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The aquatic biodiversity of springs and groundwater systems of North Africa remains largely unexplored. In an earlier field survey of Tunisian springs, a new gastropod genus, Bullaregia, was discovered as a phylogenetically independent lineage of uncertain position within the family Hydrobiidae. Here, we provide taxonomic and phylogenetic assignments for three newly collected populations of hydrobiids from springs in northern Tunisia based on morphological, anatomical and genetic (mtCOI and 18S) data. Among these and specimens of Bullaregia, major differences were observed in male and female genitalia as well as in mtCOI sequences (divergence 8.0–9.1%). Based on these findings, we describe two new genera and three new species: Belgrandiellopsis chorfensis gen. et sp. nov., Belgrandiellopsis secunda gen. et sp. nov. and Biserta putealis gen. et sp. nov. In all our phylogenetic analyses, these three new species were well resolved as a monophyletic group together with Bullaregia tunisiensis. Unexpectedly, this clade emerged as sister to the European valvatiform genera Corbellaria and Kerkia and not to the recently discovered clade of groundwater, conchologically similar, species living in Bulgaria (Balkan Peninsula). These Tunisian species are each locally endemic and form part of a newly discovered clade which in future systematic studies could eventually be identified as a distinct hydrobiid subfamily.
The freshwater snail genus Mercuria is widely distributed in lowland waters across Western Europe, Northern Africa and the Mediterranean islands. Approximately two-thirds of the currently recognised species are described based on their shell morphology, which may vary within species due to biotic and abiotic factors. Recent molecular phylogenies that included numerous previously documented populations recovered 14 species clades, nine of which correspond to nominal species and five, to undescribed taxa. Here, we formally describe the five undescribed taxa as new species and provide morphological descriptions of the shell and other anatomical structures for three of the other inferred clades and for the species M. maceana to elucidate their taxonomic status and assess the utility of morphological characters for species delimitation in Mercuria. Taken together, the morphological and molecular evidence suggest new identifications and synonymies, having implications on the known geographic range of the studied species, including the type species M. similis. Anatomical measurements and geometric morphometric analysis of shell shape revealed no clear differentiation among the species analysed, predicting the importance of molecular data in elucidating the species diversity of the genus.
The species richness of the aquatic (non-marine) gastropod family Hydrobiidae Stimpson, 1865 reaches its peak in the European region partly because other areas, such as North Africa, remain to be extensively surveyed. Most of the hydrobiid species described in North Africa have been ascribed to the subfamilies Pseudamnicolinae, Hydrobiinae and the genus Mercuria. Little is known about the presence of other hydrobiid subfamilies. This study examines several specimens of gastropods collected from two springs in Tunisia. Based on a comprehensive literature review and rigorous anatomical and molecular comparisons with known species from North Africa, Europe and Asia Minor of similar morphology, we here describe a new genus, Bullaregia gen. nov., and new species Bullaregia tunisiensis gen. et sp. nov. Although the shell shape of this Tunisian species resembles that of European hydrobiid genera such as Belgrandiella, it differs in other anatomical structures (i.e., penis with a glandular strap-like lobe, and two seminal receptacles, SR1 pedunculate, SR2 sessile and less developed). Using maximum likelihood and Bayesian approaches, phylogenetic relationships inferred from mtCOI sequences point to an independent evolutionary lineage for this new taxon outside Belgrandiella and other known hydrobiid subfamilies.