suggest that extinction is selective with respect to both patch size and dispersal rate treatments, and may be influenced by species interactions. It seems likely that in metacommunity systems of this sort, rather than mainland-island systems, the potential effect of between-patch dispersal rate in rescuing and recolonizing where local extinctions occur may be much reduced by the effect of selective extinction, relative to that expected under the assumption of random extinction.     

The effect of predator avoidance and travel time delay on the stability of predator-prey metacommunities

The stability conditions for an isolated specialist predator-prey community are fairly well understood. The spatial coupling of several such systems through dispersal of individuals can generate new dynamic behavior that is not yet completely understood. Many factors are known to be stabilizing or neutral, e.g., random dispersal or time delays, while others may induce instabilities in some cases but not others, e.g., density-dependent movement. We study the combination of two stabilizing mechanisms in a two-patch Rosenzweig-MacArthur model with a novel density-dependent movement term. Specifically, we assume that prey move between patches according to their perceived predation risk, and we include travel time between patches as a time delay. We show that the combination of mechanisms may be destabilizing even though each mechanism by itself is stabilizing. Our results show that a detailed knowledge of mechanisms and their temporal scales is necessary to correctly predict the stability of a metacommunity.     

Variations in the dispersal curves of macroalgal propagules from a source

The process of dispersal is critical to marine benthic species (i.e. invertebrates and algae) as a fundamental element of population ecology and a crucial ecological process that maintains the diversity in communities. We simultaneously sampled the abundance of spores inhabiting the water column at different distances from known parent sources at 3 sites along the coast of central Chile. From these data we constructed 258 dispersal curves for common rocky intertidal macroalgae. Only 43.8% of these curves could be predicted by the expected model, which describes the spatial distribution of propagules to be dominated by a larger concentration near the parent individual or “source site”, followed by a marked decrease in abundance with increasing distance. The curves that departed from the expected model (56.2%) were grouped into three curve types, according to the number of propagule abundance maxima observed in space. This work suggests that macroalgal propagule dispersal patterns are more variable than previously thought. The existence of several alternative curves to the expected model, as well as the presence of one to several abundance maxima associated with the differential distribution of propagule patches in the water column, suggests the idea that propagules are released in pulses which can be transported variable distances from the source site.     

The effects of resource enrichment, dispersal, and predation on local and metacommunity structure

Community structure is the observable outcome of numerous processes. We conducted a laboratory experiment using a microbial model system to disentangle effects of nutrient enrichment, dispersal, and predation on prey species richness and predator abundance at local and metacommunity scales. Prey species included: Chilomonas sp., Colpidium striatum, Colpoda cucullus, Paramecium tetraurelia, P. caudatum, Philodina sp., Spirostomum sp., Tetrahymena thermophila, and Uronema sp., and Stentor coeruleus was the predator used. We hypothesized that: (1) increased basal resources should maintain greater species richness and higher predator abundance; (2) dispersal should maintain greater species richness; and (3) predation should reduce species richness, especially in the high resource treatments relative to no-predator treatments. Our results support all three hypotheses. Further, we show that dispersal affects richness at the local community scale but not at the metacommunity scale. However, predation seems to have major effects at both the local and metacommunity scale. Overall, our results show that effects of resource enrichment, dispersal, and predation were mostly additive rather than interactive, indicating that it may be sometimes easier to understand their effects than generally thought due to complex interactive effects.     

Forecasting spatially structured populations: the role of dispersal and scale

We forecasted spatially structured population models with complex dynamics, focusing on the effect of dispersal and spatial scale on the predictive capability of nonlinear forecasting (NLF). Dispersal influences NLF ability by its influence on population dynamics. For simple 2-cell models, when dispersal is small, our ability to predict abundance in subpopulations decreased and then increased with increasing dispersal. Spatial heterogeneity, dispersal manner, and environmental noise did not qualitatively change this result. But results are not clear for complex spatial configurations because of complicated dispersal interactions across subpopulations. Populations undergoing periodic fluctuations could be forecasted perfectly for all deterministic cases that we studied, but less reliably when environmental noise was incorporated. More importantly, for all models that we have examined, NLF was much worse at larger spatial scales as a consequence of the asynchronous dynamics of subpopulations when the dispersal rate was below some critical value. The only difference among models was the critical value of dispersal rate, which varied with growth rate, carrying capacity, mode of dispersal, and spatial configuration. These results were robust even when environmental noise was incorporated. Intermittency, common in the dynamics of spatially structured populations, lowered the predictive capability of NLF. Forecasting population behaviour is of obvious value in resource exploitation and conservation. We suggest that forecasting at local scales holds promise, whereas forecasting abundance at regional scales may yield poor results. Improved understanding of dispersal can enhance the management and conservation of natural resources, and may help us to understand resource-exploitation strategies employed by local indigenous humans.     

Positive interactions and the emergence of community structure in metacommunities

The significant role of space in maintaining species coexistence and determining community structure and function is well established. However, community ecology studies have mainly focused on simple competition and predation systems, and the relative impact of positive interspecific interactions in shaping communities in a spatial context is not well understood. Here we employ a spatially explicit metacommunity model to investigate the effect of local dispersal on the structure and function of communities in which species are linked through an interaction web comprising mutualism, competition and exploitation. Our results show that function, diversity and interspecific interactions of locally linked communities undergo a phase transition with changes in the rate of species dispersal. We find that low spatial interconnectedness favors the spontaneous emergence of strongly mutualistic communities which are more stable but less productive and diverse. On the other hand, high spatial interconnectedness promotes local biodiversity at the expense of local stability and supports communities with a wide range of interspecific interactions. We argue that investigations of the relationship between spatial processes and the self-organization of complex interaction webs are critical to understanding the geographic structure of interactions in real landscapes.     

Climatic history and dispersal ability explain the relative importance of turnover and nestedness components of beta diversity

Aim We tested whether the geographic variation in the proportion of beta diversity attributed to nestedness or turnover components was explained by the effect of past glaciation events. Specifically, we tested the hypothesis that most of the beta diversity in regions retaining ice until recent periods was due to nestedness. Additionally, we tested whether the variation was influenced by thermal tolerance and the dispersal ability of species. Location This study analysed data from the New World. Methods We used presence/absence data for amphibians, birds and mammals of the New World. We calculated beta diversity among each 1°× 1° cell and the adjacent cells using the Sorensen dissimilarity index that expresses the total beta diversity. Furthermore, we partitioned it into turnover and nestedness components. The relative importance of the two latter components was expressed as the proportion of total beta diversity explained by nestedness (β_ratio). We calculated the correlation between β_ratio and the time each cell was free of ice since the last glaciation (cell age). To control the effects of spatial autocorrelation, we calculated geographically effective degrees of freedom. Results The proportion of beta diversity attributed to nestedness was negatively correlated with cell age. Moreover, this effect was stronger for amphibians than mammals, and stronger for mammals than birds. Main conclusions Our results are in accordance with the hypothesis that the nestedness component of beta diversity is more important in areas affected by glaciations until recent time. The beta diversity in high latitudes is the result of past extinctions and recent recolonization, which result in higher levels of nestedness. This process is more evident for vertebrates with lower dispersal ability and lower temperature tolerance.     

Quantifying the relative importance of natural variables,human disturbance and spatial processes in ecological status indicators of boreal lakes

Spatial processes are increasingly associated with species distributions in freshwaters. However, these processes are usually neglected in bioassessment techniques, which may introduce uncontrolled variation in ecological indicators used to express human disturbance. We used partial linear regression to quantify the relative importance of natural variables, human disturbance and spatial variables in structuring variation in boreal lake status indicators based on six biological indicator groups (phytoplankton, macrophytes, diatoms, littoral and profundal macroinvertebrates and fish). We found that, of the pure fractions, human disturbance explained most variation (7–32%) of the ecological quality ratios (EQRs) for all groups, with the exception of littoral macroinvertebrate metric, which was most controlled by natural and spatial variables (15% and 16%, respectively). In addition, pure fractions of natural and spatial variables and joint fractions of different explanatory variable groups structured all biological metrics to various degrees. Phytoplankton, diatom and profundal macroinvertebrate EQRs responded purest to human disturbance but only weakly to pure natural or spatial variation. Our work demonstrates that spatial processes and spatial structuring of the environment can bias bioassessment techniques and hinder the detection of human impact. Thus, it is important to acknowledge spatial autocorrelation, context of metacommunity dynamics, species dispersal traits and variable spatial extent when defining reference conditions and bioassessment techniques for different biological groups. More research is needed to better understand the relative role of spatial processes on ecological metrics originated from different freshwater ecosystems. To this end, our work provides an example of how sources of variation can be identified to increase accuracy in freshwater bioassessment.     

Effects of nonnative species on the stability of riverine fish communities

Despite the increasing ubiquity of biological invasions worldwide, little is known about the scale-dependent effects of nonnative species on real-world ecological dynamics. Here, using an extensive time series dataset of riverine fish communities across different biogeographic regions of the world, we assessed the effects of nonnative species on the temporal variability and synchrony in abundance at different organizational levels (population, metapopulation, community and metacommunity) and spatial scales (stream reach and river basin). At the reach scale, we found that populations of nonnative species were more variable over time than native species, and that this effect scaled up to the community level – significantly destabilizing the dynamics of riverine fish communities. Nonnative species not only contributed to reduced community stability, but also increased variability of native populations. By contrast, we found no effect of nonnative species dominance on local interspecific synchrony among native species. At the basin scale, nonnative metapopulations were again more variable than the native ones. However, neither native metapopulations nor metacommunities showed differences in temporal variability or synchrony as nonnative species dominance increased basin-wide. This suggests a ‘dilution effect’ where the contribution to regional stability of local native populations from sites displaying low levels of invasion reduced the destabilizing effects of nonnative species. Overall, our results indicate that accounting for the destabilizing effect of nonnative species is critical to understanding native species persistence and community stability.     

Scale-dependent effects of natural environmental gradients,industrial emissions and dispersal processes on zooplankton metacommunity structure: Implications for the bioassessment of boreal lakes

Environmental controls were traditionally considered as sole determinants of community assembly for freshwater bioassessment studies, whereas potential importance of dispersal processes and spatial scale have received limited attention. We conducted a bioassessment of lakes across northeast Alberta, Canada using crustacean zooplankton to develop a framework for evaluating if and how atmospheric emissions from the nearby Athabasca Oil Sands Region could impact their community assemblages. We quantified the effects of environmental gradients and spatially contingent dispersal processes for determining zooplankton community composition of 97 lakes at two spatial scales (regional and sub-regional) using constrained ordination, spatial modeling and variance partitioning techniques. Our findings indicated that effects of both environmental gradients and dispersal processes on species composition were scale-dependent. Zooplankton community composition was significantly correlated to environmental parameters that are directly and indirectly sensitive to industrial deposition including nitrate, sulphate, dissolved organic carbon, base cation, chloride, trace metal concentrations and predation regime, indicating their potential to track future environmental impacts. The relative importance of these environmental predictors varied with spatial scale, yet unraveling the effects of natural environmental heterogeneity vs. industrial deposition on this scale-dependency was not possible due to lack of regional baseline information. Dispersal processes were not important in shaping zooplankton communities at the sub-regional scale, but had limited, yet significant influence on species composition at the regional scale, emphasizing the need for cautious interpretation of broad-scale community patterns. Beyond establishing crucial regional baselines, our study highlights the necessity for explicit incorporation of dispersal effects and spatial scale in bioassessment of lakes across landscapes.     

Phenology and dispersal of Larinus minutus Gyllenhal and Larinus obtusus Gyllenhal (Coleoptera: Curculionidae), two biological control agents of Centaurea stoebe ssp. micranthos (spotted knapweed) in Michigan

Centaurea stoebe L. ssp. micranthos (Gugler) (spotted knapweed) is an invasive plant that has been the target of classical biological control in North America for more than four decades. The seedhead-feeding weevils Larinus minutus Gyllenhal and Larinus obtusus Gyllenhal (Coleoptera: Curculionidae) are two of the most-widely released C. stoebe control agents, and have more recently been introduced into the eastern US. While there have been many studies focusing on their ability to impact C. stoebe in the western US and Canada, there have been few studies from eastern North America, and basic knowledge of important aspects of their biology is lacking. Here we report on the phenology and dispersal of L. minutus and L. obtusus in Michigan. We regularly sampled two established Larinus spp. populations in southern Michigan in 2012 and 2013, and found that while adult abundance fluctuates during the growing season, they remained at easily detectable levels from mid-June through the end of August. We also used previously established populations of L. minutus and L. obtusus released in 1996 (n = 1), 2007 (n = 2), and 2010 (n = 5) to determine how dispersal of Larinus spp. into the surrounding landscape changes with time since release. Populations of Larinus spp. weevils showed little dispersal for 2 years post-release. However, after initial establishment dispersal rates increased rapidly, resulting in average dispersal rates that increased exponentially with time since release. These findings can inform future biological control release and sampling programs for Larinus spp. in eastern North America.     

Geographic variation in density-dependent dynamics impacts the synchronizing effect of dispersal and regional stochasticity

Explanations for the ubiquitous presence of spatially synchronous population dynamics have assumed that density-dependent processes governing the dynamics of local populations are identical among disjunct populations, and low levels of dispersal or small amounts of regionalized stochasticity (Moran effect) can act to synchronize populations. In this study we used historical spatially referenced data on gypsy moth (Lymantria dispar) outbreaks to document that density-dependent processes can vary substantially across geographical landscapes. This variation may be due in part to geographical variation in habitat (e.g., variation in forest composition). We then used a second-order log-linear stochastic model to explore how inter-population variation in density-dependent processes affects synchronization via either synchronous stochastic forcing or dispersal. We found that geographical variation in direct density-dependence (first order) greatly diminishes synchrony caused by stochasticity but only slightly decreases synchronization via dispersal. Variation in delayed density-dependence (second order) diluted synchrony caused by regional stochasticity to a lesser extent than first-order variation, but it did not have any influence on synchrony caused by dispersal. In general, synchronization caused by dispersal was primarily dependent upon the instability of populations and only weakly, if at all, affected by similarities in density-dependence among populations. We conclude that studies of synchrony should carefully consider both the nature of the synchronizing agents and the pattern of local density-dependent processes, including how these vary geographically.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Indicators and mechanisms of stability and resilience to climatic and landscape changes in a remnant calcareous grassland

When considering the development of conservation strategies for threatened plant communities it is crucial to understand their resilience to environmental change, taking into account current decline and the occurrence of further habitat fragmentation and climatic changes. Many recent works describe resilience character and elements, but there is little focus on the metrics and indices that describe elements of stability and specific resistance or resilience over the community composition matrix. Communities with strong niche selection might be restricted to specific resistance strategies to cope with environmental changes. This would result in a community at greater risk from increasing fragmentation and climatic changes.In a 35 years survey of relic calcareous grasslands, we looked at measures to identify the resilience mechanisms for stability in the presence and abundance of species. We used techniques of partitioning of temporal beta diversity in nestedness and turnover components, analysis of functional strategy changes and dissimilarity analysis to detect changes in between⿿plot diversity and exchanges.Contrary to expectations, we observe strong resilience with different stabilizing mechanisms both at plot level and exchanges between plots. At the scale of our grassland complex, response diversity and environmental stochasticity allow for the maintenance of high biodiversity under natural perturbations and gradual human-induced environmental changes. This highlights the importance of dispersal, recruitment dynamics and microsite diversity.Community resilience is more than just the sum of species resilience strategies; adaptive management strategies need more emphasis on the variability of conditions, as this can enable or disrupt important community resilience mechanisms.     

Speciation and the evolution of dispersal along environmental gradients

We analyze the joint evolution of an ecological character and of dispersal distance in asexual and sexual populations inhabiting an environmental gradient. Several interesting phenomena resulting from the evolutionary interplay of these characters are revealed. First, asexual and sexual populations exhibit two analogous evolutionary regimes, in which either speciation in the ecological character occurs in conjunction with evolution of short-range dispersal, or dispersal distance remains high and speciation does not occur. Second, transitions between these two regimes qualitatively differ between asexual and sexual populations, with the former showing speciation with long-range dispersal and the latter showing no speciation with short-range dispersal. Third, a phenotypic gradient following the environmental gradient occurs only in the last case, i.e., for non-speciating sexual populations evolving towards short-range dispersal. Fourth, the transition between the evolutionary regimes of long-range dispersal with no speciation and short-range dispersal with speciation is typically abrupt, mediated by a positive feedback between incipient speciation and the evolution of short-range dispersal. Fifth, even though the model of sexual evolution analyzed here does not permit assortative mating preferences, speciation occurs for a surprisingly wide range of conditions. This illustrates that dispersal evolution is a powerful alternative to preference evolution in enabling spatially distributed sexual populations to respond to frequency-dependent disruptive selection.