Genital morphology differentiates three subspecies of the terrestrial slug Arion ater (Linnæus, 1758) s.l. and reveals a continuum of intermediates with the invasive A. vulgaris Moquin-Tandon, 1855

Heike Reise, e-mail: heike.reise@senckenberg.de

Senckenberg Museum of Natural History Görlitz, Am Museum 1, 02826 Görlitz, Germany

Anne-Katrin Schwarzer, e-mail: anne.k.schwarzer@gmail.com

Senckenberg Museum of Natural History Görlitz, Am Museum 1, 02826 Görlitz, Germany

John M. C. Hutchinson, e-mail: majmch@gmail.com;john.hutchinson@senckenberg.de

Senckenberg Museum of Natural History Görlitz, Am Museum 1, 02826 Görlitz, Germany

Bettina Schlitt, e-mail: bettina.schlitt@senckenberg.de

Senckenberg Museum of Natural History Görlitz, Am Museum 1, 02826 Görlitz, Germany
Abstract

The terrestrial slug Arion vulgaris Moquin-Tandon (= A. lusitanicus auct. non Mabille) is an important agricultural pest that has invaded much of Europe. Previous work has demonstrated hybridisation with A. ater (Linnæus) s.l. We describe the genital anatomy of morphological intermediates found in eastern Saxony (Germany), comparing them with the parent species. We recommend a standard method of genital dissection and consider a set of five genital characters. The intermediates demonstrate the homology of the ligula-bearing organs of the two species; the ancestral position of the ligula in the dilated part of the oviduct has moved to the upper atrium in A. ater s.l. Furthermore we differentiate three morphotypes of A. ater s.l. in eastern Saxony, associated with different mitochondrial DNA sequences. One is A. ater ater, previously unrecognised from this part of Germany and the Czech Republic. The two other morphotypes correspond to the predominantly British and the Continental subspecies previously recognised on genetic and morphological grounds. We designate as the lectotype of Limax rufus Linnæus, 1758 a non-surviving specimen from Almondbury, England, described by Lister; thus the predominantly British subspecies becomes A. ater rufus. The appropriate name for the Continental form is A. ater ruber (Garsault, 1764).

Key words
Arion ater ruber; Arion lusitanicus; Arion rufus; Spanish slug; Pulmonata; Arionidae; COI; hybrid; lectotype; ligula; Martin Lister; pest slug
References

Adomaitis M., Skujienė G. 2016. Comparative analysis of ligulas of slugs Arion lusitanicus from Lithuania and Denmark. Scientific Bulletin of the Uzhgorod University (Series Biology) 40: 5–10.
Aguiar R., Wink M. 2005. How do slugs cope with toxic alkaloids? Chemoecology 15: 167–177. https://doi.org/10.1007/s00049-005-0309-5
Aldrovandi U. 1644. De animalibvs insectis libri septem cvm singvlorvm iconibvs ad viuum expressis. Cum indice copiosissimo. Ferronium, Bonon.
Allgaier C. 2014. How can two soft bodied animals be precisely connected? A miniature quick-connect system in the slugs, Arion lusitanicus and Arion rufus. Journal of Morphology 276: 631–648. https://doi.org/10.1002/jmor.20361
Anderson R. 2005. An annotated list of the non-marine Mollusca of Britain and Ireland. Journal of Conchology 38: 607–637.
Backeljau T., Van Beeck M. 1986. Epiphallus anatomy in the Arion hortensis species aggregate (Mollusca, Pulmonata). Zoologica Scripta 15: 61–68. https://doi.org/10.1111/j.1463-6409.1986.tb00209.x
Balashov I. 2018. Case 3685 – Arion vulgaris Moquin-Tandon, 1855 (Gastropoda, Stylommatophora, Arionidae): proposed validation of the specific name as available. Bulletin of Zoological Nomenclature 75: 12–15. https://doi.org/10.21805/bzn.v75.a006
Blattmann T., Boch S., Türke M., Knop E. 2013. Gastropod seed dispersal: an invasive slug destroys far more seeds in its gut than native gastropods. PLoS ONE 8(9): e75243. https://doi.org/10.1371/journal.pone.0075243
Boato A., Bodon M., Giusti F. 1983. On a new slug from the Northern Apennines (Pulmonata: Arionidae). Archiv für Molluskenkunde 114: 35–44.
Boettger C. R. 1949a. Über das Vorkommen einer zweiten Art von großen Wegschnecken (Arion sens. strict.) in Deutschland. Verhandlungen der deutschen Zoologen vom 24.–28. August 1948 in Kiel: 499–501.
Boettger C. R. 1949b. Zur Kenntnis der großen Wegschnecken (Arion s. str.) Deutschlands. Archiv für Molluskenkunde 78: 169–186.
Cain A. J., Williamson M. H. 1958. Variation and specific limits in the Arion ater aggregate. Proceedings of the Malacological Society of London 33: 72–86.
Castillejo J. 1997. Babosas del Noroeste Ibérico. Universidade de Santiago de Compostela, Servico de Puplicatións e Intercambio Científico, Santiago de Compostela.
Castillejo J. 1998. Guía de las babosas ibéricas. Real Academia Galega de Ciencias, Santiago.
Castillejo J., Rodríguez T. 1991. Babosas de la península ibérica y Baleares. Inventario crítico, citas y mapas de distributión. (Gastropoda, Pulmonata, Terrestria nuda). Monografías de la Universidad de Santiago de Compostela 162. Servicio de publicaciones de la Universidad de Santiago de Compostela, Santiago de Compostela.
Castillejo J., Rodríguez T. 1993. Las especies del genero Arion Férussac, 1819 en Portugal (Gastropoda, Pulmonata: Arionidae). Graellsia 49: 17–37.
Castillejo J., Rodríguez-Castro J., Iglesias-Piñeiro J. 2019. Estudio comparativo de la anatomía y caracterización del ADN de los ariónidos descritos por Torres Mínguez (1925) en Cantabria (España): Arion cendreroi y A. fulvipes, y la de A. rufus y A. vulgaris (Gastropoda Pulmonata: Arionidae). Spira 7: 49–69.
Chevallier H. 1972. Arionidae (Mollusca, Pulmonata) des Alpes et du Jura francais. Haliotis 2: 7–23.
Chevallier H. 1974. Les grands Arion de France (Mollusca, Pulmonata). Taxonomie, biogeographie, écologie, polymorphisme, croissance et cycle biologique. Ph. D. Thesis, Université de Paris.
Chevallier H. 1981. Taxonomie des “Limaces rouges” (genre Arion, sous-genre Arion s.s., Mollusca, Pulmo­nata). Haliotis 11: 87–99.
Clement M., Posada D., Crandall K. A. 2000. TCS: a computer program to estimate gene genealogies. Molecular Ecology 9: 1657–1660. https://doi.org/10.1046/j.1365-294x.2000.01020.x
Collinge W. E. 1897a. Some observations on certain species of Arion. Journal of Malacology 6: 7–10, Pl. II. https://biodiversitylibrary.org/page/16099983
Collinge W. E. 1897b. On some European slugs of the genus Arion. Proceedings of the Zoological Society of London 3: 439–450. https://doi.org/10.1111/j.1096-3642.1897.tb03102.x
Currat M., Ruedi M., Petit R. J., Exoffier L. 2008. The hidden side of invasions: massive introgression by local genes. Evolution 62: 1908–1920. https://doi.org/10.1111/j.1558-5646.2008.00413.x
Davies S. M. 1987. Arion flagellus Collinge and A. lusitanicus Mabille in the British Isles: a morphological, biological and taxonomic investigation. Journal of Conchology 32: 339–354.
Dreijers E., Reise H., Hutchinson J. M. C. 2013. Mating of the slugs Arion lusitanicus auct. non Mabille and A. rufus (L.): different genitalia and mating behaviours are incomplete barriers to interspecific sperm exchange. Journal of Molluscan Studies 79: 51–63. https://doi.org/10.1093/mollus/eys033
Dvořák L., Horsák M. 2003. Současné poznatky o plzáku Arion lusitanicus (Mollusca: Pulmonata) v České republice. Časopis Slezského Muzea Opava (A) 52: 67–71.
Ehrmann P. 1933. Kreis: Weichtiere, Mollusca. In: Brohmer P., Ehrmann P., Ulmer G. (eds). Die Tierwelt Mitteleuropas. Vol. II (1) Mollusca, Crustacea, Isopoda, Myriapoda. Quelle & Meyer, Leipzig, pp. 1–264.
Engelke S., Kömpf J., Jordaens K., Tomiuk J., Parker E. D. 2011. The genetic dynamics of the rapid and recent colonization of Denmark by Arion lusitanicus (Mollusca, Pulmonata, Arionidae). Genetica 139:709–721. https://doi.org/10.1007/s10709-011-9565-1
Evans N. J. 1986. An investigation of the status of the terrestrial slugs Arion ater ater (L.) and Arion ater rufus (L.) (Mollusca, Gastropoda, Pulmonata) in Britain. Zoologica Scripta 15: 313–322. https://doi.org/10.1111/j.1463-6409.1986.tb00232.x
Falkner G. 1981. Vier bemerkenswerte kulturfolgende Schneckenarten in Hildesheim-Ochtersum. Mitteilungen der zoologischen Gesellschaft Braunau 3: 391–396.
Férussac A. E. J. P. J. F. d’Audebard de, Deshayes G.-P. 1819–1851. Histoire naturelle générale et particulière des mollusques terrestres et fluviatiles, tant des espèces que l’on trouve aujourd’hui vivantes, que des dépouilles fossiles de celles qui n’existent plus; classés d’après les caractères essentiels que présentent ces animaux et leurs coquilles. J.-B. Bailliere, Paris. https://doi.org/10.5962/bhl.title.132272
Fischer W., Reischütz P. L. 1998. Grundsätzliche Bemerkungen zum Schadschneckenproblem. Die Bodenkultur 49: 281–292.
Fleming J. 1822. Mollusca. In: [Napier M. (ed.)] Supplement to the fourth, fifth and sixth editions of the Encyclopædia Britannica with preliminary dissertations on the history of sciences. Vol. 5. Constable, Edinburgh, pp. 567–584.
Folmer O., Black M., Hoeh W., Lutz R., Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294–299.
Frömming E. 1953. Massenvorkommen von roten Arion rufus L. in Berlin. Archiv für Molluskenkunde 82: 77–78.
Garsault F. A. P. de 1764. Les figures des plantes et animaux d’usage en medecine, décrits dans la Matiere Medicale de Mr. Geoffroy medecin, dessinés d’après nature par Mr. de Garsault, gravés par Mrs. Defehrt, Prevost, Duflos, Martinet &c. Niquet scrip. Vol. 5. Paris. https://doi.org/10.5962/bhl.title.49481
Garsault F. A. P. de 1767. Description, vertus et usages de sept cents dix-neuf plantes, tant étrangeres que de nos climats; et de cent trente-quatre animaux, en sept cents trente planches, gravées en taille-douce, sur les desseins d’après nature, de M. de Garsault, par MM. de Fehrt, Prevost, Duflos, Martinet, &c. Et rangées suivant l’ordre du livre intitulé Matiere Médicale de M. Geoffroy. Ouvrage utile à toutes matieres médicales, aux artistes, aux personnes charitables, & à tous ceux qui préparent eùx-memes leurs médicaments. Didot, Paris.
Gerhardt U. 1940. Neue biologische Nacktschnecken­studien. Zeitschrift für Morphologie und Ökologie von Tiere 36: 557–580. https://doi.org/10.1007/BF01260999
Grimm B. 2001. Life cycle and population density of the pest slug Arion lusitanicus Mabille (Mollusca: Pulmonata) on grassland. Malacologia 43: 25–32.
https://biodiversitylibrary.org/page/13142874
Hagnell J., Schander C., von Proschwitz T. 2004. Sniglarna som rör om i trädgårdar och släktträd. Fauna och Flora 99: 38–41.
Hatteland B. A., Roth S., Andersen A., Kaasa K., Støa B., Solhøy T. 2013. Distribution and spread of the invasive slug Arion vulgaris Moquin-Tandon in Norway. Fauna norvegica 32: 13–26. https://doi.org/10.5324/fn.v32i0.1473
Hatteland B. A., Solhøy T., Schander C., Skage M., von Proschwitz T., Noble L. R. 2015. Introgression and differentiation of the invasive slug Arion vulgaris from native A. ater. Malacologia 58: 303–321. https://doi.org/10.4002/040.058.0210
Hesse P. 1926. Die Nacktschnecken der palaearktischen Region. Abhandlungen des Archivs für Molluskenkunde 2(3): 1–152, Pl. I–II.
Horsák M., Juřičková L., Picka J. 2013. Molluscs of the Czech and Slovak Republics. Nakladatelství Kabourek, Zlín.
Hutchinson J. M. C., Reise H. 2015. Mating in Ariunculus isselii, an arionid slug without a spermatophore. Journal of Molluscan Studies 81: 247–258. https://doi.org/10.1093/mollus/eyu086
Hutchinson J. M. C., Schlitt B., Kořínková T., Reise H., Barker G. M. in submission. Genetic evidence illuminates the origin and global spread of the slug Deroceras invadens.
ICZN 1999. International code of zoological nomenclature. Fourth edition. International Trust for Zoological Nomenclature, London.
Jaeckel S. G. H. 1962. Ergänzungen und Berichtigungen zum rezenten und quartären Vorkommen der mitteleuropäischen Mollusken. In: Brohmer P., Ehrmann P., Ulmer G. (eds). Die Tierwelt Mitteleuropas Vol. II (1), Ergänzung. Quelle & Meyer, Leipzig, pp. 25–294.
Jensen K., Engelke S., Simpson S. J., Mayntz D., Hunt J. 2013. Balancing of specific nutrients and subsequent growth and body composition in the slug Arion lusita­nicus. Physiology & Behavior 122: 84–92. https://doi.org/10.1016/j.physbeh.2013.08.023
Kadolsky D., Welter-Schultes F., Bank R. A. 2018. Comment (Case 3685) – Arion vulgaris Moquin-Tandon, 1855 (Gastropoda, Stylommatophora, Arionidae): modified proposal to preserve the specific name in its accustomed sense. Bulletin of Zoological Nomenclature 75: 251–254. https://doi.org/10.21805/bzn.v75.a051
Kappes H., Kobialka H. 2009. Die Nacktschnecken­gesellschaften in NW-Deutschland (Gastropoda: Milacidae, Boettgerillidae, Limacidae, Agriolimacidae, Arionidae): ein Ergebnis der NRW-Kartierung. Natur und Heimat 69: 73–94.
Kappes H., Stoll S., Haase P. 2012. Differences in field behavior between native gastropods and the fast-spreading invader Arion lusitanicus auct. non Mabille. Belgian Journal of Zoology 142: 49–58.
Kerney M. 1999. Atlas of the land and freshwater molluscs of Britain and Ireland. Harley Books, Great Horkesley.
Kerney M. P., Cameron R. A. D., Jungbluth J. H. 1983. Die Landschnecken Nord- und Mitteleuropas. Paul Parey, Hamburg.
Knop E., Reusser N. 2012. Jack-of-all-trades: phenotypic plasticity facilitates the invasion of an alien slug species. Proceedings of the Royal Society B 279: 4668–4676. https://doi.org/10.1098/rspb.2012.1564
Kozłowski J. 2007. The distribution, biology, population dynamics and harmfulness of Arion lusitanicus Mabille, 1868 (Gastropoda: Pulmonata: Arionidae) in Poland. Journal of Plant Protection Research 47: 219–230. http://www.plantprotection.pl/pdf-90950-25302?filename=The%20distribution_.pdf
Kozłowski J., Kozłowski R. J. 2011. Expansion of the invasive slug species Arion lusitanicus Mabille, 1868 (Gastropoda: Pulmonata: Stylommatophora) and dangers to garden crops – a literature review with some new data. Folia Malacologica 19: 249–258. https://doi.org/10.2478/v10125-011-0005-8
Likharev I. M., Wiktor A. 1980. Slizni fauny SSSR i sopredel’nykh stran (Gastropoda terrestria nuda). Fauna SSSR: Mollyuski (Novaja serija 122) Tom 3(5). Fauna of the USSR: Mollusca (New series 122) Vol. III, Part 5]. Nauka, Leningrad.
Linnæus [Linnaeus] C. 1746. Fauna Svecica sistens animalia Sveciæ Regni: qvadrupedia, aves, amphibia, pisces, insecta, vermes, distributa per classes & ordines, genera & species. Cum differentiis specierum, synonymis autorum, nominibus incolarum, locis habitationum, descriptionibus insectorum. Wishoff, Lugduni Batavorum. https://doi.org/10.5962/bhl.title.34892
Linnæus [Linnaeus] C. 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Salvius, Holmiæ. https://doi.org/10.5962/bhl.title.542
Linné [Linnaeus] C. a 1767. Systema naturæ, Tom. I. Pars II. Editio duodecima reformata. Salvius, Holmiæ.
Lister M. 1678. Historiæ animalium Angliæ tres tractatus. Unus de araneis. Alter de cochleis tum terrestribus tum fluviatilibus. Tertius de cochleis marinis. Quibus adjectus est quartus de lapidis ejusdem insulæ ad cochlearum quandam imaginem figuratis. Memoriæ & Rationi. Martyn, Londini. https://doi.org/10.5962/bhl.title.65746
Lister M. 1685. Appendicis ad historiæ animalium Angliæ, tres tractatus, &c. Altera editiio, auctior & emendatior. Smith, Londini.
MacGregor A. 2001. The Ashmolean as a museum of natural history, 1683–1860. Journal of the History of Collections 13: 125–144. https://doi.org/10.1093/jhc/13.2.125
Moquin-Tandon A. 1855. Histoire naturelle des mollusques terrestres et fluviatiles de France. Vol. II. and Atlas J.-B. Baillière, Paris. https://doi.org/10.5962/bhl.title.13098
Muder P., Tietz O., Jechorek H. 2002. Geologische und bergbaukundliche Exkursion in den Tagebau Berzdorf. In: Tietz O. (ed.). Sammlung geologischer Exkursionsführer des Staatlichen Museums für Naturkunde Görlitz 1995–2005, pp. 72–83. Unpublished, available at library of the Senckenberg Museum of Natural History Görlitz.
Müller O. F. 1774. Vermivm terrestrium et fluviatilium, seu animalium infusoriorum, helminthicorum, et testaceorum, non marinorum, succincta historia. Volumen alterum. Heineck & Faber, Havniæ & Lipsiæ. https://doi.org/10.5962/bhl.title.46299
Noble L. R. 1992. Differentiation of large arionid slugs (Mollusca, Pulmonata) using ligula morphology. Zoologica Scripta 213: 255–263. https://doi.org/10.1111/j.1463-6409.1992.tb00330.x
Noble L. R., Jones C. S. 1996. A molecular and ecological investigation of the large arionid slugs of North-West Europe: the potential for new pests. In: Symondson, W. O. C., Liddel, J. E. (eds). The ecology of agricultural pests: biochemical approaches. The Systematics Association special volume series 53. Chapman & Hall, London, pp. 93–131.
Odhner N. H. 1951. Swedish high mountain Mollusca. In: Brink P., Wingstrand K. G. (eds). The mountain fauna of the Virihaure area in Swedish Lapland. C.W.K. Gleerup, Lund, pp. 26–50.
Păpureanu A.-M., Reise H., Varga A. 2014. First records of the invasive slug Arion lusitanicus auct. non Mabille (Gastropoda: Pulmonata: Arionidae) in Romania. Malacologica Bohemoslovaca 13: 6–11. http://mollusca.sav.sk/pdf/13/13.Papureanu.pdf
Paradis E., Jombart T., Brian K., Schliep K., Potts A., Winter D., Kamvar Z. N. 2018. Population and evolutionary genetics analysis system v. 0.11. http://ape-package.ird.fr/pegas.html accessed 17.viii.2019.
Parmakelis A., Kotsakiozi P., Rand D. 2013. Animal mitochondria, positive selection and cyto-nuclear coevolution: insights from pulmonates. PLoS ONE 8: e61970. https://doi.org/10.1371/journal.pone.0061970
Peláez M., Valdecasas A. G., Martinez D., Horreo J. L. 2018. Towards the unravelling of the slug A. ater–A. rufus complex (Gastropoda Arionidae): new genetic approaches. Web Ecology 18: 115–119. https://doi.org/10.5194/we-18-115-2018
Pfenninger M., Weigand A., Bálint M., Klussmann-Kolb A. 2014. Misperceived invasion: the Lusitanian slug (Arion lusitanicus auct. non-Mabille or Arion vulgaris Moquin-Tandon 1855) is native to Central Europe. Evolutionary Applications 7: 702–713. https://doi.org/10.1111/eva.12177
Pilsbry H. A. 1948. Land Mollusca of North America (north of Mexico). Volume II, Part 2. Academy of Natural Sciences of Philadelphia, Monographs 3.
Pollonera C. 1890: Recensement des Arionidae de la région Paléarctique. Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Università di Torino 5(87): 1–42.
Proschwitz T. von 1985. Röda skogssnigeln Arion rufus (L.), en mellaneuropeisk art stadd i snabb spridning med människan i Sverige. Fauna och Flora 80: 121–138.
Proschwitz T. von 1996. Utbredning och spridning av spansk skogssnigel [Arion lusitanicus Mabille] och röd skogssnigel [Arion rufus (L.)] – En översikt av utvecklingen i Sverige. Göteborgs Naturhistoriska Museum, Årstryck, 1996: 27–45.
Proschwitz T. von 1997. Arion lusitanicus Mabille and A. rufus (L.) in Sweden: A comparison of occurrence, spread and naturalization of two alien slug species. Heldia 4: 137–138.
Quick H. E. 1947. Arion ater (L.) and A. rufus (L.) in Britain and their specific differences. Journal of Conchology 22: 249–261.
Quick H. E. 1952. Rediscovery of Arion lusitanicus Mabille in Britain. Proceedings of the Malacological Society of London 29: 93–101.
Quick H. E. 1960. British slugs (Pulmonata; Testacellidae. Arionidae, Limacidae). Bulletin of the British Museum (Natural History), Zoology 6: 105–226. https://doi.org/10.5962/bhl.part.26845
Quinteiro J., Rodríguez-Castro J., Castillejo J., Iglesias-Piñeiro J., Rey-Méndez M. 2005. Phylogeny of slug species of the genus Arion: evidence of monophyly of Iberian endemics and of the existence of relict species in Pyrenean refuges. Journal of Zoological Systematics and Evolutionary Research 43: 139–148. https://doi.org/10.1111/j.1439-0469.2005.00307.x
Rabitsch W. 2006. DAISIE – Arion vulgaris (Moquin-Tandon, 1855) Fact Sheet. – Online Database of Delivering Alien Invasive Species Inventories for Europe. http://www.europe-aliens.org accessed 14.viii.2019.
Reischütz P. L. 1984. Zum massenhaften Auftreten von Arion lusitanicus Mabille in den Jahren 1982 und 1983. Mitteilungen der zoologischen Gesellschaft Braunau 4: 253–254.
Reischütz P. L., Seidl F. 1982. Gefährdungsstufen der Mollusken Österreichs. Mitteilungen der zoologischen Gesellschaft Braunau 4: 117–128.
Reise H. 2013. Wie seziert man Nacktschnecken? Mitteilungen der deutschen malakozoologischen Gesellschaft 88: 29–32. http://www.dmg.mollusca.de/images/mitteilungen_dmg/mitteilungen088/mitt_dmg_088_029-032_reise.pdf
Reise H., Backeljau T., Seidel D. 1996. Erstnachweise dreier Schneckenarten und weitere malakofaunistisch bemerkenswerte Funde aus der Oberlausitz. Berichte der naturforschenden Gesellschaft der Oberlausitz 5: 39–47. https://naturforschende-gesellschaft-der-oberlausitz.de/sites/default/files/pdf/bd05_03_reise-backeljau-seidel_schnecken.pdf
Reise H., Schlitt B., Hutchinson J. M. C. 2018a. Bericht über die 33. Regionaltagung des Arbeitskreises Ost der DMG in Ostritz bei Görlitz vom 23. bis 25. September 2016: Mollusken des Rotsteins bei Sohland, Sachsen. Mitteilungen der deutschen malakozoologischen Gesellschaft 98: 35–44. http://www.dmg.mollusca.de/images/mitteilungen_dmg/mitteilungen098/mitt_dmg_098_035-044_reise.pdf
Reise H., Arslangündoğdu Z., Schlitt B., Hutchinson J. M. C., Hizal E., Bacack E. 2018b. First records of the terrestrial slug Arion ater s.l. (Linnaeus, 1758) (Pulmonata: Arionidae) from Turkey. Folia Malacologica 26: 213–220. https://doi.org/10.12657/folmal.026.024
Regteren Altena C. O. van 1956. Notes sur les Limaces. 3. Sur la présence en France d’ Arion lusitanicus Mabille. Journal de Conchyliologie 45: 89–99.
Regteren Altena C. O. van 1963. Notes sur les limaces. 8. Sur l’état de deux espèces nominales du genre Arion. Basteria 27: 1–6. http://natuurtijdschriften.nl/download?type=document&docid=596445
Rhymer J. M., Simberloff D. 1996. Extinction by hybridization and introgression. Annual Review of Ecology and Systematics 27: 83–109. https://doi.org/10.1146/annurev.ecolsys.27.1.83
Risch P., Backeljau T. 1989. On the occurrence of Arion lusitanicus Mabille, 1868 in Belgium (Mollusca: Pulmonata). Annales de la Société royale zoologique de Belgique 118: 25–38.
Roth S., Hatteland B. A., Solhøy T. 2012. Some notes on reproductive biology and mating behaviour of Arion vulgaris Moquin-Tandon 1855 in Norway including a mating experiment with a hybrid of Arion rufus (Linnaeus 1758) x ater (Linnaeus 1758). Journal of Conchology 41: 249–257.
Rowson B., Anderson R., Turner J. A., Symondson W. O. C. 2014a. The slugs of Britain and Ireland: undetected and undescribed species increase a well-studied, economically important fauna by more than 20%. PLoS ONE 9(3): e91907. https://doi.org/10.1371/journal.pone.0091907
Rowson B., Turner J. A., Anderson R., Symondson W. O. C. 2014b. Slugs of Britain and Ireland. FSC Publications, Telford.
Rüetschi J., Stucki P., Müller P., Vicentini H., Claude F. 2012. Rote Liste Weichtiere (Schnecken und Muscheln). Gefährdete Arten der Schweiz, Stand 2010. Bundesamt für Umwelt, Bern und Schweizer Zentrum für die Kartographie der Fauna, Neuenburg. Umwelt-Vollzug Nr. 1216. https://www.bafu.admin.ch/bafu/de/home/themen/biodiversitaet/publikationen-studien/publikationen/rote-liste-weichtiere.html
Ryser S., Rindlisbacher N., Grüebler M. U., Knop E. 2011. Differential survival rates in a declining and an invasive farmland gastropod species. Agriculture, Ecosystems & Environment 144: 302–307. https://doi.org/10.1016/j.agee.2011.08.005
Schileyko A. A. 2007. Treatise on recent terrestrial pulmonate molluscs. Part 15: Oopeltidae, Anadenidaje, Arionidae, Philomycidae, Succineidae, Athoracophoridae. Ruthenica, Suppl. 2: 2049–2210.
Schmid G. 1972. Nacktschnecken aus Baden-Württemberg. Mitteilungen der deutschen malakozoologischen Gesellschaft 2: 332–344.
Schmid G. 1997. “Malakologische Zuckungen” Moment­aufnahmen zur Molluskenfauna Baden-Württembergs. Veröffentlichungen für Naturschutz und Landschaftspflege in Baden-Württemberg 71/72: 719–858.
Seifert B., Kulmuni J., Pamilo P. 2010. Independent hybrid populations of Formica polyctena X rufa wood ants (Hymenoptera: Formicidae) abound under conditions of forest fragmentation. Evolutionary Ecology 24: 1219–1237. https://doi.org/10.1007/s10682-010-9371-8
Simon C., Frati F., Beckenbach A., Crespi B., Liu H., Flook P. 1994. Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87: 651–701. https://doi.org/10.1093/aesa/87.6.651
Simroth H. 1885. Versuch einer Naturgeschichte der deutschen Nacktschnecken und ihrer europäischen Verwandten. Zeitschrift für wissenschaftliche Zoologie 42: 203–366, Pl. VII–XI. https://biodiversitylibrary.org/page/45468576
Slotsbo S., Fisker K. V., Hansen L. M., Holmstrup M. 2011b. Drought tolerance in eggs and juveniles of the Iberian slug, Arion lusitanicus. Journal of Comparative Physiology B 181: 1001–1009. https://doi.org/10.1007/s00360-011-0594-y
Slotsbo S., Hansen L. M., Holmstrup M. 2011a. Low temperature survival in different life stages of the Iberian slug, Arion lusitanicus. Cryobiology 62: 68–73. https://doi.org/10.1016/j.cryobiol.2010.12.005
Slotsbo S., Hansen L. M., Jordaens K., Backeljau T., Malmendal A., Nielsen N. C., Holmstrup M. 2012. Cold tolerance and freeze-induced glucose accumulation in three terrestrial slugs. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology 161: 443–449. https://doi.org/10.1016/j.cbpa.2012.01.002
Smith A. H. 1961. The place-names of the West Riding of Yorkshire. Part II. Osgoldcross and Agbrigg Wapentakes. English Place-Name Society, Vol. XXXI. Cambridge University Press, Cambridge.
Taylor J. W. 1902–1907. Monograph of the land and freshwater Mollusca of the British Isles. Testacellidae, Limacidae, Arionidae. Taylor Brothers, Leeds.
Taylor J. E. 1975. “Ambry”. In: Historic Almondbury: the village on the hill. Kirklees Libraries and Museum Service, pp. 7–10.
Welter-Schultes F. W., Klug R., Lutze A. 2008. Les figures des plantes et animaux d’usage en medecine, a rare work published by F. A. P. de Garsault in 1764. Archives of Natural History 35: 118–127. https://doi.org/10.3366/E0260954108000119
Wiktor A. 1973. Die Nacktschnecken Polens – Arionidae, Milacidae, Limacidae (Gastropoda, Stylommatophora). Monografie Fauny Polski, Vol. 1. Państwowe Wydawnictwo Naukowe, Warszawa.
Wiktor A. 2004. Ślimaki lądowe Polski. Wydawnictwo Mantis, Olsztyn.
Winter A. J. de 1984. The Arion hortensis complex (Pulmo­nata: Arionidae): designation of types, descriptions, and distributional patterns, with special reference to the Netherlands. Zoologische Mededelingen. 59: 1–17. https://www.repository.naturalis.nl/document/150316
Winter A. J. de 1989. Arion lusitanicus Mabille in Nederland (Gastropoda, Pulmonata, Arionidae). Basteria 53: 49–51.
Zemanova M. A., Broennimann O., Guisan A., Knop E., Heckel G. 2018. Slimy invasion: climatic niche and current and future biogeography of Arion slug invaders. Diversity and Distributions 2018: 1–14. https://doi.org/10.1111/ddi.12789
Zemanova M. A., Knop E., Heckel G. 2015. Development and characterization of novel microsatellite markers for Arion slug species. Conservation Genetic Resources 7: 501–503. https://doi.org/10.1007/s12686-014-0406-2
Zemanova M. A., Knop E., Heckel G. 2016. Phylogeo­graphic past and invasive presence of Arion pest slugs in Europe. Molecular Ecology 25: 5747–5764. https://doi.org/10.1111/mec.13860
Zemanova M. A., Knop E., Heckel G. 2017. Introgressive replacement of natives by invading Arion pest slugs. Scientific Reports 7: 41908. https://doi.org/10.1038/s41598-017-14619-y
Zettler M. L., Jueg U., Menzel-Harloff H., Göllnitz U., Petrick S., Weber E., Seemann R. 2006. Die Land- und Süßwassermollusken Mecklenburg-Vorpommerns. Obotritendruck, Schwerin.
Zilch A. 1962. Ergänzungen und Berichtigungen zur Nomenklatur und Systematik. In: Brohmer P., Ehr­mann P., Ulmer G. (eds). Die Tierwelt Mitteleuropas Vol. II (1), Ergänzung. Quelle & Meyer, Leipzig, pp. 1–23.

Folia Malacologica (2020) 28: 1-34
First published on-line: 2020-03-13 00:00:00
https://doi.org/10.12657/folmal.028.001
Full text (.PDF) BibTeX Mendeley Back to list