Interactions at surface-subterranean ecotones: structure and function of food webs within spring orifices.

Spring orifices are ecotones between surface and subterranean aquatic ecosystems. Invertebrates of different origins (e.g., surface, spring obligate, and subterranean) coexist in these spatially restricted environments, potentially competing for resources. However, processes that allow for population coexistence in these presumably low resource environments are not well understood. We examined invertebrate communities at two spring complexes in Texas, USA and assessed resource use and food web structure at spring orifices using stable isotopes of carbon (δ13C) and nitrogen (δ15N). Using bulk δ13C and δ15N of organisms and potential food sources, we elucidated dietary sources and found that invertebrate communities exhibited resource partitioning and contained two main food chains (periphyton versus terrestrial organic matter [OM]). In both spring complexes, several endemic spring orifice associated and subterranean taxa derived most of their diet from terrestrial OM. Analysis of compound-specific stable isotopes (i.e., δ13C of essential amino acids, EAAs) from two co-occurring elmid species indicated that the endemic spring orifice-associated species (Heterelmis comalensis) derived > 80% of its EAAs from bacteria, whereas the widespread surface species (Microcylloepus pusillus) derived its EAAs from a more equitable mix of bacteria, fungi, and algae. We additionally calculated niche overlap among of several taxonomically related groups (aquatic beetles and amphipods) that co-occur in spring ecotones and posterior probability estimates indicated little to no niche overlap among related species. Results indicate that invertebrates at subterranean-surface aquatic ecotones are partitioning food resources and highlight the importance of connections to riparian zones for persistence of several endemic invertebrates.