Spatial variation in caddisfly grazing regimes within a northern California watershed.

Ecologists seek better understanding of why species interactions change across space and time in natural communities. In streams, species effects on resources and community structure may change as physical characteristics of the stream environment change along drainage networks. We examined spatial and seasonal effects of armored grazers using a small-scale exclusion experiment that was replicated in streams of different drainage areas. Effects of grazing varied with stream size and were related to variation in grazer abundance and phenology. We identified three distinct grazing regimes and a stream size (drainage area [DA]) threshold corresponding to a shift from one to two functional trophic levels. In streams with DA < 1 km2, armored grazers did not reduce biomass of algal biofilms. In slightly larger streams (2-3 km2 DA), the armored grazer guild was dominated by bivoltine Glossosoma. These caddisflies persisted and limited algal biofilms throughout the summer in one of these streams. In the largest tributaries (DA > 10 km2), the grazer guild was dominated by univoltine caddisflies, and grazing limited algal biofilms in early summer, but not late summer, after caddisflies pupated. Drainage area is a useful predictor of spatial transitions in food web interactions within and among watersheds. Quantifying the drainage area threshold at which interactions change in catchments with differing geology, vegetation, hydrology, climate, land use, or species pools should help build the understanding we need to forecast ecological responses to environmental change.

Grazer traits, competition, and carbon sources to a headwater-stream food web.

We investigated the effect of grazing by a dominant invertebrate grazer (the caddisfly Glossosoma penitum) on the energy sources used by other consumers in a headwater-stream food web. Stable isotope studies in small, forested streams in northern California have shown that G. penitum larvae derive most of their carbon from algae, despite low algal standing crops. We hypothesized that the caddisfly competes with other primary consumers (including mayflies) for algal food and increases their reliance on terrestrial detritus. Because Glossosoma are abundant and defended from predators by stone cases, their consumption of algal energy may reduce its transfer up the food chain. We removed Glossosoma (natural densities >1000 caddisflies/m2) from five approximately 4 m2) stream sections during the summer of 2000 and measured responses of algae, invertebrate primary consumers, and invertebrate predators. The treatment reduced Glossosoma biomass by 80-90%. We observed a doubling in chlorophyll a per area in sections with reduced Glossosoma abundance and aggregative increases in the biomass of undefended primary consumers. Heptageniid mayfly larvae consumed more algae (as measured by stable carbon isotope ratios and gut content analysis) in caddisfly removal plots at the end of the 60-day experiment, although not after one month. We did not see isotopic evidence of increased algal carbon in invertebrate predators, however. Patterns of caddisfly and mayfly diets in the surrounding watershed suggested that mayfly diets are variable and include algae and detrital carbon in variable proportions, but scraping caddisflies consume primarily algae. Caddisfly and mayfly diets are more similar in larger, more productive streams where the mayflies assimilate more algae. Isotopic analysis, in combination with measurements of macroinvertebrate abundance and biomass in unmanipulated plots, suggested that a substantial portion of the invertebrate community (>50% of biomass) was supported, at least partially, by local algal carbon during midsummer. These data suggest that algae may be more important to community dynamics in headwater streams than their relatively low productivity would suggest. Through their high densities and relative invulnerability to predation, armored grazers may also affect community structure and flow of algal and detrital carbon in headwater streams.

Linkage of butterfly mate preference and wing color preference cue at the genomic location of wingless.

Sexual isolation is a critical form of reproductive isolation in the early stages of animal speciation, yet little is known about the genetic basis of divergent mate preferences and preference cues in young species. Heliconius butterflies, well known for their diversity of wing color patterns, mate assortatively as a result of divergence in male preference for wing patterns. Here we show that the specific cue used by Heliconius cydno and Heliconius pachinus males to recognize conspecific females is the color of patches on the wings. In addition, male mate preference segregates with forewing color in hybrids, indicating a genetic association between the loci responsible for preference and preference cue. Quantitative trait locus mapping places a preference locus coincident with the locus that determines forewing color, which itself is perfectly linked to the wing patterning candidate gene, wingless. Furthermore, yellow-colored males of the polymorphic race H. cydno alithea prefer to court yellow females, indicating that wing color and color preference are controlled by loci that are located in an inversion or are pleiotropic effects of a single locus. Tight genetic associations between preference and preference cue, although rare, make divergence and speciation particularly likely because the effects of natural and sexual selection on one trait are transferred to the other, leading to the coordinated evolution of mate recognition. This effect of linkage on divergence is especially important in Heliconius because differentiation of wing color patterns in the genus has been driven and maintained by natural selection for Müllerian mimicry.