Abstract
A revision of the subgenus Falsargutor Kryzhanovskij, 1983 of the genus Pterostichus Bonelli, 1810 is presented. Pterostichus (Falsargutor) dacatrai sp. nov. is described from northeastern Turkey. P. (F.) imeretinus Kryzhanovskij, 1989 is restored as a good species, P. (F.) iranianus Straneo, 1983 is raised to the species level. All known species of the subgenus are redescribed, annotated, photographed, mapped, and keyed. Both linear morphometrics and geometric morphometrics were employed to assess interspecific and intraspecific variation in shape and proportions. Both approaches are shown to be reliable techniques for taxonomic analysis with broadly similar resolution power. However, their effectiveness varies across species: Therefore, they can be effectively combined to explore the taxonomic structure of a group and to enhance diagnostic tools. Geometric morphometrics proved to be the most powerful tool for describing and quantifying shape differences. Both analyses revealed no geographic races in P. (F.) pseudopedius Reitter, 1887 and a slight west–east differentiation in P. (F.) ponticus Kirschenhofer, 1987. The geographic distribution of Falsargutor appears unique among Caucasian carabids. Members of the subgenus occur in two disjunct areas: mountains along the eastern Black Sea coast from the western extremity of the Western Caucasus to northeastern Turkey and in Mazandaran Province, northern Iran. The most primitive consubgeners are restricted to most ancient regional terrains: northeastern Pontides and Dzirula massif while the most deviated species occurs in the Western Caucasus. Species distribution models suggest a much broader distribution of Falsargutor in the past. A morphology-based phylogeny of the subgenus and related groups has been proposed, accompanied by a palaeogeographic reconstruction of its evolutionary history.
References
Afanasenkov A. R., Nikishin A. M. & Obukhov A. N. 2007. Eastern Black Sea Basin: geological structure and hydrocarbon potential. Moscow: Nauchny Mir, 172 pp. ISBN 978-589-176-457-6. (in Russian).
Avdeev B. & Niemi N. A. 2011. Rapid Pliocene exhumation of the central Greater Caucasus constrained by low-temperature thermochronometry, Tectonics, 30, TC2009, 1–16. https://doi.org/10.1029/2010TC002808
Belousov I. A. 1987. New species of carabids of the tribe Trechini (Coleoptera, Carabidae) from the Caucasus. I. Trechus liopleurus Chd. species group]. Revue d’entomologie de l’URSS, LXVI (3): 591–601. (in Russian).
Belousov I. A. 1990. New carabids of the tribe Trechini (Coleoptera, Carabidae) from the Caucasus. III. 18 new species and a new subspecies of the genus Trechus Clairv.) Revue d’entomologie de l’URSS, LXIX, 3: 609–632 (in Russian).
Belousov I. A. 1998. Le complexe générique de Nannotrechus Winkler du Caucase et de la Crimée (Сoleoptera, Carabidae, Trechini). Pentsoft, Series Faunistica, 8, 256 pp.
Belousov I. A. 1999. A new genus and species of cave dwelling trechine beetles from the West Caucasus (Coleoptera Carabidae Trechini). Advances in Carabidology (Papers Dedicated to the Memory of Prof. Dr. Oleg L. Kryzhanovskij). Edited by Alexandr Zamotajlov & Riccardo Sciaky. Krasnodar: MUISO Publishers, pp. 165–182.
Belousov I. A. & Sokolov I. M. 1996. Review of the Caucasian species of the subgenus Peryphanes Jeannel (Coleoptera: Carabidae: Bembidion). Stuttgarter Beiträge zur Naturkunde, Ser. A, 549: 1–40.
Bousquet Y. 1999. Supraspecific classification of the nearctic Pterostichini (Coleoptera: Carabidae). Fabreries, Supplément 9: 1–292.
Bousquet Y. 2017. Tribe Pterostichini Bonelli, 1810, pp. 675–755. In: Catalogue of Palaearctic Coleoptera. Vol. 1 (Löbl I. & Löbl D., editors). Archostemata – Myxophaga – Adephaga. Revised and Updated Edition. Leiden – Boston: Brill.
Boztuğ D., Jonckheere R., Wagner G. A. & Yeğingil Z. 2004. Slow Senonian and fast Palaeocene–Early Eocene uplift of the granitoids in the Central Eastern Pontides, Turkey: Apatite fission-track results. Tectonophysics, 382(3–4): 213–228. https://doi.org/10.1016/j.tecto.2004.01.001
Chaladze G., Kalatozishvili L., Janiashvili Z. & Bakuradze G. 2017. Pterostichus neilgaimani sp. nov., a new species of ground beetles (Coleoptera: Carabidae) from relict sacred grove in Eastern Georgia. Zootaxa 4329(1): 091–095. https://doi.org/10.11646/zootaxa.4329.1.6
Erwin T. L. 1985. The taxon pulse: a general pattern of lineage radiation and extinction among carabid beetles. In: Ball, G.E., ed., Taxonomy, Phylogeny, and Zoogeography of Beetles and Ants. Dr. W. Junk Publishers, Dordrecht, pp. 437–472.
Esin N. V., Yanko-Hombach V. V. & Esin N. I. 2018. Evolutionary mechanisms of the Paratethys Sea and its separation into the Black Sea and Caspian Sea. Quaternary International 465: 46–53. https://doi.org/10.1016/j.quaint.2016.06.019.
Eyüboğlu Y., Santosh M., Dudaş F. O. & Akaryalı E. 2011. Transition from shoshonitic to adakitic magmatism in the eastern Pontides, NE Turkey: Implications for slab window melting. Gondwana Research 19(2): 413–429. https://doi.org/10.1016/j.gr.2010.12.003
Hijmans R. 2023. terra: Spatial Data Analysis. R package version 1.7–55. Available from: https://CRAN.R-project.org/package=terra (accessed on 18 December 2025)
Hollister J. W. 2023. elevatr: Access Elevation Data from Various APIs. R package version 0.99.0. Available from: https://CRAN.R-project.org/package=elevatr (accessed on 18 December 2025)
Jeannel R. 1943. Les fossiles vivants des cavernes. Paris: Gallimard, L’Avenir de la science. 321 pp.
Kirschenhofer E. 1987. Untersuchungen über Pterostichus-Arten aus Kleinasien, dem Kaukasus und Zentralasien (Col. Carabidae: Pterostichinae). Folia Entomologica Hungarica. Rovartani Közlemények (S.N.) 48: 45–85.
Kryzhanovskij O. L. 1988. Review of the Caucasian species of the subgenus Oreoplatysma Jacobson, 1906 (Coleoptera: Carabidae: Pterostichus). Systematics of Insects and Mites, Proceedings of the All-Union Entomological Society 1988: 12–21. (in Russian)
Kryzhanovskij O. L. 1989. New and little-known species of the tribe Pterostichini (Coleoptera, Carabidae) from the Caucasus. Proceedings of the Zoological Institute, Leningrad, 1989, Vol. 208: 14–22 (in Russian).
Kryzhanovskij O. L. & Abdurakhmanov G. M. 1983. New and little known species of supertribe Pterostichitae (Coleoptera, Carabidae) from the Caucasus. Revue d’Entomologie de l’URSS, LXII, 3: 529–537. (in Russian).
Kryzhanovskij O. L., Belousov I. A., Kabak I. I., Kataev B. M., Makarov K. V. & Shilenkov V. G. 1995. A Checklist of the ground-beetles of Russia and adjacent Lands (Insecta, Coleoptera, Carabidae). Pentsoft, Series Faunistica, 3, 271 pp.
Lehner B. & Grill G. 2013. Global river hydrography and network routing: baseline data and new approaches to study the world’s large river systems. Hydrological Processes 27(15): 2171–2186.
Mosar J., Mauvilly J., Koiava K., Gamkrelidze I., Enna N., Lavrishev V. & Kalberguenova V. 2022. Tectonics in the Greater Caucasus (Georgia – Russia): From an intracontinental rifted basin to a doubly verging fold-and-thrust belt. Marine and Petroleum Geology 140(2022): 105630 pp. 1–14. https://doi.org/10.1016/j.marpetgeo.2022.105630
Muilwijk J., Kataev B.M. & Wrase D.W. 2024. Review of the subgenus Lyrothorax of the genus Pterostichus (Coleoptera: Carabidae) in the Western Palaearctic, with descriptions of two new species. Zootaxa 5468 (2): 255–274. https://doi.org/10.11646/zootaxa.5468.2.2
Nagy L. & Grabherr G. 2009. Chapter 6: Alpine terrestrial habitats and community types/assemblages, pp. 117–172. In: Nagy, L. & Grabherr, G. The Biology of Alpine Habitats. Oxford University Press, Oxford. https://doi.org/10.1093/oso/9780198567035.003.0006
Nakhutsrishvili G., Zazanashvili N., Batsatsashvili K. & Montalvo Mancheno C. S. 2015. Colchic and Hyrcanian forests of the Caucasus: similarities, differences and conservation status. Flora Mediterranea 25 (Special Issue): 185–192. https://doi.org/10.7320/FlMedit25SI.185
Popov S. V., Akhmetiev M. A., Lopatin A. V., Bugrova E. M., Sychevskaya E. K., Shcherba I. G., Andreeva-Grigorovich A. S., Zaporozhets N. I., Nikolaeva I. A., Kopp M. L. 2009. Paleogeografia i biogeografia bassejnov Paratetisa. Chast’ 1: Pozdnyj eocen – rannij miocen. Proceedings of the Paleontological Institute, Russian Academy of Sciences, Vol. 292. Moscow: Nauchnyi Mir, 2009. 178 pp. [Paleogeography and Biogeography of Paratethys Basins. Part 1: Late Eocene—Early Miocene]. (in Russian).
Pupier R. 2011. Tribu Pterostichini – Subgen. Cryobius Chaudoir, 1838. In: Faune de France (Coulon G, Pupier R, Queinnec E, Ollivier E, Richoux P, editors). Coléoptères Carabiques. Faune de France, Paris, 443–455.
Reitter E. 1887. Neue Coleopteren aus Europa, den angrenzenden Ländern und Sibirien, mit Bemerkungen über bekannte Arten. Dritter Theil. Deutsche Entomologische Zeitschrift 31: 241–288.
Sasakawa K. 2022. Possible differences in effects on ejaculate morphology between shape and size components of the intromittent membranous part of insect male genitalia: Phylogenetic comparative analyses in Carabidae. Frontiers in Ecology and Evolution 10: 1032378: pp. 1–9.
Sasakawa K. & Kubota K. 2007. Phylogeny and Genital Evolution of Carabid Beetles in the Genus Pterostichus and Its Allied Genera (Coleoptera: Carabidae) Inferred from Two Nuclear Gene Sequences. Annals of the Entomological Society of America 100(2): 100–109. https://doi.org/10.1603/0013-8746(2007)100[100:PAGEOC]2.0.CO;2
Sedlmeier J. E. & Faille A. 2022. First insights into the phylogeny of the subgenus Cryobius Chaudoir, 1838 (Coleoptera: Carabidae: Pterostichus). Arthropod Systematics & Phylogeny 80: 523–539. https://doi.org/10.3897/asp.80.e84114
Sciaky R. 1996. New taxa and new synonyms among Pterostichinae from Asia (Coleoptera Carabidae). Entomofauna, Zeitschrift Für Entomologie 17(29): 429–440.
Shatilova I., Mchedlishvili N., Rukhadze L. & Kvavadze E. 2011. The history of the flora and vegetation of Georgia (South Caucasus). Georgian national museum, Institute of paleobiology. Tbilisi: 200 pp.
Straneo S. L. 1983. Nuovi Pterostichini Asiatici (Coleoptera Carabidae). Bollettino della Società entomologica Italiana 115(1–3): 17–22.
Vincent S. J., Carter A., Lavrishchev V. A., Rice S. P., Barabadze T. G. & Hovius N. 2011. The exhumation of the western Greater Caucasus: a thermochronometric study. Geological Magazine 148(1): 1–21. https://doi.org/10.1017/S0016756810000257
Wickham H., Averick M., Bryan J., Chang W., McGowan L.D., François R., Grolemund G., Hayes A., Henry L., Hester J., Kuhn M., Pedersen T.L., Miller E., Bache S.M., Müller K., Ooms J., Robinson D., Seidel D.P., Spinu V., Takahashi K., Vaughan D., Wilke C., Woo K., Yutani H. 2019. “Welcome to the tidyverse.” Journal of Open Source Software 4(43): 1686. https://doi.org/10.21105/joss.01686
