The Diversity of Freshwater Stygobiotic Crustaceans in the Republic of North Ossetia–Alania Provides New Evidence for the Existence of an Ancient Glacial Refugium in the North Caucasus Region

Abstract

A review and partial revision of the diversity of freshwater stygobiotic crustaceans in the territory of the Republic of North Ossetia–Alania, in the North Caucasus, is presented here. Previously, two species of the genus Proasellus Dudich, 1925 (Isopoda, Asellidae), P. uallagirus Palatov & Sokolova, 2020 and P. irystonicus Palatov & Sokolova, 2020, and one species of the genus Niphargus Schiödte, 1849 (Amphipoda, Niphargidae), N. alanicus Marin & Palatov, 2021, were described from the hyporhean/underground habitats (hyporhea) in the area. However, further research using an integrative approach has revealed that only a single species of the genus Proasellus (P. uallagirus) is actually widely distributed in the hyporhean riverbed habitats in the area, while the diversity of the genus Niphargus is higher than previously known. Six more new Niphargus species— namely, N. ardonicus sp. nov., N. sadonicus sp. nov., N. fiagdonicus sp. nov., N. tschertschesovae sp. nov., N. osseticus sp. nov. and N. zeyensis sp. nov., were discovered from the various hypogean underground water sources (i.e., springs and seeps) and are described in this article. Their phylogenetic relationships with their congeners, as well as their ecology and known distribution, are discussed. Furthermore, molecular genetic analysis, with an interpretation of the estimated divergence time, suggests that the studied hyporheic/stygobiotic crustaceans started to diverge from related European and Balkan sister species during the Late Miocene, approximately 8–5.8 Mya, with the reduction in the Paratethys and the uplifting of the Caucasus Mountains. Local speciation was led by local geological processes and karst fragmentation during the Late Pliocene and Early Pleistocene periods, starting around 5.3 Mya. The obtained data suggest that the mountainous area of the North Ossetia–Alania could be considered as a post-Pliocene glacial refugium for subterranean and stygobiotic fauna—the first known for the North Caucasus region.