Disturbance and disease: host-parasite interactions in freshwater streams remain stable following wildfire.

Increases in the intensity and frequency of wildfires highlight the need to understand how fire disturbance affects ecological interactions. Though the effects of wildfire on free-living aquatic communities are relatively well-studied, how host-parasite interactions respond to fire disturbance is largely unexplored. Using a Before-After-Control-Impact design, we surveyed 10 stream sites (5 burned and 5 unburned) in the Willamette River Basin, Oregon and quantified snail host infection status and trematode parasite community structure 1 year before and two years after historic wildfires. Despite the severity of the wildfires, snail host populations did not show significant shifts in density or size distributions. We detected nine taxa of trematode parasites and overall probability of infection remained consistent over the three-year study period. However, at the taxon-specific level, we found evidence that infection probability by one trematode decreased and another increased after fire. In a larger dataset focusing on the first year after fire (9 burned, 8 unburned sites), we found evidence for subtle differences in trematode community structure, including higher Shannon diversity and evenness at the burned sites. Taken together, host-parasite interactions were remarkably stable for most taxa; for trematodes that did show responses, changes in abundance or behavior of definitive hosts may underlie observed patterns. These results have implications for using parasites as bioindicators of environmental change and suggest that aquatic snail-trematode interactions may be relatively resistant to wildfire disturbance in some ecosystems.

Trematode parasites exceed aquatic insect biomass in Oregon stream food webs.

Although parasites are increasingly recognized for their ecosystem roles, it is often assumed that free-living organisms dominate animal biomass in most ecosystems and therefore provide the primary pathways for energy transfer. To examine the contributions of parasites to ecosystem energetics in freshwater streams, we quantified the standing biomass of trematodes and free-living organisms at nine sites in three streams in western Oregon, USA. We then compared the rates of biomass flow from snails Juga plicifera into trematode parasites relative to aquatic vertebrate predators (sculpin, cutthroat trout and Pacific giant salamanders). The trematode parasite community had the fifth highest dry biomass density among stream organisms (0.40 g/m2 ) and exceeded the combined biomass of aquatic insects. Only host snails (3.88 g/m2 ), sculpin (1.11 g/m2 ), trout (0.73 g/m2 ) and crayfish (0.43 g/m2 ) had a greater biomass. The parasite 'extended phenotype', consisting of trematode plus castrated host biomass, exceeded the individual biomass of every taxonomic group other than snails. The substantial parasite biomass stemmed from the high snail density and infection prevalence, and the large proportional mass of infected hosts that consisted of trematode tissue (M = 31% per snail). Estimates of yearly biomass transfer from snails into trematodes were slightly higher than the combined estimate of snail biomass transfer into the three vertebrate predators. Pacific giant salamanders accounted for 90% of the snail biomass consumed by predators. These results demonstrate that trematode parasites play underappreciated roles in the ecosystem energetics of some freshwater streams.

Food-web interaction strength distributions are conserved by greater variation between than within predator-prey pairs.

Species interactions in food webs are usually recognized as dynamic, varying across species, space, and time because of biotic and abiotic drivers. Yet food webs also show emergent properties that appear consistent, such as a skewed frequency distribution of interaction strengths (many weak, few strong). Reconciling these two properties requires an understanding of the variation in pairwise interaction strengths and its underlying mechanisms. We estimated stream sculpin feeding rates in three seasons at nine sites in Oregon to examine variation in trophic interaction strengths both across and within predator-prey pairs. Predator and prey densities, prey body mass, and abiotic factors were considered as putative drivers of within-pair variation over space and time. We hypothesized that consistently skewed interaction strength distributions could result if individual interaction strengths show relatively little variation, or alternatively, if interaction strengths vary but shift in ways that conserve their overall frequency distribution. Feeding rate distributions remained consistently and positively skewed across all sites and seasons. The mean coefficient of variation in feeding rates within each of 25 focal species pairs across surveys was less than half the mean coefficient of variation seen across species pairs within a survey. The rank order of feeding rates also remained conserved across streams, seasons and individual surveys. On average, feeding rates on each prey taxon nonetheless varied by a hundredfold, with some feeding rates showing more variation in space and others in time. In general, feeding rates increased with prey density and decreased with high stream flows and low water temperatures, although for nearly half of all species pairs, factors other than prey density explained the most variation. Our findings show that although individual interaction strengths exhibit considerable variation in space and time, they can nonetheless remain relatively consistent, and thus predictable, compared to the even larger variation that occurs across species pairs. These results highlight how the ecological scale of inference can strongly shape conclusions about interaction strength consistency and help reconcile how the skewed nature of interaction strength distributions can persist in highly dynamic food webs.

What drives interaction strengths in complex food webs? A test with feeding rates of a generalist stream predator.

Describing the mechanisms that drive variation in species interaction strengths is central to understanding, predicting, and managing community dynamics. Multiple factors have been linked to trophic interaction strength variation, including species densities, species traits, and abiotic factors. Yet most empirical tests of the relative roles of multiple mechanisms that drive variation have been limited to simplified experiments that may diverge from the dynamics of natural food webs. Here, we used a field-based observational approach to quantify the roles of prey density, predator density, predator-prey body-mass ratios, prey identity, and abiotic factors in driving variation in feeding rates of reticulate sculpin (Cottus perplexus). We combined data on over 6,000 predator-prey observations with prey identification time functions to estimate 289 prey-specific feeding rates at nine stream sites in Oregon. Feeding rates on 57 prey types showed an approximately log-normal distribution, with few strong and many weak interactions. Model selection indicated that prey density, followed by prey identity, were the two most important predictors of prey-specific sculpin feeding rates. Feeding rates showed a positive relationship with prey taxon densities that was inconsistent with predator saturation predicted by current functional response models. Feeding rates also exhibited four orders-of-magnitude in variation across prey taxonomic orders, with the lowest feeding rates observed on prey with significant anti-predator defenses. Body-mass ratios were the third most important predictor variable, showing a hump-shaped relationship with the highest feeding rates at intermediate ratios. Sculpin density was negatively correlated with feeding rates, consistent with the presence of intraspecific predator interference. Our results highlight how multiple co-occurring drivers shape trophic interactions in nature and underscore ways in which simplified experiments or reliance on scaling laws alone may lead to biased inferences about the structure and dynamics of species-rich food webs.