Abstract
The Aciculidae is a family of terrestrial operculate snails with a western Palaearctic distribution. Historically three Acicula species were reported from the Caucasus Mountains, which represents the eastern edge of the family's distribution: A. limbata, A. moussoni and A. parcelineata, all from a few localities. We examined extensive material from the entire range of the Caucasus Mountains by means of classic morphology-based taxonomy and geometric morphometries. We arrived at the following conclusions: (1) At least three species inhabit the Caucasus Mountains, two of which are problematic taxonomically and regarding their nomenclature. (2) The two syntypes of A. moussoni have been lost, and both lectotype designations by the former authors were invalid. Therefore, we designate a neotype for A. moussoni. (3) Acicula limbata was described from Lower Miocene deposits of the Czech Republic. Although the fossil Central European and extant Caucasian shells differ only slightly from one another, it is highly unlikely that they would belong to the same species. Therefore, we suggest using A. limbata only for the Czech fossil species and A. moussoni for the Caucasian extant species. (4) In multiple samples, the generally smaller Acicula parcelineata was found syntopic with the generally larger A. moussoni. Therefore, the two are possibly distinct species. However, it is very difficult to distinguish, and the problem of A. moussoni and A. parcelineata needs further attention. (5) A species with a bullet (or torpedo)-shaped shell and an opisthocline apertural profile from the northern Caucasus region has been treated as A. moussoni by previous authors, but it is distinct from the other two Acicula species by consistent conchological characters and was also found with them in syntopy, and is described here as Acicula telum Páll-Gergely, n. sp.
References
Boeters, H.D., Gittenberger, E. & Subai, P. (1985) Eine neue Gattung der Aciculidae (Gastropoda: Prosobranchia) mit zwei neuen Arten. Basteria, 49 (1/3), 59–63.
Boeters, H.D., Gittenberger, E. & Subai, P. (1989) Die Aciculidae (Mollusca: Gastropoda Prosobranchia). Zoologische Verhandelingen, 252, 1–234.
Boettger, O. (1879) Kaukasische Mollusken, gesammelt von Herrn Hans Leder in Paskau. Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 6 (1), 1–42, pl. 1.
Boettger, O. (1880a) Kaukasische Mollusken gesammelt von Herrn Hans Leder, z. Z. in Tiflis. Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 7 (2), 109–150, pl. 4.
Boettger, O. (1880b) Armenische und transkaukasische Mollusken, aus einer Sendung des Hrn. Dr. G. Sievers in Tiflis. Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 7 (2), 151–161, pl. 5.
Boettger, O. (1883) Siebentes Verzeichniss von Mollusken der Kaukasusländer nach Sendungen des Hrn. Hans Leder, z. Z. in Helenendorf bei Elisabetpol (Transkaukasien). Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 10, 135–200, pls. 4–7.
Bookstein, F.L. (1991) Morphometric Tools for Landmark Data Geometric and Biology. Cambrige University, Cambrige, XVIII + 435 S. https://doi.org/10.1017/CBO9780511573064
Clessin, S. (1881) Beitrag zur Kenntniss der Molluskenfauna des Caucasus. Malakozoologische Blätter, Neue Folge, 3, 129–135.
Clessin, S. (1911) Neue Acme-Arten. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 43 (4), 165–167.
Conde-Padin, P., Grahame, J.W. & Rolán-Alvarez, E. (2007) Detecting shape differences in species of the Littorina saxatilis complex by morphometric analysis. Journal of Molluscan Studies, 73, 147–154. https://doi.org/10.1093/mollus/eym009
Dance, S.P. (1986) A History of Shell Collecting. E. J. Brill, Leiden, 265 pp.
Domokos, T., Deli, T., Varga, A., Flaviu-Crişan, H., Balashov, I., Gheoca, V., Biatov, A., Szappanos, B. & Páll-Gergely, B. (2018) Distribution of Drobacia banatica (Rossmässler, 1838) over time and space (Gastropoda: Stylommatophora: Helicidae). Brukenthal Acta Musei, 13 (3), 459–484.
Draparnaud, J.P.R. (1801) Tableau des mollusques terrestres et fluviatiles de la France. Renaud, Montpellier and Bossange, Masson & Besson, Paris, 116 pp. https://doi.org/10.5962/bhl.title.13180
Egorov, R.V. & Greíe, C. (2003) Treasure of Russian Shells. Vol. 6. Terrestrial Pectinibranchia: Helicinidae, Cyclophoridae, Diplommatinidae, Cochlostomatidae, Aciculidae, Tateidae, Terrestribythinellidae, Pomatiidae. Colus-Doverie, Moscow, 43 pp.
Giokas, S., Páll-Gergely, B. & Mettouris, O. (2014) Nonrandom variation of morphological traits across environmental gradients in a land snail. Evolutionary Ecology, 28, 323–340. https://doi.org/10.1007/s10682-013-9676-5
Gittenberger, E. (1990) Frustrating facts about area cladistics and species individuality. Bijdragen tot de dierkunde, 60 (3–4), 151–154.
Gray, J.E. (1850) [Descriptions of taxa] In: Baird, W. & Gray, J.E., Nomenclature of molluscous animals and shells in the collection of the British Museum. Part I. Cyclophoridae. British Museum, London, pp. 1–69.
Haase, M. & Misof, B. (2009) Dynamic gastropods: stable shell polymorphism despite gene flow in the land snail Arianta arbustorum. Journal of Zoological Systematics and Evolutionary Research, 47, 105–114. https://doi.org/10.1111/j.1439-0469.2008.00488.x
Hartmann, J.D.W. (1821) System der Erd- und Flußschnecken der Schweiz. Mit vergleichender Aufzählung aller auch in den benachbarten Ländern, Deutschland, Frankreich und Italien sich vorfindenden Arten. In: Steinmüller, J.R. (Ed.), Neue Alpina. Eine Schrift der Schweizerischen Naturgeschichte, Alpen- und Landwirthschaft gewiedmet. Erster Band. Steiner, Winterthur, pp. 194–268, pls. 1–2.
Harzhauser, M., Neubauer, T.A., Georgopoulou, E. & Harl, J. (2014) The Early Miocene (Burdigalian) mollusc fauna of the North Bohemian Lake (Most Basin). Bulletin of Geosciences, 89 (4), 819–908, 18 figs., 1 table. https://doi.org/10.3140/bull.geosci.1503
Hesse, P. (1920) Einige Bemerkungen über die Familie Acmidae nebst Verzeichnis der Arten. Archiv für Molluskenkunde, 52 (2), 78–89.
Horsák, M., Juřičková, L. & Picka, J. (2013) Molluscs of the Czech and Slovak Republics. Kabourek, Zlín, 264 pp. [in Czech and English]
IBM (2018) IBM SPSS Statistics for Windows, Version 25.0. IBM Corp, Armonk, NY.
Jackiewicz, M. (1974) Krajowe gatunki rodziny Aciculidae (Mollusca, Prosobranchia). Fragmenta Faunistica, 19 (17), 445–454. https://doi.org/10.3161/00159301FF1974.19.17.445
Javelidze, G. (1972) Sakartvelos khmeletis moluskebis sarkvevI [Identification keys for Georgian Terrestrial Molluscs]. Tbilisi, Tbilisis Universit'et'is. [142 pp. in Georgian]
Kalitina, Z.I. (1954) K izucheniyu nazemnykh mollyuskov tsentralnogo Kavkaza i vostochnogo Predkavkazya (Fauna, ekologiya i rol v rasprostranenii diklotselioza) [To the knowledge of the terrestrial molluscs of central and eastern Ciscaucasia (Fauna, ecology and role in disseminating dicroceliasis)]. Akademiya nauk SSSR, Zoologicheskiy institut, Leningrad. [unknown pagination]
Kalitina, Z.I. (1958) K izucheniyu zonalnogo raspredeleniya nazemnykh mollyuskov Severnykh sklonov Tsentralnogo Kavkaza i Vostochnogo Predkavkasya. Zdravookhranenie i meditsina v Severnoy Osetii, 7 (1), 157–181.
Kalitina, Z.I. & Botcharova, M.M. (1971) Molljuski vypasov i skotoprogonnyh putej kolhozov Severo-Osetinskoj ASSR [Molluscs of pastures and of the roads leading to pastures of collective farms on North Ossetian ASSR]. In: Likharev, I.M. (Ed.), Molluscs. Trends, Methods and some results of their investigation. These and Communications. Academy of Sciences USSR, Zoological Institute, Fourth meeting of the investigation of molluscs, Leningrad, pp. 111–112. [in Russian]
Kantor, Y.I. & Sysoev, A.V. (2005) [Catalogue of molluscs of Russia and adjacent countries]. KMK Scientific Press, Moskva, 627 pp.
Kijashko, P.V. (2000) Nazemnye mollyuski Lagonakskogo nagorya (Severo-zapadnyy Kavkaz) [On the Molluscs of Lagonaki mountains (Northwestern Caucasus)]. Avtoreferat, St.-Petersburg. [unknown pagination, in Russian]
Klingenberg, C.P. (2011) MorphoJ: An integrated software package for geometric morphometrics. Molecular Ecology Resources, 11 (2), 353–357. https://doi.org/10.1111/j.1755-0998.2010.02924.x
Kobelt, W. (1894–1896) Iconographie der Land- & Süsswasser-Mollusken mit vorzüglicher Berücksichtigung der europäischen noch nicht abgebildeten Arten. Neue Folge. 7. Bd. C.W.Kreidel's Verlag, Wiesbaden. [unknown pagination]
Lezhawa, G.I. (1966) Nazemnie mollyuski [Terrestrial mollusca]. Fauna bespozvonochnyh Trialetskogo khrebta [Invertebrate fauna of the Trialeti mountains], 1966, 9–20.
Lezhawa, G.I. (1973) Khmeletis moluskebi [Terrestrial molluscs]. Sakartvelos tskhovelta samkaro IV.Tbilisi, Metsniereba. [pp. 20–68, in Georgian]
Lika, R., Deli, T., Paparisto, A., Erőss, Z.P. & Fehér, Z. (2021) Contribution to the taxonomy and distribution of Aciculidae (Mollusca: Gastropoda: Caenogastropoda) in Albania and the surroundings. Zootaxa, 5004 (1), 131–150. https://doi.org/10.11646/zootaxa.5004.1.5
Moquin-Tandon, A. (1855–1856) Histoire naturelle des mollusques terrestres et fluviatiles de France. Tome Premier (1, 2, 3) & TomeSsecond (4, 5 & 6). K.-B. Baillière, Paris, pp. i–viii + 1–14, atlas: 1–16, pls. 1–9 (12 April 1855), pp. 145–256, atlas: 17–32, pls. 10–18 (4 May 1855), pp. 257–416 Atlas: 33–48, pls. 19–27 (1 August 1855), pp: i–iii + 1–176, atlas: 49–64, pls. 28–36 (10 September 1855), pp. 177–368, atlas: 65–80, pls. 37–45 (2 January 1856), pp. 369–646, atlas: i–iii + 81–92, pls. 46–54 (9 April 1856).
Mumladze, L., Asanidze, Z., Walther, F. & Hausdorf, B. (2017) Beyond elevation: testing the climatic variability hypothesis vs. Rapoport’s rule in vascular plant and snail species in the Caucasus. Biological Journal of the Linnean Society, 121, 753–763. https://doi.org/10.1093/biolinnean/blx027
Neiber, M.T., Bikashvili, A., Bananashvili, G., Shubashishvili, A., Japoshvili, B., Walther, F. & Mumladze, L. (2021) Continental molluscs collected during the second Georgian-German BioBlitz 2019 in Stepantsminda, Georgia. Mitteilungen der Deutschen Malakozoologischen Gesellschaft, 104, 23–36.
Nevill, G. (1880) On the land-shells, extinct and living, of the neighbourhood of Menton (Alpes Maritimes); with descriptions of a new genus and of several new species. Proceedings of the Zoological Society of London, 1880, 94–142, pls. 13 + 14.
Páll-Gergely, B. & Grego, J. (2022) A Georgian and an Iranian new species of Renea G. Nevill, 1880 enormously extend the genus’s distribution (Gastropoda: Caenogastropoda: Aciculidae). Zootaxa, 5188 (6), 596–600. https://doi.org/10.11646/zootaxa.5188.6.7
Páll-Gergely, B., Kornilios, P. & Giokas, S. (2012) Higher than anticipated diversity within an Albinaria species (Gastropoda, Pulmonata, Clausiliidae) in Southern Turkey. Journal of Biological Research, 18, 345–352.
Pfenninger, M. & Magnin, F. (2001) Phenotypic evolution and hidden speciation in Candidula unifasciata ssp (Helicellinae, Gastropoda) inferred by 16S variation and quantitative shell traits. Molecular Ecology, 10, 2541–2554. https://doi.org/10.1046/j.0962-1083.2001.01389.x
Pokryszko, B.M., Cameron, R.A.D., Mumladze, L. & Tarkhnishvili, D. (2011) Forest snail faunas from Georgian Transcaucasia: patterns of diversity in a Pleistocene refugium. Biological Journal of the Linnean Society, 102, 239–250. https://doi.org/10.1111/j.1095-8312.2010.01575.x
Reuss, A.E. (1861) Die fossilen Mollusken der tertiären Süßwasserkalke Böhmens. Sitzungsberichte der mathematisch-naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, 42, 55–85.
Rohlf, F.J. (2006) tpsDig, Digitize Landmarks and Outlines, Version 2.0. Department of Ecology and Evolution, State University of New York at Stony Brook.
Rohlf, F.J. & Marcus, L.F. (1993) A revolution in morphometrics. Trends in Ecology & Evolution, 8, 129–132. https://doi.org/10.1016/0169-5347(93)90024-J
Rosen, B.O. von (1905) Beitrag zur Kenntnis der Molluskenfauna des Kaukasus. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 37 (2), 49–64.
Rosen, B.O. von (1911) Die Mollusken Ciskaukasiens und speciell des Kuban-Gebietes. Annuaire du Musée Zoologique de l'Académie Impériale des Sciences de St.-Pétersbourg, 16, 86–142, pls. 2 + 3.
Skvortsova, V. & Balashov, I. (2019) Land snail Acicula parcelineata (Architaenioglossa: Cyclophoroidea: Aciculidae) in Ukraine: distribution, variability, habitat preferences and conservation status. Ruthenica, 29 (2), 94–102. https://doi.org/10.35885/ruthenica.2019.29(2).3
Steklov, A.A. (1966) Nazemnyye mollyuski neogena Predkavkaz'ya i ikh stratigraficheskoye znacheniye [Terrestrial molluscs of the Neogene Ciscaucasia and their stratigraphic significance]. Academy of Sciences of the USSR Transactions 163. Akademia Nauka S.S.S.R., Moskva, 279.
Steusloff, U. (1939) Beachtenswerte Funde am Niederrhein und im Sauerlande. Archiv für Molluskenkunde, 71, 201–209.
Sysoev, A.V. & Schileyko, A.A. (2009) Land snails and slugs of Russia and adjacent countries. Pensoft Series Faunistica No 87. Pensoft, Sofia/Moskva, 312 pp., 142 pls. [June]
Zelditch, M.L., Swiderski, D.L., Sheets, H.D. & Fink, W.L. (2004) Geometric morphometrics for biologists: a primer. Academic press, New York and London, 456 pp.
Zilch, A. (1976) Die Typen und Typoide des Natur-Museums Senckenberg, 56: Mollusca: Aciculidae. Archiv für Molluskenkunde, 107 (1/3), 123–136.