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24Freshwater aquifers are a major source of drinking water; they also possess unique 25 assemblages of organisms. However, little is known about the distributional drivers 26 of obligate groundwater organisms at the regional scale. We examine the 27 distribution and composition of stygobiont assemblages in a complex geological 28 setting and explore the relationship between groundwater fauna, hydrogeology and 29 water chemistry. In the study area we grouped similar geologies into five

Microniphargus leruthi Schellenberg, 1934 (Amphipoda: Niphargidae) was first described based on samples collected in Belgium and placed in a monotypic genus within the family Niphargidae. However, some details of its morphology as well as recent phylogenetic studies suggest that Microniphargus may be more closely related to Pseudoniphargus (Amphipoda: Pseudoniphargidae) than to Niphargus. Moreover, M. leruthi ranges over 1,469 km from Ireland to Germany, which is striking since only a few niphargids have confirmed ranges in excess of 200 km. To find out the phylogenetic position of M. leruthi and check whether it may be a complex of cryptic species, we collected material from Ireland, England and Belgium then sequenced fragments of the mitochondrial cytochrome c oxidase subunit 1 gene as well as of the nuclear 28S ribosomal gene. Phylogenetic analyses of both markers confirm that Microniphargus is closer to Pseudoniphargus than to Niphargus, leading us to reallocate Microniphargus to Pseudoniphargidae. We also identify three congruent mito-nuclear lineages present respectively in Ireland, in both Belgium and England, and in England only (with the latter found in sympatry at one location), suggesting that M. leruthi is a complex of at least three species with a putative centre of origin in England.

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The Chalk is an important water supply aquifer, yet ecosystems within it remain poorly understood. Boreholes (198) in seven areas of England (UK) were sampled to determine the importance of the Chalk aquifer as a habitat, and to improve understanding of how species are distributed. Stygobitic macro-invertebrates were remarkably common, and were recorded in 67 % of boreholes in unconcealed Chalk, although they were not recorded in Chalk that is concealed by low-permeability strata and thus likely to be confined. Most species were found in shallow boreholes (<21 m) and boreholes with deep (>50 m) water tables, indicating that the habitat is vertically extensive. Stygobites were present in more boreholes in southern England than northern England (77 % compared to 38 %). Only two species were found in northern England compared to six in southern England, but overall seven of the eight stygobitic macroinvertebrate species found in England were detected in the Chalk. Two species are common in southern England, but absent from northern England despite the presence of a continuous habitat prior to the Devensian glaciation. This suggests that either they did not survive glaciations in the north where glaciers were more extensive, or dispersal rates are slow and they have never colonised northern England. Subsurface ecosystems comprising aquatic macro-invertebrates and meiofauna, as well as the microbial organisms they interact with, are likely to be widespread in the Chalk aquifer. They represent an important contribution to biodiversity, and may influence biogeochemical cycles and provide other ecosystem services.