The role of habitat complexity has been widely neglected in the study of meiofaunal community patterns. We studied the intertidal nematode community of a structurally complex macrotidal beach exhibiting contrasting microhabitats (sandbars and runnels) to understand the influence of environmental gradients and habitat heterogeneity in the community structure. We tested whether topographical complexity affected (1) the zonation pattern in terms of abundance and diversity, and (2) local diversity by promoting compartmentalization into distinct faunal groups. Our analyses revealed three major faunal assemblages along the exposure gradient associated to differences in mean grain size and chlorophyll a. Diversity patterns involved a mid-intertidal peak, consistent with the intermediate disturbance hypothesis, and another peak at the limit with the subtidal region, consistent with the transition zone. These results highlight the predominance of environmental gradients in establishing intertidal zonation. However, microhabitats differed in environmental conditions and possessed significantly distinct nematofaunal communities. Runnels featured higher levels of taxonomic and functional diversity, many unique genera, and the community differed from the assemblage at the limit to the subtidal, stressing their role as distinct microhabitats. The nematofauna of the structurally complex beach was more diverse than the one from a homogeneous beach nearby, supporting the hypothesis that structural heterogeneity promotes diversity by compartmentalization and highlighting the importance of microhabitats in the assessment of biodiversity. Contrary to previous predictions, our results indicate potentially high regional marine nematode diversity in the Upper Gulf of California.

The deep sea is Earth's largest habitat but little is known about the nature of deep-sea parasitism. In contrast to a few characterized cases of bacterial and protistan parasites, the existence and biological significance of deep-sea parasitic fungi is yet to be understood. Here we report the discovery of a fungus-related parasitic microsporidium, Nematocenator marisprofundi n. gen. n. sp. that infects benthic nematodes at methane seeps on the Pacific Ocean floor. This infection is species-specific and has been temporally and spatially stable over 2 years of sampling, indicating an ecologically consistent host-parasite interaction. A high distribution of spores in the reproductive tracts of infected males and females and their absence from host nematodes' intestines suggests a sexual transmission strategy in contrast to the fecal-oral transmission of most microsporidia. N. marisprofundi targets the host's body wall muscles causing cell lysis, and in severe infection even muscle filament degradation. Phylogenetic analyses placed N. marisprofundi in a novel and basal clade not closely related to any described microsporidia clade, suggesting either that microsporidia-nematode parasitism occurred early in microsporidia evolution or that host specialization occurred late in an ancient deep-sea microsporidian lineage. Our findings reveal that methane seeps support complex ecosystems involving interkingdom interactions between bacteria, nematodes, and parasitic fungi and that microsporidia parasitism exists also in the deep-sea biosphere.