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.     

Density-dependent dispersal and relative dispersal affect the stability of predator-prey metacommunities

Although density-dependent dispersal and relative dispersal (the difference in dispersal rates between species) have been documented in natural systems, their effects on the stability of metacommunities are poorly understood. Here we investigate the effects of intra- and interspecific density-dependent dispersal on the regional stability in a predator-prey metacommunity model. We show that, when the dynamics of the populations reach equilibrium, the stability of the metacommunity is not affected by density-dependent dispersal. However, the regional stability, measured as the regional variability or the persistence, can be modified by density-dependent dispersal when local populations fluctuate over time. Moreover these effects depend on the relative dispersal of the predator and the prey. Regional stability is modified through changes in spatial synchrony. Interspecific density-dependent dispersal always desynchronizses local dynamics, whereas intraspecific density-dependent dispersal may either synchronize or desynchronize it depending on dispersal rates. Moreover, intra- and interspecific density-dependent dispersal strengthen the top-down control of the prey by the predator at intermediate dispersal rates. As a consequence the regional stability of the metacommunity is increased at intermediate dispersal rates. Our results show that density-dependent dispersal and relative dispersal of species are keys to understanding the response of ecosystems to fragmentation.     

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     

Climate change in metacommunities: dispersal gives double-sided effects on persistence

Climate change is increasingly affecting the structure and dynamics of ecological communities both at local and at regional scales, and this can be expected to have important consequences for their robustness and long-term persistence. The aim of the present work is to analyse how the spatial structure of the landscape and dispersal patterns of species (dispersal rate and average dispersal distance) affects metacommunity response to two disturbances: (i) increased mortality during dispersal and (ii) local species extinction. We analyse the disturbances both in isolation and in combination. Using a spatially and dynamically explicit metacommunity model, we find that the effect of dispersal on metacommunity persistence is two-sided: on the one hand, high dispersal significantly reduces the risk of bottom-up extinction cascades following the local removal of a species; on the other hand, when dispersal imposes a risk to the dispersing individuals, high dispersal increases extinction risks, especially when dispersal is global. Large-bodied species with long generation times at the highest trophic level are particularly vulnerable to extinction when dispersal involves a risk. This suggests that decreasing the mortality risk of dispersing individuals by improving the quality of the habitat matrix may greatly increase the robustness of metacommunities.     

Gene networks and metacommunities: dispersal differences can override adaptive advantage

Dispersal is an important mechanism contributing to both ecological and evolutionary dynamics. In metapopulation and metacommunity ecology, dispersal enables new patches to be colonized; in evolution, dispersal counter-acts local selection, leading to regional homogenization. Here, I consider a three-patch metacommunity in which two species, each with a limiting quantitative trait underlain by gene networks of 16 to 256 genes, compete with one another and disperse among patches. Incorporating dispersal among heterogeneous patches introduces a tradeoff not observed in single-patch simulations: if the difference between gene network size of the two species is greater than the difference in dispersal ability (e.g., if the ratio of network sizes is larger than the ratio of dispersal abilities), then genetic architecture drives community outcome. However, if the difference in dispersal abilities is greater than gene network differences, then any adaptive advantages afforded by genetic architecture are over-ridden by dispersal. Thus, in addition to the selective pressures imposed by competition that shape the genetic architecture of quantitative traits, dispersal among patches creates an escape that may further alter the effects of different genetic architectures. These results provide a theoretical expectation for what we may observe as the field of ecological genomics develops.     

Dietary shift and lowered biomass gain of a generalist herbivore in species-poor experimental plant communities

Species loss of primary producers is likely to affect processes on other trophic levels. We studied consumption and individual performance of the generalist herbivore Parapleurus alliaceus (Orthoptera) in relation to the species richness of primary producers. Adult grasshoppers were caged and left to feed for 2 weeks on experimental grassland communities ranging in plant species richness from one, two, four, eight to 32 species. Low plant diversity had a negative effect on both plant community biomass and on biomass gain of female grasshoppers, feeding to produce eggs (male grasshoppers did not gain biomass during the feeding period). This was surprising because plots with high plant diversity had a low proportion of grass biomass and grasshoppers preferentially selected grasses, leading to a greater exploitation of grasses in experimental communities of higher diversity. Thus, the concurrent increase in non-grass species in the diet from these high-diversity communities must have been beneficial to the generalist herbivore. In addition to the positive effects of plant diversity, the presence of legumes in a mixture with grasses further enhanced the biomass gain of grasshoppers at a given level of diversity. These findings suggest that plant species loss may lead to shifts in herbivore population sizes, reducing those of generalists and benefiting specialists of the remaining plant species. Our results further suggest that generalist herbivores, by having feeding preferences, can also change the relative abundances of plant species with different functional characteristics. This may feedback on both composition and diversity of plant communities.     

The role of habitat connectivity and landscape geometry in experimental zooplankton metacommunities

Theory predicts that inter-patch dispersal rates and patterns of patch heterogeneity both have the potential to alter patterns of local and regional species diversity. To test this, we manipulated both rates of habitat connectivity and the geometric arrangement of habitat heterogeneity within regions of experimental zooplankton communities. We found no effects of habitat geometry on any metric of species diversity or composition. Additionally, we found no effect of habitat connectivity rate on local species diversity. We did, however, find that increasing connectivity led to a decrease in regional diversity, as well as an increase in the percent similarity of local communities within regions. Of all of the species in these communities, the relatively large cladoceran Ceriodaphnia reticulata significantly responded to the treatments, and had a higher probability of achieving high densities when connectance was high. As such, we suggest that this species played a large role in driving the increased local community similarity and decreased regional species richness as connectivity increased. These findings are in opposition to previous experimental studies of metacommunities, but support the notion that increased connectance among local patches may decrease regional diversity when patches are heterogeneous.     

The importance of dispersal related and local factors in shaping the taxonomic structure of diatom metacommunities

To date, little is known about the relative importance of dispersal related versus local factors in shaping microbial metacommunities. A common criticism regarding existing datasets is that the level of taxonomic resolution might be too coarse to reliably assess microbial community structure and study biogeographical patterns. Moreover, few studies have assessed the importance of geographic distance between habitats, which may influence metacommunity dynamics through its effect on dispersal rates. We applied variation partitioning analyses to 15 separate regional datasets on diatoms found in lakes in Eurasia, Africa and Antarctica. These analyses quantified the relative contributions of dispersal related and local factors in determining patterns of taxonomic turnover at the species and at the genus level. In general, results were similar at both taxonomic levels. Local environmental factors accounted for most of the explained variation (median=21%), whereas dispersal related factors were much less important (median of significant fractions=5.5% variation explained) and failed to significantly explain any variation, independent of the environmental variables, in the majority of the datasets. However, the amount of variation explained by dispersal related factors increased with increasing geographic distance and increasing taxonomic resolution. We extrapolated our regional scale observations to the global scale by combining the regional datasets into a global dataset comprising 1039 freshwater lakes from both hemispheres and spanning a geographic distance of over 19 000  km. At this global scale, taxonomic turnover was lowest in highly connected habitats, once environmental factors were partialled out. In common with many other studies of macro-organisms, these analyses showed that both dispersal related and local variables significantly contribute to the structure of global lacustrine diatom communities.     

Combined effects of zooplankton grazing and dispersal on the diversity and assembly mechanisms of bacterial metacommunities

Effects of dispersal and the presence of predators on diversity, assembly and functioning of bacterial communities are well studied in isolation. In reality, however, dispersal and trophic interactions act simultaneously and can therefore have combined effects, which are poorly investigated. We performed an experiment with aquatic metacommunities consisting of three environmentally different patches and manipulated dispersal rates among them as well as the presence or absence of the keystone species Daphnia magna. Daphnia magna reduced both local and regional diversity, whereas dispersal increased local diversity but decreased beta‐diversity having no net effect on regional diversity. Dispersal modified the assembly mechanisms of bacterial communities by increasing the degree of determinism. Additionally, the combination of the D. magna and dispersal increased the importance of deterministic processes, presumably because predator‐tolerant taxa were spread in the metacommunity via dispersal. Moreover, the presence of D. magna affected community composition, increased community respiration rates but did not affect bacterial production or abundance, whereas dispersal slightly increased bacterial production. In conclusion, our study suggests that predation by a keystone species such as D. magna and dispersal additively influence bacterial diversity, assembly processes and ecosystem functioning.     

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.     

Climate change effects on plant biomass alter dominance patterns and community evenness in an experimental old‐field ecosystem

Atmospheric and climatic change can alter plant biomass production and plant community composition. However, we know little about how climate change‐induced alterations in biomass production affect plant species composition. To better understand how climate change will alter both individual plant species and community biomass, we manipulated atmospheric [CO₂], air temperature, and precipitation in a constructed old‐field ecosystem. Specifically, we compared the responses of dominant and subdominant species to our climatic treatments, and explored how changes in plant dominance patterns alter community evenness over 2 years. Our study resulted in four major findings: (1) all treatments, elevated [CO₂], warming, and increased precipitation increased plant community biomass and the effects were additive rather than interactive, (2) plant species differed in their response to the treatments, resulting in shifts in the proportional biomass of individual species, which altered the plant community composition; however, the plant community response was largely driven by the positive precipitation response of Lespedeza, the most dominant species in the community, (3) precipitation explained most of the variation in plant community composition among treatments, and (4) changes in precipitation caused a shift in the dominant species proportional biomass that resulted in lower community evenness in the wet relative to dry treatments. Interestingly, compositional and evenness responses of the subdominant community to the treatments did not always follow the responses of the whole plant community. Our data suggest that changes in plant dominance patterns and community evenness are an important part of community responses to climatic change, and generally, that such compositional shifts can alter ecosystem biomass production and nutrient inputs.     

Diversity and community biomass depend on dispersal and disturbance in microalgal communities

The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance, i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the regulation of richness and evenness in early and late successional stages.     

A shift in the Hepatitis C virus genotype dominance in blood donor samples from Thailand

Hepatitis C virus (HCV) can be classified into six major genotypes. The HCV genotypes variability accounts for its geographical distribution, its responses to treatments and the clinical outcomes. The aim of this study was to determine the distribution of HCV genotypes among volunteer blood donors in Thailand. Samples from 135 anti-HCV positive blood donors were analyzed. HCV RNA and genotyping was carried out using nested polymerase chain reaction (PCR) and genotype-specific primer PCR for a portion of the core region. HCV RNA was detected in 109 samples (80.7%). Genotype analysis demonstrated four different genotypes. The most common was genotype 3a (36.7%), followed by genotype 6 (29.4%), 1a (19.3%), 1b (6.4%) and mixed infection (1.8%). Seven samples were untyped (6.4%) in the present study. In several previous reports, the prevalence found in Thailand was HCV genotypes 3, 1 and 6. The present results show an increasing importance of the genotype 6 in HCV infections. This study has also described for the first time in Thailand mixed infections of HCV genotypes.     

Effects of connectivity and recurrent local disturbances on community structure and population density in experimental metacommunities

Metacommunity theory poses that the occurrence and abundance of species is a product of local factors, including disturbance, and regional factors, like dispersal among patches. While metacommunity ideas have been broadly tested there is relatively little work on metacommunities subject to disturbance. We focused on how localized disturbance and dispersal interact to determine species composition in metacommunities. Experiments conducted in simple two-patch habitats containing eight protozoa and rotifer species tested how dispersal altered community composition in both communities that were disturbed and communities that connected to refuge communities not subject to disturbance. While disturbance lowered population densities, in disturbed patches connected to undisturbed patches this was ameliorated by immigration. Furthermore, species with high dispersal abilities or growth rates showed the fastest post-disturbance recovery in presence of immigration. Connectivity helped to counteract the negative effect of disturbances on local populations, allowing mass-effect-driven dispersal of individuals from undisturbed to disturbed patches. In undisturbed patches, however, local population sizes were not significantly reduced by emigration. The absence of a cost of dispersal for undisturbed source populations is consistent with a lack of complex demography in our system, such as age- or sex-specific emigration. Our approach provides an improved way to separate components of population growth from organisms' movement in post-disturbance recovery of (meta)communities. Further studies are required in a variety of ecosystems to investigate the transient dynamics resulting from disturbance and dispersal.     

A propagating heat wave model of skin electroporation

The main barrier to transdermal drug delivery in human skin is the stratum corneum. Pulsed electric fields (PEFs) of sufficient amplitude can create new aqueous pathways across this barrier and enhance drug delivery through the skin. Here, we describe a model of pore formation between adjacent corneocytes that predicts the following sequence of events: (1) the PEF rapidly charges the stratum corneum near the electrode until the transepidermal potential difference is large enough to drive water into a small region of the stratum corneum, creating new aqueous pathways. (2) PEFs then drive a high current density through this newly created electropore to generate Joule heating that warms the pore perimeter. (3) This temperature rise at the perimeter increases the probability of further electroporation there as the local sphingolipids reach their phase transition temperature. (4) This heat-generated wave of further electroporation propagates outward until the surface area of the pore becomes so large that the reduced current density no longer generates sufficient heat to reach the phase transition temperature of the sphingolipids. (5) Cooling and partial recovery occurs after the field pulse.This process yields large, high permeability regions in the stratum corneum at which molecules can more readily cross this skin barrier. We present a model for this process that predicts that the initial radius of the first aqueous pathway is approximately 5 nm for a transdermal voltage of 60 V at room temperature.     

Consistent pattern of local adaptation during an experimental heat wave in a pipefish-trematode host-parasite system

Extreme climate events such as heat waves are expected to increase in frequency under global change. As one indirect effect, they can alter magnitude and direction of species interactions, for example those between hosts and parasites. We simulated a summer heat wave to investigate how a changing environment affects the interaction between the broad-nosed pipefish (Syngnathus typhle) as a host and its digenean trematode parasite (Cryptocotyle lingua). In a fully reciprocal laboratory infection experiment, pipefish from three different coastal locations were exposed to sympatric and allopatric trematode cercariae. In order to examine whether an extreme climatic event disrupts patterns of locally adapted host-parasite combinations we measured the parasite's transmission success as well as the host's adaptive and innate immune defence under control and heat wave conditions. Independent of temperature, sympatric cercariae were always more successful than allopatric ones, indicating that parasites are locally adapted to their hosts. Hosts suffered from heat stress as suggested by fewer cells of the adaptive immune system (lymphocytes) compared to the same groups that were kept at 18°C. However, the proportion of the innate immune cells (monocytes) was higher in the 18°C water. Contrary to our expectations, no interaction between host immune defence, parasite infectivity and temperature stress were found, nor did the pattern of local adaptation change due to increased water temperature. Thus, in this host-parasite interaction, the sympatric parasite keeps ahead of the coevolutionary dynamics across sites, even under increasing temperatures as expected under marine global warming.