Bathing birds bias beta-diversity: frequent dispersal by gulls homogenizes fauna in a rock-pool metacommunity.

Metacommunity theory generally predicts that regional dispersal of organisms among local habitat patches should influence spatial patterns of species diversity. In particular, increased dispersal rates are generally expected to increase local (alpha) diversity, yet homogenize local communities across the region (decreasing beta-diversity), resulting in no change in regional (gamma) diversity. Although the effect of dispersal on alpha-diversity has garnered much experimental attention, the influence of dispersal rates on diversity at larger spatial scales (beta and gamma) is poorly understood. Furthermore, these theoretical predictions are not well tested in the field, where other environmental factors (e.g., habitat size, resource density) likely also influence species diversity. Here, we used a system of freshwater rock pools on Appledore Island, Maine, USA, to test the effects of dispersal rate on species diversity in metacommunities. The pools exist in clusters (metacommunities) that experience different levels of dispersal imposed by gulls (Larus spp.), which we show to be frequent passive dispersers of rock-pool invertebrates. Although previous research has suggested that waterbirds may disperse aquatic invertebrates, our study is the first to quantify the rate at which such dispersal occurs and determine its effects on species diversity. In accordance with theory, we found that metacommunities experiencing higher dispersal rates had significantly more homogeneous local communities (reduced beta-diversity) and that gamma-diversity was not influenced by dispersal rate. Contrary to theoretical predictions, however, alpha-diversity in the rock pools was not significantly influenced by dispersal. Rather, local diversity was significantly positively related to local habitat size, and both alpha- and gamma-diversity were influenced by the physicochemical environment of the pools. These results provide an important field test of metacommunity theory, highlighting how local and regional factors interact to drive patterns of species diversity in metacommunities, and demonstrate that waterbirds are indeed important dispersal vectors for aquatic invertebrates.

Aquatic bacterial assemblage variability in the supra littoral zone of Appledore Island, Gulf of Maine.

We investigated the abundance and assemblage variability of bacteria in 10 spatially distinct freshwater pools on Appledore Island, Gulf of Maine. Assemblages were strongly heterogeneous between pools separated by even short distances. To gain insight into factors that may lead to the establishment of novel assemblages, we conducted an ecosystem-open choromophoric dissolved organic matter (CDOM) enrichment experiment within mesocosms inoculated with a standardized microbial community and observed patterns of their composition over time. Assemblages were strongly divergent from each other in composition after only 3 days of incubation. Divergence among mesocosms was significantly higher with increasing levels of CDOM. CDOM addition initially had a strong positive impact on bacterial operation taxonomic unit (OTU) richness and negative impact on bacterial OTU evenness, but no impact on total bacterial abundance, suggesting that factors controlling abundance are decoupled from those influencing overall composition.

Resource ecology of virulence in a planktonic host-parasite system: an explanation using dynamic energy budgets.

Parasites steal resources that a host would otherwise direct toward its own growth and reproduction. We use this fundamental notion to explain resource-dependent virulence in a fungal parasite (Metschnikowia)-zooplankton host (Daphnia) system and in a variety of other disease systems with invertebrate hosts. In an experiment, well-fed hosts died faster and produced more parasites than did austerely fed ones. This resource-dependent variation in virulence and other experimental results (involving growth and reproduction rate/timing of hosts) readily emerged from a model based on dynamic energy budgets. This model follows energy flow through the host, from ingestion of food, to internal energy storage, to allocation toward growth and reproduction or to a parasite that consumes these reserves. Acting as a consumer, the parasite catalyzes its own extinction, persistence with an energetically compromised host, or death of the host. In this last case, more resources for the host inadvertently fuels faster parasite growth, thereby accelerating the demise of the host (although the opposite result arises with different resource kinetics of the parasite). Thus, this model can explain how resource supply drives variation in virulence. This ecological dependence of virulence likely rivals and/or interacts with genetic mechanisms that often garner more attention in the literature on disease.

Friendly competition: evidence for a dilution effect among competitors in a planktonic host-parasite system.

The "dilution effect" concept in disease ecology offers the intriguing possibility that clever manipulation of less competent hosts could reduce disease prevalence in populations of more competent hosts. The basic concept is straightforward: host species vary in suitability (competence) for parasites, and disease transmission decreases when there are more incompetent hosts interacting with vectors or removing free-living stages of a parasite. However, host species also often interact with each other in other ecological ways, e.g., as competitors for resources. The net result of these simultaneous, multiple interactions (disease dilution and resource competition) is challenging to predict. Nonetheless, we see the signature of both roles operating concurrently in a planktonic host-parasite system. We document pronounced spatiotemporal variation in the size of epidemics of a virulent fungus (Metschnikowia bicuspidata) in Midwestern U.S. lake populations of a dominant crustacean grazer (Daphnia dentifera). We show that some of this variation is captured by changes in structure of Daphnia assemblages. Lake-years with smaller epidemics were characterized by assemblages dominated by less suitable hosts ("diluters," D. pulicaria and D. retrocurva, whose suitabilties were determined in lab experiments and field surveys) at the start of epidemics. Furthermore, within a season, less suitable hosts increased as epidemics declined. These observations are consistent with a dilution effect. However, more detailed time series analysis (using multivariate autoregressive models) of three intensively sampled epidemics show the signature of a likely interaction between dilution and resource competition between these Daphnia species. The net outcome of this interaction likely promoted termination of these fungal outbreaks. Should this outcome always arise in "friendly competition" systems where diluting hosts compete with more competent hosts? The answers to this question lie at a frontier of disease ecology.