Our understanding of species distribution and ecology is critical to properly assess their conservation status. Vertigo lilljeborgi, V. genesii, and V. geyeri have the centre of their current distribution in northern Europe, where their occurrence is relatively frequent. However, to the south their occurrence is fragmented and restricted to sites of late glacial/early Holocene origin. In the last ~30 years, there has been an increase in records, connected with the listing of the latter two species in Annex II of the EU Habitats Directive (94/43/EEC). However, there is no comprehensive publication documenting their pan-European distribution. Therefore, we assembled all available data from online databases, books, and scientific literature and combined them with our unpublished records to create distribution maps. The results show a more frequent occurrence in temperate Europe than previously known, especially for V. geyeri. Analyses performed on data from 327 ecologically potentially suitable sites, covering the entire distribution range of the species, have improved our knowledge of their ecology. Vertigo lilljeborgi and especially V. genesii are restricted to areas with lower summer and winter temperatures, and therefore, their further decline is expected in the face of rising temperatures due to climate change. The preference of V. geyeri for higher temperatures, in comparison to the latter two species, may explain its relatively frequent distribution in temperate Europe. Vertigo lilljeborgi favors base-poor sites, while V. genesii and V. geyeri prefer calciumrich sites, with the latter being the most calcicolous. Their need for a stable water regime and lowproductive sites, known from previous studies, was not conspicuous in our results, probably due to the selection of sites well within the species range. Despite the increase in record frequency, these species are still endangered, especially in temperate Europe. Their sites should therefore be strictly protected as sites of high biological diversity and conservation value. Because of their relict nature, these land snails should be considered umbrella species and indicators of well-preserved groundwater-dependent ecosystems in temperate Europe.

Conservatoire Botanique National de Franche Comté Observatoire Régional des Invertébrés Besançon France E mail

Department of Botany and Zoology Faculty of Science Masaryk University Brno Czechia E mail radovan

Department of Zoology Institute of Biosciences Life Sciences Center Vilnius University Saulėtekio av 7 LT 10223 Vilnius Lithuania E mail

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Balashov I. 2016. Conservation of terrestrial molluscs in Ukraine. Institute of Zoology NAS, Kyiv, UA.

Bech M. 1992. Noves aportacions a la malacofauna de Catalunya. Natura B-N 2:164–167.

Breiman L, Friedman JH, Olshen RA, Stone CJ. 1984. Classification and Regression Trees. Chapman and Hall/CRC, New York, USA.

Cadevall J, Corbella J, Bros V, Orozco A, Guillén G, Prats L, Capdevila M. 2020. Els molluscs continentals de Catalunya i Andorra (península Ibèrica): llista comentada. Spira 7:117–159.

Cadevall J, Orozco A. 2016. Caracoles y babosas de la Península Ibérica y Baleares. Ediciones Omega, Barcelona, ES.

Cameron RAD, Colville B, Falkner G, Holyoak GA, Hornung E, Killeen IJ, Moorkens E, Pokryszko B, von Proschwitz T, Tattersfield P, Valovirta I. 2003. Species accounts for snails of the genus Vertigo listed in Annex II of the Habitats Directive: V. angustior, V. genesii, V. geyeri and V. moulinsiana (Gastropoda, Pulmonata: Vertiginidae). Heldia 5 (Sonderheft 7), pp. 151–170.

Čejka T, Beran L, Korábek O, Hlaváč JČ, Horáčková J, Coufal R, Drvotová M, Maňas M, Horsáková V, Horsák M. 2020. Malacological news from the Czech and Slovak Republics in 2015–2019. Malacol Bohemoslov 19: 71–106. doi:10.5817/MaB2020-19-71. DOI

Chytrý M, Tichý L, Hennekens SM, Knollová I, Janssen JAM et al. 2020. EUNIS Habitat Classification: Expert system, characteristic species combinations and distribution maps of European habitats. Appl Veg Sci 23:648–675. doi:10.1111/avsc.12519. DOI

Claude F, Gonseth Y. 2021. Swiss National Mollusca Databank. Swiss National Biodiversity Data and Information Centres –infospecies.ch. Occurrence dataset Accessed via GBIF.org on 13 Dec. 2022. doi:10.15468/zfspwn. DOI

Combrisson D, Vuinée L. 2017. Note sur la présence du vertigo des aulnes Vertigo lilljeborgi (Westerlund, 1871) sur le massif du Taillefer en Isère (France). MalaCo 13:5–7.

Conchological Society of Great Britain & Ireland. 2020. Conchological Society of Great Britain & Ireland: non-marine molluscs (rare and scarce species). Occurrence dataset Accessed via GBIF.org on 13 Dec. 2022. doi:10.15468/e9fnjh. DOI

Coufal R, Hájková P, Hájek M, Jiroušek M, Polášek M, Horsáková V, Horsák M. 2023. Compositional variation of endangered spring fen biota reflects within-site variation in soil temperature. Plant Soil, pp. 1–17. doi:10.1007/s11104-022-05841-3. DOI

Coufal R. 2019. Ekologické determinanty slatiništních společenstev měkkýšů a glaciálně reliktních plžů na Českomoravské vrchovině. –Diploma thesis, Palacký University, Faculty of Science, Department of Ecology and Environmental Sciences.

Duda M. 2015. Interessante Funde zweier Arten der Gattung Vertigo O. F. MÜLLER 1773 (Mollusca, Gastropoda) im süd-westlichen Niederösterreich. Nachrichtenblatt der Ersten Vorarlberger Malakologische Gesellschaft 22:3–4.

Ellenberg H, Leuschner C. 2010. Vegetation Mitteleuropas mit den Alpen in ökologischer, dynamischer und historischer Sicht. Ed. 6. Ulmer, Stuttgart, DE.

Essl F, Dullinger S, Moser D, Rabitsch W, Kleinbauer I. 2012. Vulnerability of mires under climate change: implications for nature conservation and climate change adaptation. Biodivers Conserv 21:655–669. doi:10.1007/s10531-011-0206-x. DOI

Gabriel M. 2020. Zum Vorkommen der Moorwindelschnecke Vertigo lilljeborgi (WESTERLUND 1871) in Bayern und BadenWürttemberg, mit einem neuen Fundort in der Oberpfalz, Bayern. Dtsch Malakozool Ges 103:49–60.

Gong J, Wang K, Kellomäki S, Zhang C, Martikainen PJ, Shurpali N. 2012. Modeling water table changes in boreal peatlands of Finland under changing climate conditions. Ecol Model 244:65–78. doi:10.1016/j.ecolmodel.2012.06.031. DOI

Gural-Sverlova NV, Gural RI. 2012. Vyznachnyk nazemnykh moliuskiv Ukrainy [Guide to the Land Snails of the Ukraine]. National Academy of Sciences of Ukraine, Lviv, UA.

Hájek M, Dítě D, Horsáková V, Mikulášková E, Peterka T, Navrátilová J, Jiménez-Alfaro B, Hájková P, Tichý L, Horsák M. 2020. Towards the pan-European bioindication system: assessing and testing updated hydrological indicator values for vascular plants and bryophytes in mires. Ecol Indic 116:106527. doi:10.1016/j.ecolind.2020.106527. DOI

Hájek M, Hájková P, Goia I, Dítě D, Plášek V. 2021. Variability and classification of Carpathian calcium-rich fens: breaking the state borders. Preslia 93:203–235. doi:10.23855/preslia.2021.203. DOI

Hájek M, Horsák M, Tichý L, Hájková P, Dítě D, Jamrichová E. 2011. Testing a relict distributional pattern of fen plant and terrestrial snail species at the Holocene scale: a null model approach. J Biogeogr 38:742–755. doi:10.1111/j.1365-2699.2010.02424.x. DOI

Hájek M, Těšitel J, Tahvanainen T, Peterka T, Jiménez-Alfaro B, Jansen F, Pérez-Haase A, Garbolino E, Carbognani M, Kolari THM, Hájková P, Jandt U, Aunina L, Pawlikowski P, Ivchenko T, Tomaselli M, Tichý L, Dítě D, Plesková Z, Mikulášková E. 2022. Rising temperature modulates pH niches of fen species. Glob Chang Biol 28:1023–1037. doi:10.1111/gcb.15980. PubMed DOI

Hájková P, Petr L, Horsák M, Jamrichová E, Roleček J. 2022. Holocene history of the landscape at the biogeographical and cultural crossroads between Central and Eastern Europe (Western Podillia, Ukraine). Quat Sci Rev 288: 107610. doi:10.1016/j.quascirev.2022.107610. DOI

Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A. 2005. Very high resolution interpolated climate surfaces for global land areas. Int J Climatol 25:1965–1978. doi:10.1002/joc.1276. DOI

Horsák M, Hájek M, Horsáková V, Hlaváč J, Hájková P, Dítě D, Peterka T, Divíšek J, Potůčková A, Preece RC. 2017. Refugial occurrence and ecology of the land snail Vertigo lilljeborgi in fen habitats in temperate mainland Europe. J Molluscan Stud 83:451–460. doi:10.1093/mollus/eyx028. DOI

Horsák M, Hájek M, Spitale D, Hájková P, Dítě D, Nekola JC. 2012. The age of island‐like habitats impacts habitat specialist species richness. Ecology 93: 1106–1114. doi:10.1890/0012-9658- 93.5.1106. PubMed DOI

Horsák M, Hájek M. 2003. Composition and species richness of molluscan communities in relation to vegetation and water chemistry in the western Carpathian spring fens: the poor rich gradient. J Molluscan Stud 69: 349–357. doi:10.1093/mollus/69.4.349. DOI

Horsák M, Hájek M. 2005. Habitat requirements and distribution of Vertigo geyeri (Gastropoda: Pulmonata) in Western Carpathian rich fens. J Conchol 38:683–700.

Horsák M. 2003. How to sample mollusc communities in mires easily. Malacol Bohemoslov 2:11–14. doi:10.5817/MaB2003-2-11. DOI

Horsák M. 2006. Mollusc community patterns and species response curves along a mineral richness gradient: a case study in fens. J Biogeogr 33:98–107. doi:10.1111/j.1365-2699.2005.01359.x. DOI

Horsáková V, Horsák M. 2018. Vrkoč Geyerův a vrkoč útlý –první nálezy celoevropsky ohrožených plžů ve Slavkovském lese [Geyer’s Whorl Snail and Narrow-Mouthed Whorl Snail –the first records of pan-European endangered snails in the Slavkovský les Forest]. Arnika 1:30–33.

Horsák M, Ortiz D, Nekola JC, Van Bocxlaer B. 2024. Intercontinental dispersal and niche fidelity drive 50 million years of global diversification in Vertigo land snails. Glob Ecol Biogeogr 33:e13820. doi:10.1111/geb.13820. DOI

Hughes PD, Gibbard PL, Ehlers J. 2013. Timing of glaciation during the last glacial cycle: evaluating the concept of a global ‘Last Glacial Maximum’ (LGM). Earth-Sci Rev 125: 171–198. doi: 10.1016/j.earscirev.2013.07.003. DOI

Jaeckel SGA. 1962. Ergänzungen und Berichtigungen zum rezenten und quartären Vorkommen der mitteleuropäischen Mollusken. In: Brohmer P, Ehrmann P, Ulmer G, (eds.), Die Tierwelt Mitteleuropas 2 (1 Ergänzung). Quelle and Meyer, Leipzig, DE.

Jamrichová E, Hájková P, Horsák M, Rybníčková E, Lacina A, Hájek M. 2014. Landscape history, calcareous fen development and historical events in the Slovak Eastern Carpathians. Veget Hist Archaeobot 23:497–513. doi:10.1007/s00334-013-0416-0. DOI

Jensen K, Schrautzer J. 1999. Consequences of abandonment for a regional fen flora and mechanisms of successional change. Appl Veg Sci 2:79–88. doi:10.2307/1478884. DOI

Jiménez‐Alfaro B, Hájek M, Ejrnaes R, Rodwell J, Pawlikowski P, Weeda EJ, Laitinen J, Moen A, Bergamini A, Aunina L, Sekulová L, Tehvanainen T, Gillet F, Jandt U, Dítě D, Hájková P, Corriol G, Kondelin H, Díaz TE. 2014. Biogeographic patterns of base‐rich fen vegetation across Europe. Appl Veg Sci 17:367–380. doi:10.1111/avsc.12065. DOI

Joosten H, Tanneberger F, Moen A. (Eds.) 2017. Mires and Peatlands of Europe. Schweizerbart Science Publishers, Stuttgart, DE.

Kerney MP, Cameron RAD, Jungbluth JH. 1983. Die Landschnecken Nord-und Mitteleuropas: Ein Bestimmungsbuch für Biologen und Naturfreunde. Parey, Hamburg, DE.

Killeen IJ, Willing M, Moorkens E. 2019. Site Condition Monitoring of Vertigo geyeri and Vertigo genesii 2017. Scottish Natural Heritage Research Report No. 1161.

Killeen IJ. 2003. A review of EUHSD Vertigo species in England and Scotland (Gastropoda, Pulmonata: Vertiginidae). In: Speight MCD, Moorkens E, Falkner G (eds) Proceedings of the workshop on conservation biology of European Vertigo species (Dublin, Ireland, April 2002). Heldia 5 (Sonderheft 7), Munich, Germany, pp. 73–84.

Krausp C. 1940. Beitrag zur Molluskenfauna Lettlands. Tartu Ülikooli juures oleva Loodusuurijate Seltsi aruanded, 45:217–270, Tartu.

Książkiewicz Z, Biereznoj-Bazille U, Krajewski L, Goldyn B. 2015. New records of Vertigo geyeri Lindholm, 1925, V. moulinsiana (Dupuy, 1849) and V. angustior Jeffreys, 1830 (Gastropoda: Pulmonata: Vertiginidae) in Poland. Folia Malacol 23:121–136. doi:10.12657/folmal.023.006. DOI

Kuczyńska A, Moorkens E. 2010. Micro-hydrological and micrometeorological controls on survival and population growth of the whorl snail Vertigo geyeri Lindholm, 1925 in groundwater fed wetlands. Biol Conserv 143: 1868–1875. doi:10.1016/j.biocon.2010.04.033. DOI

Kurylyk BL, Bourque CA, MacQuarrie KT. 2013. Potential surface temperature and shallow groundwater temperature response to climate change: an example from a small forested catchment in east-central New Brunswick (Canada). Hydrol Earth Syst Sci 17:2701–2716. doi:10.5194/hess-17-2701-2013. DOI

Kuznecova V, Skujienė G. 2011. Biodiversity of terrestrial molluscs in urbophytocoenoses of Vilnius, capital of Lithuania. IOBC/wprs Bulletin 64:41–51.

Kuznecova V, Skujienė G. 2012. Rare terrestrial molluscs’ species of Kaunas and Kaišiadoriai districts’ reserves. Acta Biol Univ Daugavpiliensis 12:69–83.

Lasne O, Ryelandt J, Horsák M, Horsáková V. 2021. Étude de l’habitat et des exigences écologiques du Vertigo geyeri Lindholm, 1925 dans le Massif jurassien. In: Léonard L (ed): Colloque national de malacologie continentale. Nantes 6: 21–33. doi:10.5852/naturae2021a2. DOI

Lasne O. 2018. Étude de l’habitat et des exigences écologiques du Vertigo geyeri Lindholm, 1925 dans le massif jurassien. Master’s thesis, Conservatoire botanique national de Franche-Comté –Observatoire régional des Invertébrés, Besançon.

Lecaplain B. 2013. Un nouveau mollusque de la Directive HabitatsFaune-Flore pour la France: découverte du Vertigo septentrional Vertigo geyeri Lindholm, 1925 (Gastropoda, Vertiginidae) en Franche-Comté et en Haute-Savoie. MalaCo 9:453–456.

Ložek V. 1964. Quartärmollusken der Tschechoslowakei. ČSAV, Prague, CZ.

Ložek V. 1992. [Molluscs (Mollusca)]. [Red data book of endangered and rare plant and animal species of the ČSFR. 3. Invertebrates]. Príroda, Bratislava, SK.

Ložek V. 1993. Vertigo geyeri in Böhmen. Dtsch Malakozool Ges 50/51:53–54.

Menberg K, Blum P, Kurylyk BL, Bayer P. 2014. Observed groundwater temperature response to recent climate change. Hydrol Earth Syst Sci 18: 4453–4466. doi:10.5194/hess-18-4453 2014. DOI

Myšák J, Horsák M, Hlaváč J. 2012. Jedna špatná a jedna dobrá zpráva o vrkoči Geyerově –z červené knihy našich měkkýšů. Živa 60:73–74.

Natural Resources Wales. 2020. Welsh Invertebrate Database (WID). Occurrence dataset. Accessed via GBIF.org on 13 Dec. 2022. doi:10.15468/bv8fcj. DOI

Nekola JC, Chiba S, Coles BF, Drost CA, von Proschwitz T, Horsák M. 2018. A phylogenetic overview of the genus Vertigo O. F. Müller, 1773 (Gastropoda: Pulmonata: Pupillidae: Vertigininae). Malacologia 62:21–161. doi:10.4002/040.062.0104. DOI

Nekola JC, Coles BF. 2010. Pupillid land snails of eastern North America. Am Malacol Bull 28:29–57. doi:10.4003/006.028.0221. DOI

Nekola JC, Coles BF. 2016. Supraspecific taxonomy in the Vertiginidae (Gastropoda: Stylommatophora). J Molluscan Stud 82:208–212. doi:10.1093/mollus/eyv034. DOI

Ogle DH, Doll JC, Wheeler AP, Dinno A. 2023. FSA: Simple Fisheries Stock Assessment Methods. R package version 0.9.4. Available at: https://CRAN.R-project.org/package=FSA.

Peterka T, Tichý L, Horsáková V, Hájková P, Coufal R, Petr L, Dítě D, Hradílek Z, Hrivnák R, Jiroušek M, Plášek V, Plesková Z, Singh P, Šmerdová E, Štechová T, Mikulášková E, Horsák M, Hájek M. 2022. The long history of rich fens supports persistence of plant and snail habitat specialists. Biodiv Conserv 31:39–57. doi:10.1007/s10531-021-02318-0. DOI

Pilāte D. 2000. Spermodea lamellata (Jeffreys, 1830) un Vertigo genesii (Gredler, 1856) (Gastropoda: Pulmonata) Latvijā. [Spermodea lamellata (Jeffreys, 1830) and Vertigo genesii (Gredler, 1856) (Gastropoda: Pulmonata) in Latvia]. Raksti par dabu 1:3–5.

Pilāte D, Spuņģis V, Skuja A, Ozoliņš D. 2022. Investigation of occurrence of specially protected whorl snails Vertigo spp. in the habitats of EU importance. 80th International Scientific Conference of the University of Latvia. Innovative and applied research in Biology, Proceedings 4: 23–25 doi:10.22364/iarb.2022.03. DOI

Pokryszko BM. 2003. Vertigo of continental Europe—autecology, threats and conservation status. In: Speight MCD, Moorkens E, Falkner G (eds) Proceedings of the workshop on conservation biology of European Vertigo species (Dublin, Ireland, April 2002). Heldia 5 (Sonderheft 7), Munich, Germany, pp. 13–25.

Pokryszko BM, Ruta R, Książkiewicz-Parulska Z. 2016. The first record of Vertigo geyeri Lindholm, 1925 (Gastropoda: Pulmonata: Vertiginidae) in north-western Poland. Folia Malacol 24:63–68. doi:10.12657/folmal.024.009. DOI

Pokryszko BM. 1990. The Vertinigidae of Poland (Gastropoda: Pulmonata: Pupilloidea) -a systematic monograph. Ann Zool 8(43).

R Core Team. 2022. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available at: https://www.R-project.org/.

Roy C, Vanderpert H. 2016. Découverte de Vertigo geyeri Lindholm, 1925 en Provence-Alpes-Côte d’Azur. Folia Conchyliologica 35:1–5.

Rudzīte M, Dreijers E, Ozoliņa-Moll L, Parele E, Pilāte D, Rudzītis M, Stalažs A. 2010. Latvijas gliemji Sugu noteicejs [A Guide to the Molluscs of Latvia]. Akadēmiskais apgāds, Rīga, LV.

Rydin H, Sjörs H, Löfroth M. 1999. Mires. Acta Phytogeogr Suec 84:91–112.

Šatkauskienė I. 2001. Naujos šliužų rūšies Lietuvoje Boettgerilla pallens ir retųjų Lietuvos sausumos moliuskų rūšių apžvalga. [A review of rare Lithuanian terrestrial malacofauna species and the Boettgerilla pallens species new to Lithuania]. Ekologija 2:56–60.

Schenková V, Horsák M, Plesková Z, Pawlikowski P. 2012. Habitat preferences and conservation of Vertigo geyeri (Gastropoda: Pulmonata) in Slovakia and Poland. J Molluscan Stud 78:105–111. doi:10.1093/mollus/eyr046. DOI

Schenková V, Horsák M. 2013a. Refugial Populations of Vertigo lilljeborgi and V. genesii (Vertiginidae): New Isolated Occurrences in Central Europe, Ecology and Distribution. Am Malacol Bull 31:323–329. doi:10.4003/006.031.0211. DOI

Schenková V, Horsák M. 2013b. Nové nálezy vrkoče Geyerova potvrzují jeho ohroženost –z červené knihy našich měkkýšů. Živa 5:238–239.

Skujienė G, Kuznecova V, Jukonis B, Juzėnas S. 2020. New records of rare Vertiginidae (Mollusca: Gastropoda) in Lithuania. Bull Lithuanian Entomol Soc 4:132–140.

Skujienė G, Kuznecova V, Juzėnas S. 2019. New records of Vertigo geyeri (Gastropoda: Vertiginidae) in Lithuania. Bull Lithuanian Entomol Soc 3:132–139.

Skujienė G, Kuznecova V, Juzėnas S. 2021. Additional data on rare Vertiginidae (Mollusca: Gastropoda) in Lithuania. Bull Lithuanian Entomol Soc 5:118–125.

Stalažs A, Dreijers E. 2016. Annotated checklist of the molluscs of the Baltic countries. Raksti par Dabu 2:9–20.

Therneau T, Atkinson B, Ripley B. 2022. rpart: Recursive partitioning for classification, regression and survival trees. An implementation of most of the functionality of the 1984 book by Breiman, Friedman, Olshen and Stone. R package version 4. 1. 19.

Turner H, Kuiper JGJ, Thew N, Bernasconi R, Ruetschi J, Wuthrich M, Gosteli M. 1998. Atlas of the Mollusca of Switzerland and Liechtenstein. Centre Suisse de cartographie de la faune 1998 (CSCF/SZKF), Neuchâtel, Switzerland.

Valovirta I. 2003. The habitat and status of Vertigo angustior, V. genesii and V. geyeri in Finland and nearby Russian Karelia (Gastropoda, Pulmonata: Vertiginidae). In: Speight MCD, Moorkens E, Falkner G (eds) Proceedings of the workshop on conservation biology of European Vertigo species (Dublin, Ireland, April 2002). Heldia 5 (Sonderheft 7), Munich, Germany, pp. 85–94.

van Regteren Altena CO. 1934. Note sur une récolte de mollusques aux environs de Font-Romeu. Journal de Conchyliologie 78:262–269.

Vavrová LU, Horsák M, Šteffek J, Čejka T. 2009. Ecology, distribution and conservation of Vertigo species or European importance in Slovakia. J Conchol 40:61.

von Proschwitz T. 1993. Habitat selection and distribution of ten vertiginid species in the province of Dalsland (SW. Sweden) (Gastropoda, Pulmonata: Vertiginidae). Malakologische Abhandlungen Staatlisches Museum für Tierkunde Dresden 16:177–212.

von Proschwitz T. 2003. A review of the distribution, habitat selection and conservation status of the species of the genus Vertigo in Scandinavia (Denmark, Norway and Sweden) (Gastropoda, Pulmonata: Vertiginidae). In: Speight MCD, Moorkens E, Falkner G (eds) Proceedings of the workshop on conservation biology of European Vertigo species (Dublin, Ireland, April 2002). Heldia 5 (Sonderheft 7), Munich, Germany, pp. 27–50.

von Proschwitz T. 2004. On the distribution and ecology of Vertigo substriata (Jeffreys), Vertigo modestaarctica (Wallenberg), Vertigo lilljeborgi (Westerlund) and Vertigo alpestris Alder in France and on the Iberian Peninsula. J Conchol 38:411–420.

von Proschwitz T, Roth J, Lundin K, Back R. 2023. Nationalnyckeln till Sveriges flora och fauna. Blötdjur: Snyltsnäckor-skivsnäckor. Mollusca: Pyramidellidae-Planorbidae. SLU Artdatabanken, Uppsala, Sweden.

Vrignaud S. 2012. Inventaire des mollusques de la réserve naturelle nationale de Chastreix-Sancy (Puy-de-Dôme, France). Rev Sci Nat Auvergne 76:39–52.

Waldén HW. 1966. Einige Bemerkungen zum Erganzungsband zu Ehrmann’s “Mollusca”, in “Die Tierwelt Mitteleuropas”. Arch Moll 95(1/2):49–68.

Wickham H. 2016. ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. ISBN 978-3-319-24277-4. Available at: https://ggplot2.tidyverse.org.

Wood S. 2022. mgcv: Mixed GAM Computation Vehicle with Automatic Smoothness Estimation. R package version 1.8-41. Available at: https://CRAN.R-project.org/package=mgcv.

Wright HE Jr, Ammann B, Stefanova I, Atanassova J, Margalitadze N, Wick L, Blyakharchuk T. 2003. Late-glacial and Early-Holocene Dry Climates from the Balkan Peninsula to Southern Siberia. In: Tonkov S (ed) Aspects of palynology and palaeoecology. Pensoft Publishing, Sofia, Moscow, BG.

Pyrenees as the southernmost European refugium of glacial relict land snails