Elemental content of a host-parasite relationship in the threespine stickleback.

Parasite infections are ubiquitous and their effects on hosts could play a role in ecosystem processes. Ecological stoichiometry provides a framework to study linkages between consumers and their resource, such as parasites and their host, and ecosystem process; however, the stoichiometric traits of host-parasite associations are rarely quantified. Specifically, it is unclear whether parasites' elemental ratios closely resemble those of their host or if infection is related to host stoichiometry, especially in vertebrate hosts. To answer such questions, we measured the elemental content (%C, %N, and %P) and molar ratios (C:N, C:P, and N:P) of parasitized and unparasitized Gasterosteus aculeatus (three-spined stickleback) and their cestode parasite, Schistocephalus solidus. Host and parasite elemental content were distinct from each other, and parasites were generally higher in %C and lower in %N and %P. Parasite infections were related to host C:N, with infected hosts being lower in C:N. Parasite elemental content was independent of their host, but parasite body mass and parasite density were important drivers of parasite stoichiometry. Overall, these potential effects of parasite infections on host stoichiometry along with parasites' distinct elemental compositions suggest parasites may further contribute to differences in how individual hosts store and recycle nutrients.

Host ecology and biogeography drive parasite community composition in Atlantic killifishes.

Understanding the mechanisms of parasite community assembly can be confounded by phylogenetic distance among host species. Addressing this requires focusing on parasite communities within closely related taxa. Thus, we took a macroecological approach to examining parasite community structure within Killifish species in the genus Fundulus to disentangle the effects of host phylogeny and ecological variables. We constructed a database of parasite communities within Fundulus species from 15 published and unpublished surveys covering the Atlantic coast of the US and Canada. The database was expanded by sampling sites in underrepresented provinces and states, totaling 10 Fundulus species from 57 unique geographic sites. Univariate analysis of observed parasite species richness among Fundulus populations in the dataset found that latitude, climate type, and salinity were the dominant factors determining parasite species richness. Multivariate analysis found that host species and landscape type were the most important factors in determining the similarity of parasite assemblages. Unexpectedly, parasite species richness decreased in low latitudes, and host phylogenetic distance was not found to be a significant factor in the similarity of parasite communities. These results indicate that commonly reported large-scale drivers of parasite community structure, such as latitude and phylogeny, could have diminished significance at the host genus level relative to host ecology, biogeography, and local landscape factors.

Host Energetics Explain Variation in Parasite Productivity across Hosts and Ecosystems.

AbstractParasites are thought to play a role in ecosystem energetics, in part because some ecosystems harbor a substantial amount of parasite biomass. Nevertheless, the extent to which parasite biomass accurately reflects the flow of energy from hosts to parasites-and the linkages between their energetics-remains unclear. Here, we estimate parasite community energetics at the host and ecosystem level and test predictions for parasite energetics using the metabolic theory of ecology. Across 27 host species, parasite community abundance declines with average individual parasite energy use Rp as Rp-0.50 and increases with host metabolic rate Rh as Rh0.64, which is inconsistent with metabolic theory. We next test whether the fraction of host energy that is allocated to parasitism is invariant across hosts. Our empirical analysis demonstrates that 85% of the variation in parasite community energy use can be explained by differences in host metabolic rate. However, parasite community energy use increases allometrically with host metabolic rate Rh as Rh0.67, suggesting that the fraction of host energy used by parasites declines with host metabolic rate. At the ecosystem level, we show that the energy flowing through parasite communities scales allometrically with the total rate of energy use by their fish hosts across three ecosystems. Importantly, directly examining energy flux revealed variation in parasite energy use among ecosystems that was not apparent when examining differences in biomass. Taken together, these results establish strong empirical links between host and parasite energetics, but our findings often did not align with predictions based on metabolic theory.

Historical contingency in parasite community assembly: Community divergence results from early host exposure to symbionts and ecological drift.

Host individuals are commonly coinfected with multiple parasite species that may interact to shape within-host parasite community structure. In addition to within-host species interactions, parasite communities may also be structured by other processes like dispersal and ecological drift. The timing of dispersal (in particular, the temporal sequence in which parasite species infect a host individual) can alter within-host species interactions, setting the stage for historical contingency by priority effects, but how persistently such effects drive the trajectory of parasite community assembly is unclear, particularly under continued dispersal and ecological drift. We tested the role of species interactions under continued dispersal and ecological drift by simultaneously inoculating individual plants of tall fescue with a factorial combination of three symbionts (two foliar fungal parasites and a mutualistic endophyte), then deploying the plants in the field and tracking parasite communities as they assembled within host individuals. In the field, hosts were exposed to continued dispersal from a common pool of parasites, which should promote convergence in the structure of within-host parasite communities. Yet, analysis of parasite community trajectories found no signal of convergence. Instead, parasite community trajectories generally diverged from each other, and the magnitude of divergence depended on the initial composition of symbionts within each host, indicating historical contingency. Early in assembly, parasite communities also showed evidence of drift, revealing another source of among-host divergence in parasite community structure. Overall, these results show that both historical contingency and ecological drift contributed to divergence in parasite community assembly within hosts.

Disease decreases variation in host community structure in an old-field grassland.

Disease may drive variation in host community structure by modifying the interplay of deterministic and stochastic processes that shape communities. For instance, deterministic processes like ecological selection can benefit species less impacted by disease. When communities have higher levels of disease and disease consistently selects for certain host species, this can reduce variation in host community composition. On the other hand, when host communities are less impacted by disease and selection is weaker, stochastic processes (e.g., drift, dispersal) may play a bigger role in host community structure, which can increase variation among communities. While effects of disease on host community structure have been quantified in field experiments, few have addressed the role of disease in modulating variation in structure among host communities. To address this, we conducted a field experiment spanning three years, using a tractable system: foliar fungal pathogens in an old-field grassland community dominated by the grass Lolium arundinaceum, tall fescue. We reduced foliar fungal disease burden in replicate host communities (experimental plots in intact vegetation) in three fungicide regimens that varied in the seasonal duration of fungicide treatment and included a fungicide-free control. We measured host diversity, biomass, and variation in community structure among replicate communities. Disease reduction generally decreased plant richness and increased aboveground biomass relative to communities experiencing ambient levels of disease. These changes in richness and aboveground biomass were consistent across years despite changes in structure of the plant communities over the experiment's three years. Importantly, disease reduction amplified host community variation, suggesting that disease diminished the degree to which host communities were structured by stochastic processes. These results of experimental disease reduction both highlight the potential importance of stochastic processes in plant communities and reveal the potential for disease to regulate variation in host community structure.

Historical contingency and the role of post-invasion evolution in alternative community states.

Historical contingency has long figured prominently in the conceptual frameworks of evolutionary biology and community ecology. Evolutionary biologists typically consider the effects of chance mutation and historical contingency in driving divergence and convergence of traits in populations, whereas ecologists instead are often interested in the role of historical contingency in community assembly and succession. Although genetic differences among individuals in populations can influence community interactions, variability among populations of the same species has received relatively little attention for its potential role in community assembly and succession. We used a community-level study of experimental evolution in two compositionally different assemblages of protists and rotifers to explore whether initial differences in species abundances among communities attributed to differences in evolutionary history, persisted as species that continued to evolve over time. In each assemblage, we observed significant convergence between two invaded treatments initially differing in evolutionary history over an observation period equal to ~40-80 generations for most species. Nonetheless, community structure failed to converge completely across all invaded treatments within an assemblage to a single structure. This suggests that whereas the species in the assemblage represent a common selective regime, differences in populations reflecting their evolutionary history can produce long-lasting transient alternative community states. In one assemblage, we also observed increasing within-treatment variability among replicate communities over time, suggesting that ecological drift may be another factor contributing to community change. Although subtle, these transient alternative states, in which communities differed in the abundance of interacting species, could nonetheless have important functional consequences, suggesting that the role of evolution in driving these states deserves greater attention.

Temporal Community Structure in Two Gregarines (Rotundula gammari and Heliospora longissima) Co-Infecting the Amphipod Gammarus fasciatus.

This study surveyed gregarine parasites that infect the amphipod, Gammarus fasciatus , to investigate temporal dynamics in infracommunity structure. We sampled a population of hosts for 2 yr from the north branch of the Raritan River in New Jersey. These hosts were infected with 2 direct life cycle gregarine parasites, Rotundula gammari and Heliospora longissima. Infections were separated temporally, with the prevalence of R. gammari peaking within the amphipod population in the fall (prevalence = 78% year 1 and 97% year 2) and H. longissima peaking in early spring (prevalence = 41% year 1 and 52% year 2). Increases in host population density did not significantly correlate with the abundance of these 2 parasites. However, H. longissima abundance was positively correlated with host body weight while R. gammari showed no significant relationship. The mean body mass of amphipods infected with H. longissima was 20.7 ± 1. 2 mg, and with R. gammari 8.1 ± 0.2 mg, which suggests a sized-based infection pattern. Mixed species infections were infrequent with an overall prevalence of 4.6%. When both gregarine species co-infected the same host, the R. gammari but not the H. longissima infrapopulation size was significantly lower when compared to single-species infections, suggesting asymmetric interactions. We conclude that the observed temporal patterns of infection by the 2 parasites are driven by a seasonal change in host demographics and size-dependent infections. We argue that specificity for host developmental stages may have arisen as a mechanism to avoid overlap between these gregarine species.