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The past climatic changes caused repeated distribution shifts within insect populations leading to a highly diverse fauna in the mountain regions, which have acted as a refuge for many groups. There, some taxa have adapted to high altitudes and cold climatic conditions. One of those is the highly diverse and Holarctic subgenus Cryobius Chaudoir, 1838 (Carabidae: Pterostichus) including both locally and widely distributed species. Isolated and morphologically divergent populations of the same species led to the description of many subspecies. Until now, there has been no comprehensive work concerning the phylogeny of Cryobius, and genetic data on this taxon are sparse. This study is the first to provide insights into the molecular phylogeny of this subgenus, focusing on species from the Pyrenean and Cantabrian mountain systems. Cryobius specimens were sequenced targeting mitochondrial and nuclear genes. A molecular phylogeny was then built, merging the new data with genetic data from online public databases. All species of Cryobius included in this study form a monophyletic clade within Pterostichus. The synonymy of the two former taxa Pyreneorites and Haptoderus with Cryobius is confirmed by this study. Cryobius of the Pyreneo-Cantabrian area are closely related. Moreover, several well-supported clades of local species were found. The results further indicate a relation between Nearctic and Eastern Palearctic Cryobius, in agreement with the theory of faunal and floral colonization of North America via the Bering land bridge.

Genus Speoplanes Müller Apholeuonus (Speoplanes) Müller, 1911: 2, type species: Apholeuonus (Speoplanes) giganteus Müuller, 1911, (original designation). Speoplanes Müller: Jeannel, 1924: 381 (monography); Pretner, 1968: 35 (catalogue); Pretner, 1973: 184 (catalogue); Guéorguiev, 1976: 117 (phylogeny, classification); Newton, 1998: 124 (catalogue); Perreau, 2000: 220 (catalogue); Perreau, 2015: 220 (catalogue); Hlaváč, Perreau & Čeplík, 2017: 120 (catalogue)., Published as part of Mizerakis, Vangelis, Hlaváč, Peter, Čeplík, Dávid, Jalžić, Branko & Faille, Arnaud, 2022, New insights into the subterranean genus Speoplanes Müller, 1911 (Coleoptera, Leiodidae, Cholevinae) from the Dinarides, pp. 581-594 in Zootaxa 5214 (4) on page 590, DOI: 10.11646/zootaxa.5214.4.6, http://zenodo.org/record/7397860, {"references":["Muller, J. (1911) Neue Hohlenkafer aus dem osterreichischen Karst. Wiener Entomologische Zeitung, 30 (1), 1 - 4.","Jeannel, R. (1924) Monographie des Bathysciinae. Archives de Zoologie experimentale et generale, 63 (1), 436.","Pretner, E. (1968) Catalogus Faunae Jugoslaviae. Vol. III / 6. Coleoptera: Catopidae: Bathysciinae. Academia scientiarum et artium Slovenica, Ljubljana, 59 pp.","Pretner, E. (1973) Koleopteroloska fauna pecina i jama Hrvatske: s historijskim pregledom istrazivanja. Jugoslavenska Akademija Znanosti i Umjetnosti, 8 / 6, 101 - 239.","Gueorguiev, V. B. (1976) Recherches sur la taxonomie, la classification et la phylogenie des Bathysciinae (Coleoptera: Catopidae). Razprave Sazu, 19 (4), 91 - 129.","Newton, A. F. (1998) Phylogenetic problems, current classification and generic catalogue of world Leiodidae (including Cholevinae). In: Phylogeny and Evolution of Subterranean and Endogean Cholevidae (= Leiodidae Cholevinae). Proceedings of XX International Congress of Entomology, Firenze, 1996. Atti del Museo Regionale di Scienze Naturali, Torino, pp. 41 - 178.","Perreau, M. (2000) Catalogue des Coleopteres Leiodidae Cholevinae et Platypsyllinae. Memoires de la Societe Entomologique de France, 4, 1 - 460.","Perreau, M. (2015) Leiodidae. In: Lobl, I. & Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Hydrophiloidea - Staphylinoidea. Revised and Updated Edition. Brill, Leiden and Boston, pp. 180 - 290.","Hlavac, P., Perreau, M. & Ceplik, D. (2017) The subterranean beetles of the Balkan peninsula. Czech University of Life Sciences, Faculty of Forestry and Wood Sciences, Praha, 267 pp."]}

Speoplanes biocovensis stat. nov. Müller, 1934 Speoplanes giganteus biocovensis Müller, 1934: 180. Speoplanes giganteus biocovensis Müller, 1934: Pretner, 1968: 36; Perreau, 2000: 220; Perreau, 2015: 220; Hlaváč, Perreau & Čeplík, 2017: 120. Type locality: Dalmatia centralis, in antro glacialis Japaga u Zelenim docima (1300 m) Montis Biokovo (= Crna Ledenica, Zeleni doci) (Bedek et al 2006) Distribution: Croatia: Central Dinaric Alps, glacial caves of mount Biokovo, Mali Šibenik and Rašćane Gornje.

Speoplanes giganteus (Müller 1911) Apholeuonus (Speoplanes) giganteus Müller, 1911: 2. Speoplanes giganteus Müller: Jeannel, 1914: 50; Jeannel, 1924: 384; Newton, 1998: 124. Speoplanes giganteus giganteus Müller, 1911: Pretner, 1968: 35; Perreau, 2000: 221; Perreau, 2015: 220; Hlaváč, Perreau & Čeplík, 2017: 120. Type locality: Zentraldalmatien, in einer Höhle im Mosorgebirge (= Ledenica pod Jabukovcem) (Bedek et al 2006) Distribution: Croatia: Central Dinaric Alps, glacial caves of mount Mosor., Published as part of Mizerakis, Vangelis, Hlaváč, Peter, Čeplík, Dávid, Jalžić, Branko & Faille, Arnaud, 2022, New insights into the subterranean genus Speoplanes Müller, 1911 (Coleoptera, Leiodidae, Cholevinae) from the Dinarides, pp. 581-594 in Zootaxa 5214 (4) on page 590, DOI: 10.11646/zootaxa.5214.4.6, http://zenodo.org/record/7397860, {"references":["Muller, J. (1911) Neue Hohlenkafer aus dem osterreichischen Karst. Wiener Entomologische Zeitung, 30 (1), 1 - 4.","Jeannel, R. (1914) Coleopterorum catalogus, Silphidae: Subfam. Bathysciinae. Pars 60. W. Junk, Berlin, 62 pp.","Jeannel, R. (1924) Monographie des Bathysciinae. Archives de Zoologie experimentale et generale, 63 (1), 436.","Newton, A. F. (1998) Phylogenetic problems, current classification and generic catalogue of world Leiodidae (including Cholevinae). In: Phylogeny and Evolution of Subterranean and Endogean Cholevidae (= Leiodidae Cholevinae). Proceedings of XX International Congress of Entomology, Firenze, 1996. Atti del Museo Regionale di Scienze Naturali, Torino, pp. 41 - 178.","Pretner, E. (1968) Catalogus Faunae Jugoslaviae. Vol. III / 6. Coleoptera: Catopidae: Bathysciinae. Academia scientiarum et artium Slovenica, Ljubljana, 59 pp.","Perreau, M. (2000) Catalogue des Coleopteres Leiodidae Cholevinae et Platypsyllinae. Memoires de la Societe Entomologique de France, 4, 1 - 460.","Perreau, M. (2015) Leiodidae. In: Lobl, I. & Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Hydrophiloidea - Staphylinoidea. Revised and Updated Edition. Brill, Leiden and Boston, pp. 180 - 290.","Hlavac, P., Perreau, M. & Ceplik, D. (2017) The subterranean beetles of the Balkan peninsula. Czech University of Life Sciences, Faculty of Forestry and Wood Sciences, Praha, 267 pp.","Bedek, J., Matocec, S. G., Jalzic, B., Ozimec, R. & Stamol, V. (2006) Catalogue of Cave type localities of Croatian fauna. Natura Croatica, 15 (Supplement 1), 1."]}