Jenny M. Schmid-Araya

THE IMPORTANCE OF MEIOFAUNA IN FOOD WEBS: EVIDENCE FROM AN ACID STREAM

Seasonal food webs were constructed for the whole invertebrate assemblage (meio- and macrofauna) inhabiting Broadstone Stream (southeast England). High and uniform taxonomic resolution was applied in a dietary analysis, by resolving the complete benthic community to species, including the meiofauna, protozoa, and algae. Meiofauna accounted for 70% of all species in the summary web and for 73% and 63% of those in the summer/autumn and spring webs, respectively. The web structure changed between summer/autumn and winter/spring, due to differences in species composition. Many stream invertebrates fed on meiofauna and organic matter. Addition of meiofauna to the Broadstone web increased the percentage of intermediate species. Seasonal webs contained between 54 (spring 1997) and 86 (autumn 1996) interactive taxa and 229–378 trophic links. Marked differences in web complexity were found between the summer/autumn and winter/spring periods. Meiofauna accounted for most of the links in the web with a high proportion of intermediate–intermediate links in summer and autumn (0.421–0.440) and also of intermediate–basal links during winter and spring (0.509–0.628). In general, the summary web showed that intermediate species and basal resources were numerically dominant components in this stream. Web connectance rose slightly between summer (0.052) and winter (0.061) and increased further in spring (0.079), coinciding with a reduction in species number. A high fraction of detritivores was combined with omnivorous predators, many of which supplemented their diets with organic matter and, depending on season, with algae and invertebrate eggs. In addition, a wide range of feeding modes was found among meiofaunal species. The diversity of the Broadstone community suggests that the impact of top predators tends to dissipate. A low proportion of top predators in the web was combined with a low mean number of prey items, other than detritus, in their guts (large predators, 1.08–1.26 prey/individual gut; small-sized tanypods, 2.15–2.32 prey/individual gut). Dietary similarity was highest in autumn and winter 1996, and observed feeding links of the most common predatory species showed low overlap in their diets. The web architecture of this stream is reticulate and complex, and the patterns observed in these seasonal webs differed from previous stream webs, resulting in low connectance, high linkage density, long food chains, and a high proportion of intermediate species and of intermediate–intermediate links. The food web derived from Broadstone Stream clearly demonstrates that the meiofauna increases web complexity and thus, taking into account their functional diversity, may be crucial to the understanding of food web properties and ecosystems processes in streams.

Scaling in stream communities

Scaling relationships between population density (N) and body size (W), and of their underlying size distributions, can contribute to an understanding of how species use resources as a function of size. In an attempt to resolve the controversy over the form of scaling relationships, an extensive dataset, comprising 602 invertebrate species, was obtained from two geographically separate stream communities (Seebach in Austria and Mynach in Wales). We analysed the temporal consistency of the N–W relationship, which was subjected to ordinary least squares (OLS), bisector (OLSBIS) and quantile regressions, and species–size spectra with seasonally collated data. Slopes of seasonal OLSBIS regressions did not depart from −1 in either community, indicating a seasonally convergent scaling relationship, which is not energetically constrained. Species–size spectra may scale with habitat complexity, providing an alternative explanation for the observed body–size scaling. In contrast to the right–skewed species–size frequency distributions of single–species assemblages, the size spectra of these benthic communities exhibited ‘central tendencies’, reflecting their phyletic constitution. The shape of species body–mass spectra differed between the two communities, with a bimodal and seasonally convergent pattern in the Seebach community and a seasonally shifting unimodality in the Mynach community. The body–size spectra of large, mostly insect, species (greater than or equal to 1 mm) scaled to seasonal variations in habitat complexity (i.e. fractal D), suggesting that habitat structure constrains the community organization of stream benthos.

Life history allometries and production of small fauna

The production of heterotrophic biomass is an important aspect of overall ecosystem functioning. However, single-celled organisms or microscopic metazoans are often ignored in studies of secondary production, despite being very abundant and possessing high mass-specific population growth rates, relative to the more widely studied larger taxa. Here, we focused on how life history parameters scale with body size of ciliates and meiofauna (body mass range from approximately 0.001 to 90 mg C/individual) and integrated experimental and survey data to calculate secondary production of these groups. First, we derived a single allometric scaling relationship between the intrinsic rate of population increase and body mass in a laboratory experiment. We then used this relationship to calculate secondary production for over 260 of these small species in the field, using survey data from two contrasting streams; one of which was nutrient rich, the other nutrient poor. Results from laboratory cultures showed that the scaling relationship between body mass and both daily intrinsic rate of population increase and generation time followed a power law. The relationship between body mass and annual secondary production was consistent in both streams, but the number of taxa was greater in the more productive site. Both ciliates and meiofauna had high rates of biomass production, with annual P/B ratios (production divided by biomass) for the whole assemblage exceeding 11 in both streams. We conclude that a large fraction of benthic production is overlooked when protozoans and microscopic metazoans are excluded from estimates of biomass turnover.

Trophic positioning of meiofauna revealed by stable isotopes and food web analyses

Despite important advances in the ecology of river food webs, the strength and nature of the connection between the meio- and macrofaunal components of the web are still debated. Some unresolved issues are the effects of the inclusion of meiofaunal links and their temporal variations on the overall river food web properties, and the significance of autochthonous and allochthonous material for these components. In the present study, we conducted analyses of gut content of macro- and meiofauna and stable isotope analyses of meiofauna to examine seasonal food webs of a chalk stream. The results of the gut content analyses, confirmed by the δ13 C signatures, revealed a seasonal shift from a dependence on autochthonous (biofilm) to allochthonous food sources. Here, we demonstrate that aggregating basal or meiofaunal species into single categories affects key web properties such as web size, links, linkage density, and predator-prey ratios. More importantly, seasonal variation in attributes characterized the entire web and these changes persist regardless of taxonomic resolution. Furthermore, our analyses evidenced discrete variations in δ15 N across the meiofauna community with a trophic structure that confirms gut content analyses, placing the meiofauna high in the food web. We, therefore, conclude that small-body-sized taxa can occur high in dynamic river food webs, questioning assumptions that trophic position increases with body size and that webs are static.