Current biodiversity patterns are considered largely the result of past climatic and tectonic changes. In an integrative approach, we combine taxonomic and phylogenetic hypotheses to analyze temporal and geographic diversification of epigean (Carychium) and subterranean (Zospeum) evolutionary lineages in Carychiidae (Eupulmonata, Ellobioidea). We explicitly test three hypotheses: 1) morphospecies encompass unrecognized evolutionary lineages, 2) limited dispersal results in a close genetic relationship of geographical proximally distributed taxa and 3) major climatic and tectonic events had an impact on lineage diversification within Carychiidae.

Though recent investigations have contributed substantially to our understanding of the Alpine-Dinaric radiation of the genus Zospeum Bourguignat, 1856, its southernmost member, Zospeum troglobalcanicum Absolon, 1916, has remained a taxonomic ghost. The assumed absence of type material, the insufficient original description, and the lack of new samples from its Western Balkan type locality have stymied further clarification. The recent discovery of a single syntype shell housed at the Natural History Museum Vienna now enables the first morphological assessment via 3D X-ray and SEM imaging. Based on this image data, different characters for assessing the southernmost members of the genus are determined and a lectotype is designated. Eleven allied species from 15 Western Balkan populations are described from museum material and recent sampling efforts: Z. amplioscutum Jochum & Ruthensteiner sp. nov., Z. biokovoense Jochum & Ruthensteiner sp. nov., Z. constrictum Jochum & Ruthensteiner sp. nov., Z. dubokidoense Jochum & Ruthensteiner sp. nov., Z. intermedium Jochum & Ruthensteiner sp. nov., Z. kolbae Jochum, Inäbnit, Kneubühler & Ruthensteiner sp. nov., Z. neuberti Jochum & Ruthensteiner sp. nov., Z. njegusiense Jochum & Ruthensteiner sp. nov., Z. njunjicae Jochum, Schilthuizen & Ruthensteiner sp. nov., Z. tortuosum Jochum & Ruthensteiner sp. nov. and Z. tumidum Jochum, Schilthuizen & Ruthensteiner sp. nov. One species, Z. kolbae, is described using DNA sequence data and one species, Z. simplex Inäbnit, Jochum & Neubert, 2021 for which DNA sequence data is already available, is supported by morphological data presented in this study. The DNA sequence dataset (COI, 16S and H3) is included here and implemented in the most recent phylogenetic reconstruction of the genus. A translation of Karel Absolon’s notes from the Balkan scientific expeditions is provided.

The Dinaric Karst is a global hotspot for subterranean diversity, with two distinct peaks of species richness in the northwest and southeast, and an area of a lower species richness in the central part. In this article, we present a species list and describe the ecological conditions of the Lukina jama–Trojama cave system, located in the central part of the Dinaric Karst. This cave system is the deepest and one of the most logistically challenging cave systems sampled so far in the Dinaric Karst. Repeated sampling resulted in a list of 45 species, including 25 troglobionts, 3 troglophiles, 16 stygobionts, and 1 stygophile. Most of the recorded species are endemic to the Velebit Mountain, while three species are endemic to the Lukina jama–Trojama cave system. Within the system, species richness peaks in the deepest third of the cave, most likely reflecting the harsh ecological conditions in the upper parts, including ice, cold winds, and occasional waterfalls. Milder and more stable deeper parts of the cave contain a rich subterranean species community, part of which is associated with two very distinct aquatic habitats, the cave hygropetric and the phreatic zone. The newly recognized hotspot of subterranean biodiversity in the central Dinaric Karst, which has emerged between the two known centers of biodiversity, further highlights the species richness in large cave systems, but also challenges the diversity patterns in the Dinaric Karst overall.