Compiler and date details
31 May 2002 - Brian J. Smith, Queen Victoria Museum, Launceston, and Shannon Reid and Winston F. Ponder, Australian Museum, Sydney
This treatment of the Australian pulmonate fauna is a revision of the pulmonates in Smith (1992) with the addition of four pulmonate families.
The majority of Australian land molluscs and some of its freshwater groups are pulmonates. Smith (1992) dealt with the entire non-marine mollusc fauna, then comprising some 57 families, over 400 valid genus names with about 600 genus available names, and over 1 000 valid species names with over 2 000 species available names. It is estimated that about 30-40% of the non-marine fauna remains to be described, mainly from tropical regions. Details of the life cycle, food preferences and basic biology of almost the whole fauna awaits description. This volume, therefore, is not the definitive statement on the status of the fauna. It is another step in the documentation of the Australian fauna started by Cox (1864, 1868), in his catalogue and monograph of the Australian land shells, and continued by Iredale (1937a, 1937b, 1938, 1943a) in his basic lists of the entire non-marine mollusc fauna of Australia.
The family classification used is based on that outlined by Beesley, Ross & Wells (1998) and their classification has been modified where more recent information is available, or where our views differ in matters of emphasis or detail from this. Other arrangements are given by Zilch (1959), Burch (1976), Solem (1978), Hubendick (1978), Boss (1982) and Barker (1999).
The description of the Australian fauna has not proceeded at a uniform rate. Over the first 80 years of European settlement about 5-10% of the fauna was described from collections made by European voyages of exploration. During the next 50 years a further 25-30% of the fauna was described, primarily by specialists in Europe working on material collected by early settlers and sent over or purchased by European dealers. Gradually, a number of Australian specialists emerged who did the detailed work of documenting the various regional faunas. At the start of this century these regional bursts of activity were on the wane. In the 1930's and the 1940's, however, Iredale described a further 5-10% of the fauna and, from the 1970's up to the present, Solem, Stanisic, Kershaw, Smith and others have greatly increased our knowledge of the fauna.
Tom Iredale's view of the fauna has been criticised, both in print (e.g. Solem 1959) and privately by later workers on Australian molluscs, for his brief descriptions and lack of justification of his many new names and family and generic placements. Notwithstanding these criticisms, most of which are probably fully justified, without the ground-work laid by him in his basic lists (Iredale 1937a, 1937b, 1938, 1943a) and his major regional revisions (Iredale 1937c, 1939, 1940, 1941a, 1941b, 1942, 1943b, 1944, 1945) the compilation of Smith (1992) and this revision would have been much more difficult. We hope that this compilation will stimulate the much needed detailed study of the Australian molluscan fauna and give workers a firm basis on which continue to build on.
The pulmonates are contained within the Class Gastropoda. The gastropods comprise the largest and most diverse class of the Phylum Mollusca in terms of their taxonomic diversity and the range of habitats they occupy. Besides being found in every part of the marine environment, from high energy surge zones to the abyssal depths, gastropods are found in estuarine and freshwater habitats and are the only group of molluscs to have successfully colonized dry terrestrial habitats. The Subclass Heterobranchia contains the marine opisthobranchs and the largely terrestrial pulmonates.
The Pulmonata are so called because the gills or ctenidia seen in the prosobranchs have been greatly reduced or lost and respiration is carried out by the blood vessels in the wall of the mantle cavity, forming a "lung". Most pulmonates are non-marine in habit, either living in freshwater or on land. A few pulmonate families live in marine habitats. Five families, the Amphibolidae, Onchidiidae, Siphonariidae, Smeagolidae and Trimusculidae are completely marine while the Ellobiidae are marine and semi-terrestrial in the supralittoral zone. In some tropical areas outside Australia the ellobiids have a wider habitat preference, with some species being found away from areas of marine influence in forest litter. The Australian non-marine mollusc fauna contains 46 families of pulmonates, of which five only live in freshwater.
The single most striking feature of the pulmonate fauna is that 16 of the 46 families are not native to Australia and have been introduced, mostly in the last 200 years. In addition, the species from a further four families contain both native and introduced species, bringing the introduced pulmonate fauna to over 40 species in 20 families. Many of these are significant crop or garden pests or the carriers of major stock diseases. The most notable are the Helicidae, Hygromiidae, Agrolimacidae, Limacidae, Milacidae and Lymnaeidae. Thus far, no significant snail-borne disease of man has been identified in Australia.
The aquatic fauna is dominated by the Planorbidae in terms of diversity and distribution. The most unusual members of this family are the so-called "giant limpets" belonging to the genus Ancylastrum which are only found in the lakes and streams of the Central Plateau, Tasmania. The Glacidorbidae contains a diverse fauna in southern Australia, especially Tasmania and is otherwise known only from southern Chile. A species of the introduced family Physidae which is spread through the freshwater habitats of southern Australia.
The terrestrial pulmonate fauna can be divided into six regional subfaunas, each with a high degree of regional endemism. The family composition of the faunas are also different; each fauna is dominated by one or more families or genus-groups in terms of species diversity within the faunas. The family with the most genera and species in Australia is the Camaenidae and different genera make up the dominant groups of snails over much of tropical and subtropical Australia. The number of species declines as you move south and the family is absent from south-western Australia and Tasmania. Southern Western Australia is dominated by the Bulimulidae and the wetter parts of eastern Australia, including Tasmania, have large radiations of species of the Charopidae, Helicarionidae and Rhytididae. There are two families which are endemic to eastern Australia, the Caryodidae and the unusual group of slugs, the Cystopeltidae. The Australian pulmonate fauna contains 11 families of slugs, six of which are introduced.
For the majority of the Australian pulmonate fauna, the basic biology is poorly known and the habitat preferences and distribution can only be broadly estimated. Many groups are urgently in need of a modern taxononic revision to fully understand the composition of the fauna.
Many of the non-marine land snails are significantly threatened by habitat destruction, such as the coastal rainforests of eastern Australia. Many also have very narrow ranges and consequently, present special conservation concerns (Ponder 1998).
This work represents an updated and expanded treatment of the information presented in Smith (1992).
Classification of Pulmonata
Family Veronicellidae (= Vaginulidae)
Family Cionellidae (= Cochlicopidae)
The following acknowledgements were given by B.J. Smith for the 1992 Catalogue, which forms the basis of this work.
I am very grateful to Ms Penny Morison who carried out much of the initial library searches, wrote to many overseas museums concerning their type holdings and drafted most of the available name data sheets. Thanks are due to the Australian Biological Resources Study for the grant enabling Ms Morison to be employed for this vital preparatory work. I would also like to thank the Director and Council of the Museum of Victoria for permitting this work to be undertaken and for making all the facilities of the Museum available. Several of the staff of the Division of Zoology, particularly Ms Rhyll Plant, greatly assisted by carrying out some of my routine work allowing me time to work on this project.
Staff of the Natural History Library of the Museum of Victoria deserve special thanks for their help in finding many obscure references and processing innumerable requests for inter-library loans. I would also like to thank the libraries of the Australian Museum; the British Museum (Natural History); Christchurch College, Canterbury, UK; and the Royal Museum of Scotland for their special help in providing assistance and facilities.
I am grateful to the staff in charge of mollusc collections in both Australian and overseas museums for their hospitality, which allowed me to examine type holdings and reference collections. I am grateful, also, for their courteous assistance in replying to written enquiries about particular type holdings. In particular, I am grateful to Dr M. Bishop (University Museum, Cambridge, UK); Dr A. Bogan (ANSP); Dr K. Boss (MCZ); Dr J. Burch (UMMZ); Mr P. Colman (AM); Dr G. Davis (ANSP); Miss A. Green (TMH); Dr R. Green (QVM); Dr D. Heppell (RSM); Dr R. Janssen (SMF); Dr J. Knudsen (MZC); Mr I. Loch (AM); Dr A. Mead (University of Arizona, USA); Dr P. Mordan (BMNH); Ms S. Morris (BMNH); Mr F. Naggs (BMNH); Dr G. Oliver (NMW); Dr O. Paget (NHMW); Dr W. Ponder (AM); Dr R. Robertson (ANSP); Dr C. Roper (USNM); Mrs S. Slack-Smith (WAM); the late Dr A. Solem (FMNH); Dr J. Stanisic (QM); Dr J. Taylor (BMNH); Dr S. Tillier (MNHP); Mrs E. Turner (TMH); Mr W. Zeidler (SAMA); and Dr A. Zilch (SMF). For financial assistance, which enabled me to visit many overseas museums to consult reference collections, I would like to thank the Science and Industry Endowment Fund, the CSIRO, the British Council, the Myer Foundation, the Ian Potter Foundation and the Council of the Museum of Victoria.
Though the decisions about many controversial details in the taxonomic arrangements used in this work are mine, I am grateful for the many helpful discussions with colleagues about these problems. In particular, I would like to express my gratitude to Mr R. Burn, Dr F. Climo, Mr R. Kershaw, Dr W. Ponder, the late Dr A. Solem, Dr J. Stanisic and Dr S. Tillier. I am also greatly indebted to staff of the Australian Biological Resources Study, (formerly the Bureau of Flora and Fauna), for their help with this work, especially Dr W.W.K. Houston, Dr B. Richardson, Dr D. Walton, D. Brunckhorst and Dr G. Dyne, whose editorial attention to detail materially improved the Catalogue.
Finally, I would like to thank my many friends who helped me to cope with the volume of paper and the many lists and indices generated during this project. In particular I would like to thank Mrs E. Johns, Mrs B. Argo, Mr K. Bell, my two daughters Vivienne and Susan, and my wife Dr Helen Malcolm.
Acknowledgements for the On-line Catalogue
We greatly acknowledge the input from comments on drafts from Dr John Stanisic, Michael Shea, Ian Loch and Stephanie Clark. Information on some types was provided by Ms Kathie Way, Natural History Museum London and by Ian Loch of the Australian Museum.
We also indebted to Keith Houston and Kathy Tsang of ABRS for technical advice and editorial input, and to ABRS for funds which asssited with the update to this section of the Mollusca.
Synonymy and Incertae sedis
Complete Australian synonymy is given for each valid genus-group and species-group name. Only those synonyms that either have their type locality in Australia or have been recorded as occurring in Australia have been included. The relevant overseas lists should be consulted for the full international synonymies of those genera and species whose distribution extends outside Australia.
In most cases, the synonymies follow a published revision of the group and the authority for these synonymies is given. In a few cases expert opinions have been given by specialists that are based on unpublished research and these are given as personal communications (e.g. Stanisic, J., pers. comm.), meaning that the responsibility for the decision is ours but we have relied heavily on the advice of that particular worker. Where a worker is credited with the decision (e.g. Stanisic, J., this work), that person takes full responsibility for that decision. Where one of us takes credit for the decision without reference to another authority, we have given a brief justification for the decision.
There are a number of cases where there has been no recent work on a group and where neither I, nor any other specialist, was willing to make a decision about placement. In these cases, the doubtful taxa are placed under Incertae sedis at the end of the family, sometimes with qualifying notes to the likely placements or relationships of the taxon. It is often apparent to which genus a particular species-level taxon should be referred and this is indicated as a qualification. Families where this has been done to a significant extent, because of the poor state of knowledge of the group, include the Bulimulidae, Planorbidae and Succineidae. Other families require revisionary work in particular geographic regions of their distributions.
The original reference describing the name is given for each genus and species available name, including the page number on which the name first appears. In almost all cases the original description has been verified by one of us and every effort made to confirm the date of publication.
The type species, with author and date, of each genus available name, and its method of designation, is given. Where the type species was subsequently designated, the authority for that designation is given. Every effort has been made to check the original reference to all the designations, but these should be confirmed when revisionary work is being carried out. In the case of type species by subsequent designation, authoritative secondary references have been followed to get a pointer to that designation. If there was a conflict between different secondary references, further investigation was undertaken to settle the matter wherever possible. Where no conflict was encountered, however, exhaustive searches for unlikely possible earlier designations were not carried out.
Considerable effort was expended to find the types of every species available name. For many species the search was in vain and the types are presumed lost. Where the types were found, the status shown is the one put on the material by the holding institution (e.g. holotype, syntype), unless any reason was found to question that status. Where possible, all the relevant associated documentation, such as labels and register entries, was examined to confirm the standing of the material as types. Comparisons were made between the original description, any figures and the material itself. When disputes arose as to the status of any type material, such as claims by more than one institution to hold the holotype of a particular species, the scientists with curatorial responsibility for those collections were consulted and a mutually acceptable resolution achieved. Any worker revising any of these groups should personally recheck these findings rather than take these listings at face value.
Particular difficulties were encountered in trying to trace some of the type material of species introduced into Australia, especially those species native to Europe. Most of these species were described fairly soon after 1758 and, for many, type material never existed or has long since been lost. There are fewer details regarding types of the marine taxa owing to the lack of opportunity to check these in overseas institutions during the updating of this list.
In most cases, references other than the original reference helped to establish some of the information about the taxon. Many of these references are given with the taxon as secondary references to assist subsequent workers to retrace the steps taken to accumulate the data. In particular, the page number to the taxon in one of Iredale's basic lists (1937a, 1937b, 1938, 1943a) is given, together with references in Zilch (1959) and other standard listings or other works.
The Australian distribution of all valid species and subspecies can only be a best guess. Very few of these taxa have been accurately mapped over their whole range. In some cases, where the entire range of a species is much smaller than that suggested by the standard region and state listing, a brief description of the range also is given. The species information is based on a mixture of museum specimen records, published data and any additional information we could gain. The information should only be taken as a guide.
Data on the ecology and general biology of each valid species are drawn from published works, from museum specimen notes, from personal communications from colleagues and from my own observations and experience. Very little of this information is published. A large proportion of the species have never been observed in their natural habitat and no details are available of even their basic biology. The many species with "nothing known" or with incomplete listings reflect the extremely poor state of our knowledge. Occasionally, a few extra observations are inserted after a descriptor. Entries in this section are to be considered tentative and as a guide only.
A selection of references to the general biological literature of the species and its relatives is given. This list, however, is not complete. Many species have no citation in this section due to the lack of published information other than that given in the secondary reference section.
Barker, G.M. 1999. Naturalised terrestrial Stylommatophora (Mollusca: Gastropoda). Fauna of New Zealand, 38. Lincoln, Canterbury, New Zealand : Manaaki Whenua Press 253 pp.
Barker, G.M. 2001. The Biology of Terrestrial Molluscs. Wallingford, UK : CABI Publishing 558 pp.
Boss, K.J. 1982. Mollusca. pp. 945-1166 in Parker, S.P. (ed.). Synopsis and Classification of Living Organisms. New York : McGraw-Hill Book Company Vol. 1.
Burch, J.B. 1976. Outline of classification of Australian terrestrial molluscs (native and introduced). Journal of the Malacological Society of Australia 3: 127-156
Cox, J.C. 1864. Catalogue of Australian Land Shells. Sydney : John Alex Engel 44 pp.
Cox, J.C. 1868. A Monograph of Australian Land Shells. Sydney : Maddock 111 pp.
Fretter, V. & Peake, J. 1978. Pulmonates. Systematics, Evolution and Ecology. London New York San Francisco : Academic Press Vol. 2A 540 pp.
Fretter, V. & Peake, J. (eds.) 1975. Pulmonates. Functional Anatomy and Physiology. London New York San Francisco : Academic Press Vol. 1 417 pp.
Hubendick, B. 1978. Systematics and comparative morphology of the Basommatophora. pp. 1-47 in Fretter, V. & Peake, J. Pulmonates. Systematics, Evolution and Ecology. London New York San Francisco : Academic Press Vol. 2A 540 pp.
Iredale, T. 1937a. A basic list of the land Mollusca of Australia. The Australian Zoologist 8: 287-333 [12 Mar. 1937]
Iredale, T. 1937b. A basic list of the land Mollusca of Australia. Pt II. The Australian Zoologist 9: 1-39 [12 Nov. 1937]
Iredale, T. 1937c. An annotated check list of the land shells of South and Central Australia. South Australian Naturalist 18: 6-59 [30 Sept. 1937]
Iredale, T. 1938. A basic list of the land Mollusca of Australia. Pt III. The Australian Zoologist 9: 83-124 [Date published 30 Nov. 1938]
Iredale, T. 1939. A review of the land Mollusca of Western Australia. Records of the Western Australian Museum 2(1): 1-88 [published 1 Aug 1939; same text re-published 21 Aug.1939 in Journal of the Royal Society of Western Australia 25:1–88]
Iredale, T. 1940. Guide to the land shells of New South Wales. Australian Naturalist 10: 227-236
Iredale, T. 1941a. A basic list of the land Mollusca of Papua. The Australian Zoologist 10: 51-94
Iredale, T. 1941b. Guide to the land shells of New South Wales. Pt III. Australian Naturalist 11: 1-8
Iredale, T. 1942. Guide to the land shells of New South Wales. Pt IV (to be continued). Australian Naturalist 11: 33-40
Iredale, T. 1943a. A basic list of the freshwater Mollusca of Australia. The Australian Zoologist 10(2): 188-230
Iredale, T. 1943b. Guide to the freshwater shells of New South Wales, Part I. Australian Naturalist 11: 85-95
Iredale, T. 1944. The land Mollusca of Lord Howe Island. The Australian Zoologist 10: 299-334 [10 May 1944]
Iredale, T. 1945. The land Mollusca of Norfolk Island. The Australian Zoologist 11: 46-71
Kershaw, R.C. 1991. Snail and Slug Pests of Tasmania. Launceston : Queen Victoria Museum and Art Gallery 67 pp.
Mead, A.R., Fretter, V. & Peake, J. (eds) 1979. Pulmonates. Economic Malacology. London, New York, San Francisco : Academic Press Vol. 2B 150 pp.
Ponder, W.F. 1998. Conservation. pp. 105-115 in Beesley, P.L., Ross, G.J.B. & Wells, A. (eds). Mollusca: The Southern Synthesis. Fauna of Australia. Melbourne : CSIRO Publishing Vol. 5(Part A) pp. xvi, 1-563.
Smith, B.J. 1992. Non-Marine Mollusca. In, Houston, W.W.K. (ed.). Zoological Catalogue of Australia. Non-marine Mollusca. Canberra : Australian Government Publishing Service Vol. 8 xii 408 pp.
Smith, B.J. & Stanisic, J. 1998. Pulmonata. pp. 1037-1061 in Beesley, P.L., Ross, G.J.B. & Wells, A. (eds). Mollusca: The Southern Synthesis. Fauna of Australia. Melbourne : CSIRO Publishing Vol. 5(Part B) pp. vi-viii, 565-1234.
Solem, A. 1959. Systematics of the land and freshwater Mollusca of the New Hebrides. Fieldiana Zoology 43: 1-234
Solem, A. 1974. The Shell Makers. New York : John Wiley & Sons xii 289 pp.
Solem, A. 1978. Classification of the land Mollusca. pp. 49-97 in Fretter, V. & Peake, J. Pulmonates. Systematics, Evolution and Ecology. London New York San Francisco : Academic Press Vol. 2A 540 pp.
Solem, A. 1991. Distribution and diversity patterns of Australian pupilloid land snails (Mollusca: Pulmonata: Pupillidae, s.l.). Veliger 34(3): 233-252
Stanisic, J. 1990. Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pulmonata) from subtropical rainforests. Memoirs of the Queensland Museum 30: 1-241
Walker, J.C. 1988. Classification of Australian buliniform planorbids (Mollusca: Pulmonata). Records of the Australian Museum 40: 61-89
Zilch, A. 1959. Euthyneura. In, Schindewolf, O.H. (ed.). Handbuch der Paläozoologie. Berlin : Bornträger Vol. 6 (2) xii 834 pp.