Groundwater copepods: diversity patterns over ecological and evolutionary scales

Copepods are common components of the groundwater fauna, and greatly increase the diversity of groundwater communities. With more than 900 species/subspecies known from continental groundwaters, stygobiont copepods inhabit all kinds of aquifers (karstic, fissured, porous), as well as surface/subsurface ecotones (land/water and water/water). The polyhedral and varied structure of the stygohabitats is reflected in the surprising mixture of functional morphologies and habitat exploitations experienced by groundwater copepods. Morphological adaptations and specializations are discussed, as well as the chronology of their appearance in the evolutionary history of several taxa. Diversity patterns of copepod assemblages in groundwater are examined under both structural and functional profiles, as well as across a range of scales. Structure and function operate in an interactive, sometimes hierarchical ways, as well as scales. On the ecological scale, local heterogeneity and patchiness in geomorphic and hydrologic characteristics, as well as biotic interactions, are to be considered causal factors affecting the diversity patterns over a range of spatial and temporal scales. On the evolutionary scale, it is widely accepted that stygobiont copepods evolved from ancestors living in marine, freshwater and semiterrestrial environments. They gained access to the groundwater through major highways represented by the interstitial and the crevicular/karstic corridors. `Phylogenetic diversity' in groundwater copepod taxocoenoses is viewed as a heterogeneous assemblage of species belonging to different phylogenetic lineages, which entered groundwater at different times and by different ways.