The South American genus Cnemalobus (Coleoptera: Carabidae: Cnemalobini): phylogeny and biogeographic analysis with the description of four new species from extra-Andean Patagonian mountains

The carabid beetle Cnemalobus Gu & eacute;rin-M & eacute;n & eacute;ville, 1838 inhabits high- and lowland grasslands of southern South America. The highest diversity is found in the Patagonian Steppe, where distribution patterns are associated with latitude and elevation. Northern Patagonia, a large volcanic region with a complex geoclimatic history, exhibits elevated grades of endemism. However, a great deal remains unknown regarding diversification and biogeographical patterns for most of the endemic groups. We describe new Cnemalobus species restricted to isolated volcanoes from these extra-Andean mountain systems. We assess the phylogenetic relationships by updating the phylogeny of the genus and conduct a Bayesian binary Markov chain-Monte Carlo (MCMC) analysis on the resulting phylogenetic tree to discuss the biogeographical distribution patterns. We also provide a taxonomic key to all currently known species of Cnemalobus from the Patagonian Steppe. Our phylogenetic analysis supports the monophyly of the new species Cnemalobus tromen sp. nov., Cnemalobus silviae sp. nov., Cnemalobus aucamahuida sp. nov. and Cnemalobus domuyo sp. nov. grouped with C. diamante and C. nevado, referred to as the 'Extra-Andean' mountain lineage. Biogeographical analysis recognises vicariant events as the most plausible explanation for the allopatric distributions of the new species. We hypothesise that these vicariant events could be related to climatic barriers that likely promoted speciation processes by generating geographical isolation in ancestral populations. Our findings contribute significantly to the biogeographical understanding of the Patagonian volcanic region, prompting new inquiries to unravel the speciation processes of the endemic biota in extra-Andean mountain systems.