Functional trait sorting increases over succession in metacommunity mosaics of fish assemblages.

Metacommunity theory predicts that the relative importance of regional and local processes structuring communities will change over time since initiation of community assembly. Determining effects of these processes on species and trait diversity over succession remains largely unaddressed in metacommunity ecology to date, yet could confer an improved mechanistic understanding of community assembly. To test theoretical predictions of the increasing importance of local processes in structuring communities over successional stages in metacommunities, we evaluated fish species and trait diversity in three pond metacommunities undergoing secondary succession from beaver (Castor canadensis) disturbance. Processes influencing taxonomic and trait diversity were contrasted across pond communities of different ages and in reference streams. Counter to predictions, the local environment became less important in structuring communities over succession but did exert a stronger effect on trait sorting. Beta diversity and trait richness declined over succession while there was no influence on species richness or trait dispersion. The trait filtering in older habitats was likely a response to the larger and deeper pond ecosystems characteristic of late succession. In contrast to these observed effects in ponds, the local environment primarily structured species and trait diversity in streams. Analyses of the relative importance of regional and local processes in structuring fish assemblages within each pond metacommunity suggests that habitat age and connectivity were more important than the environment in structuring communities but contributions were region and scale-dependent. Together, these findings highlight that regional and local processes can differentially influence taxonomic and trait diversity in successional metacommunity mosaics.

Native species dispersal reduces community invasibility by increasing species richness and biotic resistance.

Recent studies indicate that diversity-invasibility relationships can depend on spatial scale, but the contributing role of native species dispersal among local communities in mediating these relationships remains unaddressed. Metacommunity ecology highlights the effects of species dispersal rates on local diversity, thereby suggesting that native species dispersal may influence local biotic resistance to invasion by non-native species. However, the effects of native species dispersal rates on local native diversity and invasibility could depend on any intraspecific differences of the invader that may alter establishment success. Here, I experimentally tested for the influence of native dispersal-diversity relationships on the invasibility of native communities by a non-native species represented by core, midrange and peripheral regions of the introduced geographic range. In mesocosms, native plankton communities were connected by low or moderate rates of dispersal to yield dispersal rate-driven differences in native species richness prior to invasion by a non-native zooplankter, Daphnia lumholtzi. After invasion, establishment success and effects of the non-native species on native community structure and ecosystem properties were evaluated as a function of dispersal rate and invader source region relative to a control without native species. Native species richness was greater at the moderate dispersal rate than the low dispersal rate and yielded a dispersal rate-dependent diversity-invasibility relationship that was robust to invader source region. There was almost no establishment success of the non-native species at moderate dispersal and reduced success at low dispersal relative to the control. Invader population growth rates were negative only at the moderate dispersal rate. Effects of species dispersal on native community and ecosystem response were more influential than effects of invasion and impacts associated with invader source region. The results demonstrate that dispersal-diversity relationships can influence diversity-invasibility relationships at the local spatial scale. These dispersal-driven responses of invasion were unaffected by any ecological differences associated with invasion history-related intraspecific variation of the non-native species. This study emphasizes that dispersal rates of native species in metacommunities can differentially alter local biotic resistance to invasion. Thus, native species dispersal rates have largely been an underappreciated local diversity maintenance mechanism that can confer insurance against biological invasions.

Emergence of a novel prey life history promotes contemporary sympatric diversification in a top predator.

Intraspecific phenotypic variation can strongly impact community and ecosystem dynamics. Effects of intraspecific variation in keystone species have been shown to propagate down through the food web by altering the adaptive landscape for other species and creating a cascade of ecological and evolutionary change. However, similar bottom-up eco-evolutionary effects are poorly described. Here we show that life history diversification in a keystone prey species, the alewife (Alosa pseudoharengus), propagates up through the food web to promote phenotypic diversification in its native top predator, the chain pickerel (Esox niger), on contemporary timescales. The landlocking of alewife by human dam construction has repeatedly created a stable open water prey resource, novel to coastal lakes, that has promoted the parallel emergence of a habitat polymorphism in chain pickerel. Understanding how strong interactions propagate through food webs to influence diversification across multiple trophic levels is critical to understand eco-evolutionary interactions in complex natural ecosystems.

Intraspecific phenotypic variation among alewife populations drives parallel phenotypic shifts in bluegill.

Evolutionary diversification within consumer species may generate selection on local ecological communities, affecting prey community structure. However, the extent to which this niche construction can propagate across food webs and shape trait variation in competing species is unknown. Here, we tested whether niche construction by different life-history variants of the planktivorous fish alewife (Alosa pseudoharengus) can drive phenotypic divergence and resource use in the competing species bluegill (Lepomis macrochirus). Using a combination of common garden experiments and a comparative field study, we found that bluegill from landlocked alewife lakes grew relatively better when fed small than large zooplankton, had gill rakers better adapted for feeding on small-bodied prey and selected smaller zooplankton compared with bluegill from lakes with anadromous or no alewife. Observed shifts in bluegill foraging traits in lakes with landlocked alewife parallel those in alewife, suggesting interspecific competition leading to parallel phenotypic changes rather than to divergence (which is commonly predicted). Our findings suggest that species may be locally adapted to prey communities structured by different life-history variants of a competing dominant species.

Predation inhibits the positive effect of dispersal on intraspecific and interspecific synchrony in pond metacommunities.

Recent interest in the ecological drivers of compensatory and synchronous population dynamics has provided an improved yet incomplete understanding of local and regional population oscillations in response to variable environments. Here, we evaluate the effect of dispersal rate and spatiotemporal heterogeneity in predation by the selective planktivore, bluegill sunfish (Lepomis macrochirus), on local and regional dynamics of zooplankton in pond metacommunities. A metacommunity consisted of three pond mesocosm communities, one with constant presence of predators, one without predators, and one with alternating presence-absence of predators. The three communities were connected at either no, low (0.7% per day), or high (20% per day) planktonic dispersal. Results demonstrate that heterogeneous predation (1) prevents spatial synchrony among prey populations across local communities, (2) disrupts the synchronous population dynamics within communities produced by dispersal, and (3) induces local compensatory dynamics between species within communities regardless of dispersal rate. Taken together, the results emphasize that spatiotemporal heterogeneity in selective predation can inhibit both intraspecific and interspecific synchrony in metacommunities.

Intraspecific phenotypic variation in a fish predator affects multitrophic lake metacommunity structure.

Contemporary insights from evolutionary ecology suggest that population divergence in ecologically important traits within predators can generate diversifying ecological selection on local community structure. Many studies acknowledging these effects of intraspecific variation assume that local populations are situated in communities that are unconnected to similar communities within a shared region. Recent work from metacommunity ecology suggests that species dispersal among communities can also influence species diversity and composition but can depend upon the relative importance of the local environment. Here, we study the relative effects of intraspecific phenotypic variation in a fish predator and spatial processes related to plankton species dispersal on multitrophic lake plankton metacommunity structure. Intraspecific diversification in foraging traits and residence time of the planktivorous fish alewife (Alosa pseudoharengus) among coastal lakes yields lake metacommunities supporting three lake types which differ in the phenotype and incidence of alewife: lakes with anadromous, landlocked, or no alewives. In coastal lakes, plankton community composition was attributed to dispersal versus local environmental predictors, including intraspecific variation in alewives. Local and beta diversity of zooplankton and phytoplankton was additionally measured in response to intraspecific variation in alewives. Zooplankton communities were structured by species sorting, with a strong influence of intraspecific variation in A. pseudoharengus. Intraspecific variation altered zooplankton species richness and beta diversity, where lake communities with landlocked alewives exhibited intermediate richness between lakes with anadromous alewives and without alewives, and greater community similarity. Phytoplankton diversity, in contrast, was highest in lakes with landlocked alewives. The results indicate that plankton dispersal in the region supplied a migrant pool that was strongly structured by intraspecific variation in alewives. This is one of the first studies to demonstrate that intraspecific phenotypic variation in a predator can maintain contrasting patterns of multitrophic diversity in metacommunities.

Species dispersal rates alter diversity and ecosystem stability in pond metacommunities.

Metacommunity theory suggests that relationships between diversity and ecosystem stability can be determined by the rate of species dispersal among local communities. The predicted relationships, however, may depend upon the relative strength of local environmental processes and disturbance. Here we evaluate the role of dispersal frequency and local predation perturbations in affecting patterns of diversity and stability in pond plankton metacommunities. Pond metacommunities were composed of three mesocosm communities: one of the three communities maintained constant "press" predation from a selective predator, bluegill sunfish (Lepomis macrochirus); the second community maintained "press" conditions without predation; and the third community experienced recurrent "pulsed" predation from bluegill sunfish. The triads of pond communities were connected at either no, low (0.7%/d), or high (20%/d) planktonic dispersal. Richness and composition of zooplankton and stability of plankton biomass and ecosystem productivity were measured at local and regional spatial scales. Dispersal significantly affected diversity such that local and regional biotas at the low dispersal rate maintained the greatest number of species. The unimodal local dispersal-diversity relationship was predator-dependent, however, as selective press predation excluded species regardless of dispersal. Further, there was no effect of dispersal on beta diversity because predation generated local conditions that selected for distinct community assemblages. Spatial and temporal ecosystem stability responded to dispersal frequency but not predation. Low dispersal destabilized the spatial stability of producer biomass but stabilized temporal ecosystem productivity. The results indicate that selective predation can prevent species augmentation from mass effects but has no apparent influence on stability. Dispersal rates, in contrast, can have significant effects on both species diversity and ecosystem stability at multiple spatial scales in metacommunities.

Prey dispersal rate affects prey species composition and trait diversity in response to multiple predators in metacommunities.

1. Recent studies indicate that large-scale spatial processes can alter local community structuring mechanisms to determine local and regional assemblages of predators and their prey. In metacommunities, this may occur when the functional diversity represented in the regional predator species pool interacts with the rate of prey dispersal among local communities to affect prey species diversity and trait composition at multiple scales. 2. Here, we test for effects of prey dispersal rate and spatially and temporally heterogeneous predation from functionally dissimilar predators on prey structure in pond mesocosm metacommunities. An experimental metacommunity consisted of three pond mesocosm communities supporting two differentially size-selective invertebrate predators and their zooplankton prey. In each metacommunity, two communities maintained constant predation and supported either Gyrinus sp. (Coleoptera) or Notonecta ungulata (Hemiptera) predators generating a spatial prey refuge while the third community supported alternating predation from Gyrinus sp. and N. ungulata generating a temporal prey refuge. Mesocosm metacommunities were connected at either low (0.7% day(-1)) or high (10% day(-1)) planktonic prey dispersal. The diversity, composition and body size of zooplankton prey were measured at local and regional (metacommunity) scales. 3. Metacommunities experiencing the low prey dispersal rate supported the greatest regional prey species diversity (H') and evenness (J'). Neither dispersal rate nor predation regime affected local prey diversity or evenness. The spatial prey refuge at low dispersal maintained the largest difference in species composition and body size diversity between communities under Gyrinus and Notonecta predation, suggesting that species sorting was operating at the low dispersal rate. There was no effect of dispersal rate on species diversity or body size distribution in the temporal prey refuge. 4. The frequency distribution, but not the range, of prey body sizes within communities depended upon prey dispersal rate and predator identity. Taken together, these results demonstrate that prey dispersal rate can moderate the strength of predation to influence prey species diversity and the local frequency distribution of prey traits in metacommunities supporting ecologically different predators.

Planktonic dispersal dampens temporal trophic cascades in pond metacommunities.

Trophic cascades, in which changes in predation affect the biomass of lower trophic levels, vary substantially in strength and incidence. Most work to explain this variation has focused on local factors and has ignored larger regional effects. To study how metacommunity dynamics can alter trophic cascades, we constructed mesocosm metacommunities consisting of three pond communities with heterogeneous levels of fish predation and examined how planktonic dispersal rate (5-140% per week) affected biomass partitioning. Two of the three communities differed continually in the occurrence of fish and supported different but constant environments in a 'spatial trophic cascade,' while the third community supported temporally variable fish occurrence in a 'temporal trophic cascade.' We find that the presence, but the not the magnitude, of dispersal dampens temporal trophic cascades through an increase in grazer biomass. In contrast, dispersal has no effect on the strength of spatial cascades due to strong sorting pressures in the communities with constant presence or absence of fish as top predators.

Contrasting demographic and genetic estimates of dispersal in the endangered Coahuilan box turtle: a contemporary approach to conservation.

The evolutionary viability of an endangered species depends upon gene flow among subpopulations and the degree of habitat patch connectivity. Contrasting population connectivity over ecological and evolutionary timescales may provide novel insight into what maintains genetic diversity within threatened species. We employed this integrative approach to evaluating dispersal in the critically endangered Coahuilan box turtle (Terrapene coahuila) that inhabits isolated wetlands in the desert-spring ecosystem of Cuatro Ciénegas, Mexico. Recent wetland habitat loss has altered the spatial distribution and connectivity of habitat patches; and we therefore predicted that T. coahuila would exhibit limited movement relative to estimates of historic gene flow. To evaluate contemporary dispersal patterns, we employed mark-recapture techniques at both local (wetland complex) and regional (intercomplex) spatial scales. Gene flow estimates were obtained by surveying genetic variation at nine microsatellite loci in seven subpopulations located across the species' geographical range. The mark-recapture results at the local spatial scale reveal frequent movement among wetlands that was unaffected by interwetland distance. At the regional spatial scale, dispersal events were relatively less frequent between wetland complexes. The complementary analysis of population genetic substructure indicates strong historic gene flow (global F(ST) = 0.01). However, a relationship of genetic isolation by distance across the geographical range suggests that dispersal limitation exists at the regional scale. Our approach of contrasting direct and indirect estimates of dispersal at multiple spatial scales in T. coahuila conveys a sustainable evolutionary trajectory of the species pending preservation of threatened wetland habitats and a range-wide network of corridors.