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Type: Articles
Published: 2010-05-14
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A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family

School of Biological Sciences, University of Sydney, NSW 2006, Australia
Australian Museum, 6 College St Sydney, NSW 2010, Australia
Mollusca Helicarionoidea Ariophantidae Urocyclidae Gastrodontoidea Euconulidae Euconulinae Microcystinae Trochomorphidae taxonomy systematics morphology phylogeny

Abstract

Helicarionidae is a diverse and widely distributed group whose delineation and phylogenetic relationships are poorly understood. We review the helicarionid fauna of Australia (excluding Lord Howe Island and Norfolk Island) and carry out a phylogenetic analysis based on 54 morphological characters. The morphology of 22 Australian helicarionid species from 17 genera is described, including one new genus (Stanisicarion) and two new combinations (Stanisicarion freycineti, Stanisicarion virens). The taxonomic positions of the remaining Australian genera are discussed. In addition, morphological descriptions of 24 taxa closely related to Helicarionidae are presented. Morphological characters indicate that Euconulidae (with subfamilies Euconulinae and Microcystinae), Trochomorphidae and Cystopeltidae can be excluded from Helicarionoidea, supporting recent phylogenies based on molecular data. The Helicarionidae is redefined and most endemic Australian mainland taxa previously assigned to Euconulinae, Durgellinae and Sesarinae can be included within the family as redefined. Five Australian genera (Coneuplecta, Euconulus, Turrisitala, Wilhelminaia and Queridomus) are moved to Euconulinae, one (Liardetia) to Microcystinae and one (Theskelomensor) to Trochomorphidae, leaving 24 genera and 47 species of mainland Australian Helicarionidae. Our results show poor support for the monophyly of Helicarionidae and no synapomorphies can be identified. Helicarionoidea, which is monophyletic in some other recent morphological and molecular analyses, is supported by the shared presence of a flagellum with glandular walls and an axial filament and the presence of an epiphallic retractor caecum. However, one or both of these characters are missing in some helicarionoid taxa. Morphological characters alone give little support for branching patterns among the Australian taxa. This is partly due to high levels of convergence and loss in the data.

References

  1. Albers, J.C. & Martens, E. von. (1860) Die Heliceen nach natürlicher Verwandtschaft systematisch geordnet. Zweite Ausgabe. Engelmann, Leipzig, 1–359.

    Baker, H.B. (1938) Zonitid snails from Pacific Islands Part 1: Southern genera of Microcystinae. Bernice P. Bishop Museum Bulletin, 158, 1–102.

    Baker, H.B. (1940) Zonitid snails from Pacific Islands Part 2: Hawaiian genera of Microcystinae. Bernice P. Bishop Museum Bulletin, 165, 105–201.

    Baker, H.B. (1941a) Zonitid snails from Pacific Islands Part 3: Genera other than Microcystinae. Bernice P. Bishop Museum Bulletin, 166, 205–346.

    Baker, H.B. (1941b) Zonitid snails from Pacific Islands Part 4: Distribution and indexes. Bernice P. Bishop Museum Bulletin, 166, 347–370.

    Baker, H.B. (1955) Heterurethrous and aulacopod. The Nautilus, 68, 109–112.

    Barr, R.A. (1927) Mucous and skin glands of Arion. Quarterly Journal of Microscopical Science, 71, 503–525.

    Beck, H. (1837) Index Molluscorum Praesentis Aevi Musei Principis Augustissimi Christianni Frederici. Vol.1. Mollusca Gastropoda Pulmonata. Hafniae, Beck, 124 pp.

    Benthem-Jutting, W.S.S. van (1950) Systematic studies on the non-marine Mollusca of the Indo-Australian Archipelago. II. Critical revision of the Javanese pulmonate land-shells of the families Helicarionidae, Pleurodontidae, Fruticolidae and Streptaxidae. Treubia, 21, 381–505.

    Benthem-Jutting, W.S.S. van (1964) Non-marine Mollusca of West New Guinea. Part 3, Pulmonata, I. Nova Guinea, Zoology, 32, 205–304.

    Blainville, H.M.D. de (1824) Dictionnaire des sciences naturelles dans lequel on traite méthodiquement des différens êtres de la nature. Levrault (1816–30), Paris & Strasbourg.

    Blanford, W.T. & Godwin-Austen, H.H. (1908) Fauna of British India, including Ceylon and Burma; Mollusca, Volume 1: Testacella and Zonitidae. Taylor and Francis, London.

    Bonham, K. (2003) Biogeography of Tasmanian Native Land Snails. University of Tasmania, Hobart.

    Boss, K.J. (1982) Mollusca. In: Parker, S.P. (Ed.) Synopsis and Classification of Living Organisms. McGraw-Hill, New York, pp. 945–1166.

    Bouchet, P. & Rocroi, J.-P. (2005) Classification and nomenclator of gastropod families. Malacologia, 47, 1–397.

    Bourguignat, J.-R. (1877) Description de deux nouveaux genres algériens, suivi d’une classification des familles et des genres de Mollusques terrestres et fluviatiles du système européen. Bulletin de la Société des Sciences Physiques et Naturelles de Toulouse, 3, 49–101.

    Brazier, J. (1875a) Description of fourteen new species of terrestrial, fluviatile and marine shells from Australia and the Solomon Islands. Proceedings of the Linnaean Society of New South Wales, 1, 1–9.

    Brazier, J. (1875b) Descriptions of eleven new species of terrestrial and marine shells from north-east Australia. Proceedings of the Zoological Society of London, 1874, 668–672.

    Brazier, J. (1876a) Descriptions of thirty-five new species of land shells from New Guinea, Australia, and islands in the Torres Straits, collected during the Chevert Expedition. Proceedings of the Linnean Society of New South Wales, 1, 98–113.

    Brazier, J. (1876b) List of land shells collected during the Chevert Expedition. Proceedings of the Linnean Society of New South Wales, 1, 117–133.

    Burch, J.B. (1976) Outline of classification of Australian terrestrial molluscs (native and introduced). Journal of the Malacological Society of Australia, 3, 127–156.

    Chung, D.J.D. (1987) Courtship and dart shooting behaviour of the land snail Helix aspersa. The Veliger, 30, 24–39.

    Climo, F.M. (1969) Classification of New Zealand Arionacea (Mollusca, Pulmonata): I. The higher classification. Records of the Dominion Museum, 6, 145–158.

    Cox, J.C. (1864) Catalogue of Specimens of the Australian Land Shells in the Collection of James C. Cox. John Alex Engel, Sydney, 44 pp.

    Cox, J.C. (1865) Descriptions of four new species of Australian land shells. Proceedings of the Zoological Society of London 1864, 594–596.

    Cox, J.C. (1866) Description d'espèces nouvelles provenant d'Australie et des îles Solomon et Norfolk. Journal de Conchyliologie, 14, 45–48.

    Cox, J.C. (1868) A Monograph of Australian Land Shells. William Maddock, Sydney, 110 pp.

    Cox, J.C. (1871) Descriptions of seven new species of Australian land shells. Proceedings of the Zoological Society of London, 1871, 53–55.

    Cox, J.C. (1873) Descriptions of new land-shells from Australia and the Solomon Islands. Proceedings of the Zoological Society of London, 1873, 146–152.

    Cox, J.C. (1887) Contributions to Conchology, No. 1. Proceedings of the Linnaean Society of New South Wales, Series 2, 2, 1061–1064.

    Cox, J.C. (1909) A list of the subclass Pulmonata found in Australia, Part I., not including Tasmania, Lord Howe's Island (under the NSW Government), or the New Guinea Mainland and adjacent islands. Frederick W. White, Sydney, 23 pp.

    Dallwitz, M.J., Paine, T.A. & Zurcher, E.A. (2000) DELTA: Description Language for Taxonomy. CSIRO, Canberra.

    Dartnall, A.J. & Kershaw, R.C. (1978) Description of a new species of Helicarion (Stylommatophora: Helicarionidae) in Tasmania. Records of the Queen Victoria Museum, 62, 1–17.

    Dasen, D.D. (1933) Structure and function of the reproductive system in Ariophanta ligulata. Proceedings of the Zoological Society of London, 1933, 97–118.

    Emberton, K. (1991) Polygrid relations: a phylogenetic analysis of 17 subfamilies of land snails (Mollusca: Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society, 103, 207–224.

    Emberton, K.C., Kuncio, G.S., Davis, G.M., Phillips, S.M., Monderewicz, K.M. & Guo, Y.H. (1990) Comparison of recent classifications of stylommatophoran land-snail families, and evaluation of large-ribosomal-RNA sequencing for their phylogenetics. Malacologia, 31, 327–352.

    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.

    Férussac, A.E.J.P.J.F.d'Audebard de (1821) Tableaux systématiques des animaux mollusques classés en familles naturelles, dans lesquels on a établi la concordance de tous les systèmes; suivis d'un Prodrome général pour tous les mollusques terrestres ou fluviatiles, vivants ou fossiles. A. Bertrand, Paris, 184 pp.

    Fischer-Piette, E., Blanc, C.P., Blanc, F. & Salvat, F. (1994) Gastéropodes Terrestres Pulmonés (excl. Veronicellidae et g. Elisolimax). Faune de Madagascar 83. Muséum Natural d'Histoire Naturelle, Paris, 1–552 pp.

    Forbes, E. (1851) On the Mollusca collected by Mr MacGillivray during the voyage of the Rattlesnake. In: MacGillivray, J. (Ed.), Narrative of the Voyage of the H.M.S. Rattlensnake, Commanded by the Late Captain Owen Stanley, During the Years 1846–1850, 2. T. & W. Boone, London.

    Franc, A. (1957) Mollusques terrestres et fluviatiles de l'Archipel Néo–Calédonien. Memoires du Muséum National d'Histoire Naturelle, S érie A, Zoologie., 13, 1–200.

    Gabriel, C.J. (1930) Catalogue of the land shells of Victoria. Proceedings of the Royal Society of Victoria, 43, 62–88.

    Godwin-Austen, H.H. (1882a) Land and Freshwater Mollusca of India, Including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burmah, Pegu, Tenasserim, Malay Peninsula, Ceylon, and Other Islands of the Indian Ocean. Part I. Supplementary to Messrs. Theobald and Hanley's Conchologica Indica. Taylor and Francis, London.

    Godwin-Austen, H.H. (1882b) Land and Freshwater Mollusca of India, Including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burmah, Pegu, Tenasserim, Malay Peninsula, Ceylon, and Other Islands of the Indian Ocean. Part II. Supplementary to Messrs. Theobald and Hanley's Conchologica Indica. Taylor and Francis, London.

    Godwin-Austen, H.H. (1883a) Land and Freshwater Mollusca of India, Including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burmah, Pegu, Tenasserim, Malay Peninsula, Ceylon, and Other Islands of the Indian Ocean. Part III. Supplementary to Messrs. Theobald and Hanley's Conchologica Indica. Taylor and Francis, London.

    Godwin-Austen, H.H. (1883b) Land and Freshwater Mollusca of India, Including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burmah, Pegu, Tenasserim, Malay Peninsula, Ceylon, and Other Islands of the Indian Ocean. Part IV. Supplementary to Messrs. Theobald and Hanley's Conchologica Indica. Taylor and Francis, London.

    Godwin-Austen, H.H. (1888) Land and Freshwater Mollusca of India, Including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burmah, Pegu, Tenasserim, Malay Peninsula, Ceylon, and Other Islands of the Indian Ocean. Part VI. Supplementary to Messrs. Theobald and Hanley's Conchologica Indica. Taylor and Francis, London.

    Gray, J.E. (1834) Characters of new species of shells. Proceedings of the Zoological Society of London, 1834, 63–68.

    Gude, G.K. (1905) Descriptions of nine new species of helicoid land shells. Journal of Malacology, 12, 11–14.

    Gude, G.K. (1911) Note on some preoccupied molluscan generic names and proposed new genera of the family Zonitidae. Proceedings of the Malacological Society of London, 9, 269–273.

    Gude, G.K. & Woodward, B.B. (1921) On Helicella, Férussac. Proceedings of the Malacological Society of London, 14, 174–190.

    Hausdorf, B. (1995) A preliminary phylogenetic and biogeographical analysis of the Dyakiidae (Gastropoda: Stylommatophora) and a biogeographic analysis of other Sundaland taxa. Cladistics, 11, 359–376.

    Hausdorf, B. (1998) Phylogeny of the Limacoidea sensu lato (Gastropoda: Stylommatophora). Journal of Molluscan Studies, 64, 35–66.

    Hedley, C. (1888) A list of the land shells recorded from Queensland. Royal Society of Queensland, 5, 45–70.

    Hedley, C. (1890) On Parmella etheridgei, Brazier. Records of the Australian Museum, 1, 78–80.

    Hedley, C. (1891a) The land and freshwater shells of Lord Howe Island. Records of the Australian Museum, 1, 134–143.

    Hedley, C. (1891b) On the anatomy of some Tasmanian snails. Proceedings of the Linnean Society of New South Wales (Series 2), 6, 19–26.

    Hedley, C. (1912) On some land shells collected in Queensland by Mr Sidney W. Jackson. Proceedings of the Linnean Society of New South Wales, 37, 259–270.

    Hedley, C. (1916) A preliminary index of the Mollusca of Western Australia. Journal of the Royal Society of Western Australia, 1, 1–77.

    Hollingsworth, R.G., Kaneta, R.K., Sullivan, J.J., Bishop, H.S., Qvarnstrom, Y., Da Silva, A.J., Robinson, D.G. (2007) Distribution of Parmarion cf. martensi (Pulmonata: Helicarionidae), a new semi-slug pest on Hawai`i Island, and its potential as a vector for Human Angiostrongyliasis. Pacific Science, 61, 457–467.

    Hyman, I.T. (2005) Taxonomy, systematics and evolutionary trends in Helicarionidae (Mollusca, Pulmonata), PhD thesis, University of Sydney, 583pp.

    Hyman, I.T. (2007) Three new genera and five new species of Helicarionidae from southeastern Australia (Pulmonata: Stylommatophora: Helicarionoidea). Molluscan Research, 27, 89–104.

    Hyman, I.T., Ho, S.Y.W. & Jermiin, L.S. (2007) Molecular phylogeny of Australian Helicarionidae, Euconulidae and related groups (Gastropoda: Pulmonata: Stylommatophora) based on mitochondrial DNA. Molecular Phylogenetics and Evolution, 45, 792–812.

    ICZN. (1992) Opinion 1678. Helicarion Férussac, 1821 (Mollusca, Gastropoda) conserved, and Helicarion cuvieri Férussac, 1821 as the type species. Bulletin of Zoological Nomenclature, 49, 160–161.

    Iredale, T. (1933) Systematic notes on Australian land shells. Records of the Australian Museum, 19, 37–59.

    Iredale, T. (1937a) An annotated checklist of the land shells of South and Central Australia. South Australian Naturalist, 18, 6–59.

    Iredale, T. (1937b) A basic list of the land Mollusca of Australia. Australian Zoologist, 8, 287–333.

    Iredale, T. (1937c) A basic list of the land Mollusca of Australia―part II. Australian Zoologist, 9, 1–39.

    Iredale, T. (1939) A review of the land Mollusca of Western Australia. Journal of the Royal Society of Western Australia, 25, 1–88.

    Iredale, T. (1941) Guide to the land shells of New South Wales―part III. The Australian Naturalist, 11, 1–8.

    Iredale, T. (1942) Guide to the land shells of New South Wales―part IV. The Australian Naturalist, 11, 33–40.

    Iredale, T. (1944) The land Mollusca of Lord Howe Island. Australian Zoologist, 10, 299–334.

    Kershaw, R.C. (1979) Redescription of Helicarion cuvieri from southern Tasmania and Helicarion freycineti from New South Wales (Pulmonata: Helicarionidae). Journal of the Malacological Society of Australia, 4, 145–156.

    Kershaw, R.C. (1980) Notes on Helicarion rubicundus (Pulmonata: Helicarionidae). Journal of the Malacological Society of Australia, 4, 213–214.

    Kershaw, R.C. (1981) Redescription of the genus Helicarion and of Helicarion niger (Quoy & Gaimard, 1832) from Victoria (Stylommatophora: Helicarionidae). Journal of the Malacological Society of Australia, 5, 17–31.

    Landolfa, M.A., Green, D.M. & Chase, R. (2001) Dart shooting influences paternal reproductive success in the snail Helix aspersa (Pulmonata, Stylommatophora). Behavioural Ecology, 12, 773–777.

    Legrand, W. (1871) Collections for a Monograph of Tasmanian Land Shells. W. Legrand, Hobart.

    Moellendorff, O.F. von. (1893) Materialien zur Fauna der Philippinen. XI. Die Insel Leyte. Bericht der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main, 1893, 51–154.

    Muratov, I.V. (1999) Analysis of the phylogenetic relationships and their systematic implications in the Limacoinei (= Zonitinia) infraorder (Gastropoda, Pulmonata, Geophila). Ruthenica, 9, 5–26.

    Nordsieck, H. (1985) The system of the Stylommatophora (Gastropoda), with special regard to the systematic plan of the Clausiliidae, 1. Importance of the excretory and genital systems. Archiv für Molluskenkunde, 116, 1–24.

    Nordsieck, H. (1986) The system of the Stylommatophora (Gastropoda), with special regard to the systematic position of the Clausiliidae, 2. Importance of the shell and distribution. Archiv für Molluskenkunde, 117, 93–116.

    Nordsieck, H. (1993) Phylogeny and system of the Pulmonata (Gastropoda). Archiv für Molluskenkunde, 121, 31–52.

    Odhner, N.H.J. (1917) Results of Dr. E. Mjöbergs Swedish Scientific Expeditions to Australia 1910–1913. XVII. Mollusca. Kungliga Svenska Vetenskapsakademiens Handlingar., 52, 1–115.

    Petterd, W.F. (1879) A Monograph of the Land Shells of Tasmania. Privately published, Launceston.

    Pfeiffer, L. (1846) Die Schnirkelschnecken (Gattung Helix) in Abbildungen nach der Natur, mit Beschreibungen. Systematisches Conchylien-Cabinet von Martini und Chemnitz. 1 (12). Bauer und Raspe, Nürnberg. 400 pp.

    Pfeiffer, L. (1849) Description of a new Helix and Streptaxis, from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London, 1848, 108–109.

    Pfeiffer, L. (1850) Descriptions of twelve new species of Vitrina and Succinea from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London, 1849, 132–134.

    Pfeiffer, L. (1852) Bemerkungen über Deshayes's Bearbeitung des Férussacschen Werkes. Zeitschrift für Malakozoologie, 12, 97–112.

    Pfeiffer, L. (1853) Monographia Heliceorum Viventium. Volumen Tertium. F. A. Brockhaus, Lipsiae. 711 pp.

    Pfeiffer, L. (1855) Descriptions of fifty-seven new species of Helicea, from Mr Cuming's collection. Proceedings of the Zoological Society of London, 1854, 286–298.

    Pfeiffer, L. (1876) Monographia Heliceorum Viventium. Volumen Septimus. F. A. Brockhaus, Lipsiae. 674 pp.

    Pfeiffer, L. & Clessin, S. 1881. Nomenclator heliceorum viventium quo continetur nomina omnium hujus familiae generum et specierum hodie cognitarum, disposita ex affinitate naturali. Th. Fischer, Cassellis. 617 pp.

    Pilsbry, H.A. (1900) On the zoological position of Partula and Achatinella. Proceedings of the Academy of Natural Sciences, Philadelphia, 52, 561–567.

    Quoy, J.R.C. & Gaimard, J.P. (1824) Description des Mollusques. In: Freycinet, M.L. (Ed.), Voyage Autour du Monde, entrepris par ordre du Roi, sous le Ministère et conformément aux la instructions de S. Exc. M. le Viscomte de Bouchage, Secrétaire d'État au Département de la Marine, exécute sur les Corvettes de S. M. l'Uraine et la Physicienne, pendant les années 1817–1820. Zoologie. Pillet Ainé, Paris. 671 pp.

    Raheem, D. & Naggs, F. (2005) The Sri Lankan endemic semi-slug Ratnadvipia (Limacoidea: Ariophantidae) and a new species from southwestern Sri Lanka. Systematics and Biodiversity, 4, 99–126.

    Reeve, L.A. (1852) Monograph of the genus Helix, part I. In: Reeve, L.A. (Ed.) Conchologia Iconica: or, Illustrations of the Shells of Molluscous Animals, VII. Lovell Reeve: London.

    Reeve, L.A. (1854) Monograph of the genus Helix, part II. In: Reeve, L.A. (Ed.) Conchologia Iconica: or, Illustrations of the Shells of Molluscous Animals, VII. Lovell Reeve, London.

    Reeve, L.A. (1862) Monograph of the genus Vitrina. In: Reeve, L.A. (Ed.) Conchologia Iconica: or, Illustrations of the Shells of Molluscous Animals, XIII. Lovell Reeve, London.

    Rensch, I. (1937) Systematische und tiergeographische Untersuchungen über die Landschneckenfauna des Bismarck-Archipels. II. Archiv für Naturgeschichte, 6, 526–644.

    Rensch, I. & Rensch, B. (1936) Systematische und tiergeographische Studien über die Landschnecken der Salomonen auf Grund der Sammlungen von Dr. A. Paravicini und Dr. H. Hediger. Revue Suisse de Zoologie, 43, 653–695.

    Richter, K.O. (1980) Movement, reproduction, defense and nutrition as functions of the caudal mucous in Ariolimax columbianus. Veliger, 23, 43–47.

    Schileyko, A.A. (1979) The system of the order Geophila (=Helicida) (Gastropoda Pulmonata). Department of Molluscs, Harvard University, Special Occasional Publication, 6 [1985], 1–45.

    Schileyko, A.A. (1984) Nazemnye mollyuski podotrada Pupillina fauny SSSR (Gastropoda, Pulmonata, Geophila). Fauna SSSR, Mollyuski, III (3), Nauka, Leningrad.

    Schileyko, A.A. (2002a) Treatise on Recent Terrestrial Pulmonate Molluscs, Part 8: Punctidae, Helicodiscidae, Discidae, Cystopeltidae, Euconulidae, Trochomorphidae. Ruthenica, Supplement 2, Moscow.

    Schileyko, A.A. (2002b) Treatise on Recent Terrestrial Pulmonate Molluscs, Part 9: Helicarionidae, Gymnarionidae, Rhysotinidae, Ariophantidae. Ruthenica, Supplement 2, Moscow.

    Schileyko, A.A. (2003a) Treatise on Recent Terrestrial Pulmonate Molluscs, Part 10: Ariophantidae, Ostracolethidae, Ryssotidae, Milacidae, Dyakiidae, Staffordiidae, Gastrodontidae, Zonitidae, Daudebardiidae, Parmacellidae. Ruthenica, Supplement 2, Moscow.

    Schileyko, A.A. (2003b) Treatise on Recent Terrestrial Pulmonate Molluscs, Part 11: Trigonochlamydidae, Papillodermidae, Vitrinidae, Limacidae, Bielziidae, Agriolimacidae, Boettgerillidae, Camaenidae. Ruthenica, Supplement 2, Moscow.

    Scott, B. (1995) Redescription of Fastosarion superba (Cox, 1871) with descriptions of two new species of Fastosarion from northern Queensland (Pulmonata: Helicarionidae). Molluscan Research, 16, 69–80.

    Semper, C. (1870) Reisen im Archipel der Philippinen 2(3). Wissenschaftliche Resultate. Landmollusken. C. W. Kreidel, Wiesbaden, 327 pp.

    Simroth, H. (1889) Beiträge zur Kenntnis der Nachtschnecken. Nova Acta. Abhandlungen der Kaiserlich Leopoldinischen-Carolinischen Deutschen Akademie der Naturforscher, 54, 1–91.

    Simroth, H. (1898) Über die Gattungen Parmacochlea, Parmarion und Microparmarion. Zoologischen Jahrbüchern. Abteilung für Systematik, Geographie und Biologie der Thiere, 11, 151–172.

    Smith, B.J. (1965) The secretions of the reproductive tract of the garden slug Arion ater. Annals of the New York Academy of Sciences, 118, 997–1014.

    Smith, B.J. (1992) Non-marine Mollusca. In: Houston, W.W.K. (Ed.) Zoological Catalogue of Australia. Australian Government Publishing Service, Canberra, pp. 408.

    Smith, B.J. (1998) Family Helicarionidae. In: Beesley, P.L., Ross, G.J.B. & Wells, A. (Eds.) Mollusca: The Southern Synthesis. Fauna of Australia. Vol. 5. CSIRO Publishing, Melbourne, pp. 1105–1106.

    Smith, B.J. & Stanisic, J. (1998) Pulmonata Introduction. In: Beesley, P.L., Ross, G.J.B. & Wells, A. (Eds.) Mollusca: The Southern Synthesis. Fauna of Australia. Vol. 5. CSIRO Publishing, Melbourne, pp. 1037–1061.

    Smith, B.J., Reid, S. & Ponder, W.F. (2002) Australian Faunal Directory, Mollusca: Pulmonata. Australian Biological Resources Study, http://www.deh.gov.au/cgi-bin/abrs/abif-fauna/tree.pl?pstrVol=PULMONATA&pintMode=1.

    Smith, E.A. (1884) An account of the land and freshwater Mollusca collected during a voyage of the "Challenger" from December 1872 to May 1876. Proceedings of the Zoological Society of London, 1884, 258–281.

    Smith, E.A. (1894) On the land shells of Western Australia. Proceedings of the Malacological Society of London, 1, 84–99.

    Solem, A. (1958) New land snail from Queensland. Nautilus, 72, 20–22.

    Solem, A. (1959) Systematics of the land and fresh-water Mollusca of the New Hebrides. Fieldiana Zoology, 43, 1–238.

    Solem, A. (1961) New Caledonian land and freshwater snails. An annotated check list. Fieldiana Zoology, 41, 415–501.

    Solem, A. (1966) Some non-marine mollusks from Thailand, with notes on classification of the Helicarionidae. Spolia Zoologica Musei Hauniensis, 24, 1–110.

    Solem, A. (1978) Chapter 2: Classification of the land Mollusca. In: Fretter, V. & Peake, J. (Eds.) Pulmonates Volume 2A. Systematics, Evolution and Ecology. Academic Press, London, New York, pp. 49–97.

    Solem, A. (1982) Small land snails from Northern Australia, II: Species of Westracystis Iredale, 1939 (Mollusca: Pulmonata, Helicarionidae). Journal of the Malacological Society of Australia, 5, 175–193.

    Solem, A. (1988) Non-camaenid land snails of the Kimberly and Northern Territory, Australia. I. Systematics, affinities and ranges. Invertebrate Taxonomy, 4, 455–604.

    Stanisic, J. (1988) A re-evaluation of the relationships of Helix (Thalassia) gayndahensis Brazier, 1875 (Mollusca, Pulmonata, Helicarionidae). Memoirs of the Queensland Museum, 25, 475–479.

    Stanisic, J. (1993a) Eungarion mcdonaldi gen. et n. sp., a montane semi-slug from mideastern Queensland rainforests (Pulmonata: Helicarionidae). Memoirs of the Queensland Museum, 34, 27–34.

    Stanisic, J. (1993b) The identity of Helicarion semoni Marten, 1894: a large semi-slug from the Wet Tropics, northeastern Queensland (Pulmonata: Helicarionidae). Memoirs of the Queensland Museum, 34, 1–9.

    Stanisic, J. (1998) A new semi-slug from Tamborine Mountain, Southeastern Queensland (Mollusca: Eupulmonata: Helicarionidae). Memoirs of the Queensland Museum, 42, 597–603.

    Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, Massachusetts.

    Thiele, J. (1928) Mollusken vom Bismarck-Archipel, von Neu-Guinea und Nachbar-Inseln. Zoologische Jahrbücher Abteilung für Systematik, Ökologie und Geographie der Tiere, 55, 119–146.

    Thiele, J. (1929–1931) Handbuch der Systematischen Weichtierkunde. Vol. 1. G. Fischer, Jena, 1189 pp.

    Tillier, S. (1984) Patterns of digestive tract morphology in the limacisation of helicarionid, succineid and athoracophorid snails (Mollusca: Pulmonata). Malacologia, 25, 173–192.

    Tillier, S. (1989) Comparative morphology, phylogeny and classification of land snails and slugs (Gastropoda: Pulmonata: Stylommatophora). Malacologia, 30, 1–303.

    Thon, D. (1829) Helixarion. In: Ersche, J.C. & Gruber, J.G. (Eds) Allgemeine Encyclopadie der Wissenschasten und Künste. 2. Section (H–N). Johann Friedrich Gleditsch, Leipzig, 149–150 pp.

    Tryon, G.W.j. (1885) Manual of Conchology; Structural and Systematic. Second Series: Pulmonata. Volume I. Testacellidae, Oleacinidae, Streptaxidae, Helicoidea, Vitrinidae, Limacidae, Arionidae. G.W. Tryon, jnr, Philadelphia.

    Tryon, G.W.j. (1886) Manual of Conchology; Structural and Systematic. Second Series: Pulmonata. Volume 2. Zonitidae. G.W. Tryon, jnr, Philadelphia.

    Van Goethem, J.L. (1977) Révision systématiques des Urocyclinae (Mollusca, Pulmonata, Urocyclidae). Musée Royal de l'Afrique Centrale, Tervuren, Belgique, Annales, Sciences Zoologiques, 218, 1–355.

    Van Mol, J.-J. (1968) Notes anatomiques sur les Helicarionidae (Gasteropodes Pulmonés). I. Etudes des genres Macrochlamys, Parmarion et Xesta. Annales de la Société Royale Zoologique de Belgique, 98, 189–215.

    Van Mol, J.-J. (1970a) Anatomical studies in the families Urocyclidae and Helicarionidae: Pulmonata, Stylommatophora, pp. 57–58. The American Malacological Union―Annual Reports for 1970.

    Van Mol, J.-J. (1970b) Révision des Urocyclidae (Mollusca, Gastropoda, Pulmonata): anatomie, systématique, zoogéographie. Première Partie. Musée Royal de l'Afrique Centrale, Tervuren, Belgique, Annales, Sciences Zoologiques, 180, 1–234.

    Van Mol, J.-J. (1973) Notes anatomiques sur les Helicarionidae (Mollusques, Gastéropodes. Pulmonés). II. Études des genres Pseudaustenia, Dyakia, Helicarion et comprenant la description de Papuarion, genre nouveau. Discussion sur la classification des Helicarionidae et les affinités des Urocyclidae. Annales de la Société Royale Zoologique de Belgique, 103, 209–237.

    Voigt, F.S. (1834) Das Thierreich, geordnet nach seiner Organisation als Grundlage der Naturgeschichte der Thiere und Einleitung in die vergleichende Anatomie. vom Baron von Cuvier. F. A. Brockhaus, Liepzig.

    Wächtler, W. (1935) Zur äusseren Morphologie des Fusses der monotrematen Land-Lungenschnecken. Jahrbücher der Akademie Gemeinnütziger Wissenschaften zu Erfurt, NF 52, 107–135.

    Wade, C.M., Mordan, P.B. & Clarke, B. (2001) A phylogeny of the land snails (Gastropoda: Pulmonata). Proceedings of the Royal Society of London, Series B: Biological Sciences, 268, 413–422.

    Wade, C.M., Mordan, P.B. & Naggs, F. (2006) Evolutionary relationships among the Pulmonate land snails and slugs (Pulmonata, Stylommatophora). Biological Journal of the Linnean Society, 87, 593–610.

    Zilch, A. (1959) Euthyneura. In: Schindewolf, O.H. (Ed.) Handbuch der Paläozoologie. Bornträger, Berlin-Zehlendorf.