Mark F. Dybdahl

In theory, parasites can create time-lagged, frequency-dependent selection in their hosts, resulting in oscillatory gene-frequency dynamics in both the host and the parasite (the Red Queen hypothesis). However, oscillatory dynamics have not been observed in natural populations. In the present study, we evaluated the dynamics of asexual clones of a New Zealand snail, Potamopyrgus antipodarum, and its trematode parasites over a five-year period. During the summer of each year, we determined host-clone frequencies in random samples of the snail to track genetic changes in the snail population. Similarly, we monitored changes in the parasite population, focusing on the dominant parasite, Microphallus sp., by calculating the frequency of clones in samples of infected individuals from the same collections. We then compared these results to the results of a computer model that was designed to examine clone frequency dynamics for various levels of parasite virulence. Consistent with these simulations and with ideas regarding dynamic coevolution, parasites responded to common clones in a time-lagged fashion. Finally, in a laboratory experiment, we found that clones that had been rare during the previous five years were significantly less infectible by Microphallus when compared to the common clones. Taken together, these results confirm that rare host genotypes are more likely to escape infection by parasites; they also show that host-parasite interactions produce, in a natural population, some of the dynamics anticipated by the Red Queen hypothesis.

Individual animal species can impact ecosystem processes, but few exotic invaders have demonstrated ecosystem-scale impacts, even when population sizes are large. We combined whole-stream measures of carbon and nitrogen fluxes with rates of consumption and ammonium excretion to show that an exotic freshwater snail, Potamopyrgus antipodarum, dominated these fluxes in a highly productive stream. The snails consumed 75% of gross primary productivity, and their excretion accounted for two-thirds of ammonium demand. Such large fluxes were due to high snail biomass rather than high rates of excretion or consumption. This exotic species may dramatically alter ecosystem function in rivers, with potential consequences for food web structure and element transport.

Sexual populations should be vulnerable to invasion and replacement by ecologically similar asexual females because asexual lineages have higher per capita growth rates. However, as asexual genotypes become common, they may also become disproportionately infected by parasites. The Red Queen hypothesis postulates that high infection rates in the common asexual clones could periodically favor the genetically diverse sexual individuals and promote the short-term coexistence of sexual and asexual populations. Testing this idea requires comparison of competing sexual and asexual lineages that are attacked by natural parasites. To date no such data have been available. Here, we report on long-term dynamics and parasite coevolution in a "mixed" (sexual and asexual) population of snails (Potamopyrgus antipodarum). We found that, within 7-10 years, the most common clones were almost completely replaced by initially rare clones in two different habitats, while sexuals persisted throughout the study period. The common clones, which were initially more resistant to infection, also became more susceptible to infection by sympatric (but not allopatric) parasites over the course of the study. These results are consistent with the Red Queen hypothesis and show that the coevolutionary dynamics predicted by the theory may also favor sexual reproduction in natural populations.

The functional importance of invasive animals may be measured as the degree to which they dominate secondary production, relative to native animals. We used this approach to examine dominance of invertebrate secondary production by invasive New Zealand mudsnails (Potamopyrgus antipodarum) in rivers. We measured secondary production of mudsnails and native invertebrates in three rivers in the Greater Yellowstone Area (Wyoming, USA): Gibbon River, Firehole River, and Polecat Creek. Potamopyrgus production was estimated by measuring in situ growth rates and multiplying by monthly biomass; native invertebrate production was estimated using size frequency and instantaneous growth methods. Mudsnail growth rates were high (up to 0.06 d(-1)) for juvenile snails and much lower for adult females (0.003 d(-1)). Potamopyrgus production in Polecat Creek (194 g x m(-2) x yr(-1)) was one of the highest values ever reported for a stream invertebrate. Native invertebrate production ranged from 4.4 to 51 g x m(-2) x yr(-1). Potamopyrgus was the most productive taxon and constituted 65-92% of total invertebrate productivity. Native invertebrate production was low in all streams. Based on a survey of production measures from uninvaded rivers, the distribution of secondary production across taxa was much more highly skewed toward the invasive dominant Potamopyrgus in the three rivers. We suggest that this invasive herbivorous snail is sequestering a large fraction of the carbon available for invertebrate production and altering food web function.