Memoirs of the Museum of Victoria 56(2):42 1-430 (1997) 28 February 1997 https://doi.org/10.24199/j.mmv.1997.56.33 CONSERVATION STATUS, THREATS AND HABITAT REQUIREMENTS OF AUSTRALIAN TERRESTRIAL AND FRESHWATER MOLLUSCA

Winston F. Ponder

Division of Invertebrate Zoology, Australian Museum, 6 College Street, Sydney, NSW 2000, Australia

Abstract

Ponder, W.F., 1997. Conservation status, threats and habitat requirements of Australian terrestrial and freshwater molluscs. Memoirs of the Museum of Victoria 56(2): 421-430. World-wide, non-marine molluscs have the largest number of documented extinctions, and of IUCN — listed threatened species, of any major group. Despite this, as with other invertebrate groups, they attract little or no attention from organisations and government departments concerned with conservation. Major impediments to the conservation of non-marine molluscs in Australia, and else- where, include the lack of appropriate legislation, lack of concordance with existing pro- tected areas and little public and political interest. An Action Plan for Australian molluscs is being prepared as part of an Action Plan for non-marine molluscs world-wide being coor- dinated through the Mollusc Specialist Group of IUCN. The plan will bring otherwise inaccessible data to public attention and its recommendations will have application in the conservation of non-marine invertebrates in general. Many Australian non-marine molluscs have very small ranges and are therefore vulner- able to a wide range of threatening processes. The Australian non-marine molluscan fauna currently comprises 1020 named species-group taxa, 80.2% of which are endemic to one State or Territory. 255 species-group taxa (25% of the total fauna) are currently listed by IUCN as threatened in Australia.

Introduction ified. Such areas are easily missed in a strictly habitat-based approach to conservation. Invertebrates comprise a large proportion of Non-marine molluscs are one of the best biodiversity but are very rarely considered in known groups of non-marine invertebrates (see reserve selection and other conservation initiat- Smith, 1992, for a reasonably up to date listing ives in Australia (e.g., Yen and Butcher, 1992; of named taxa). Non-marine molluscs, as a Ponder, 1992a; New, 1995). A major impedi- group, comprise the largest number of recorded ment in utilizing invertebrates is that many extinctions in the last 300 years [284, listed by groups are poorly known both taxonomically IUCN (Groombridge, 1 993)] — this being far and biologically and few people have the exper- more than combined world-wide bird and mam- tise to accurately identify them. In addition, mal extinctions in the same period. To date, unlike the situation with vertebrates and higher mainland Australia has no absolutely confirmed will plants, funding has not generally been made non-marine mollusc extinctions but this available to place important collections on data undoubtedly change, given increasing knowl- of nar- bases. Some of the better known groups, includ- edge about the fauna, the large numbers the rapid rate of habitat ing some insect groups (e.g. butterflies), are row range endemics and will undoubtedly affect a known to contain rare and threatened species. destruction that non-marine However many of these taxa, like many threat- significant number of Australian conservation con- ened vertebrates and higher plants, have con- molluscs. However, mollusc profile, even in current siderable distributional ranges. By way of tinues to have a low invertebrate conservation. For contrast, some invertebrates, including certain discussions on recent issue of Victorian Naturalist groups of insects and spiders show marked example, in a 1 1995) devoted to invertebrate conser- regional endemicity. This is especially true in (vol. 12, molluscs hardly rated a mention. land and freshwater molluscs and some aquatic vation, important aspect of IUCN conservation crustacean groups, many of which have very An 2 is to produce 'action plans' for particu- restricted ranges (often less than 2 km — e.g., strategy concentration lar groups. To date (Nov 1995). there has been Solem, 1 988). Areas which have a IUCN Action Plan developed for of narrow range endemics ('hot-spots') should only one invertebrates, that being for one small group of have a high priority for conservation once ident-

421 422 W.F. PONDER

insects. Swallowtail Butterflies. The Mollusc and the overall total of named terrestrial and Specialist Group (one of the IUCN specialist freshwater taxa 1170. Of these 72.1% of the groups) is well advanced in its produc- named freshwater fauna (176 species-group tion of an Action Plan for all non-marine mol- taxa) and 69.3% of the named terrestrial fauna luscs of the world. A draft Australian plan (642 species group taxa) are restricted to a single has been developed as part of this exercise state or territory. Only 14 (1.5%) of these species and some of the findings are summarized also occur outside Australia. below. For most groups, the work to date has only been descriptive. There are no substantive The Australian non-marine molluscan fauna reviews of most families, and very few phylogen- etic treatments. Only two cladistic analyses of The non-marine fauna of Australia, Lord Howe non-marine groups have been published, both and Norfolk Islands was catalogued by Smith involving Hydrobiidae (Ponder and Clark, (1992) who recognised 190 freshwater and 920 1990; Ponder et al., 1993). Very little biological terrestrial species, including 52 introduced taxa. information is available, even for common taxa On the Australian mainland (including and few detailed distributions have been Tasmania), only 14(1 .4%) of the native land and published. freshwater taxa occur naturally outside Australia, thus the molluscan fauna has a very Terrestrial molluscan fauna high endemicity even by Australian standards (98.6% compared with mammals 82%; birds Of the 23 families recognised in the described 45%; reptiles 89% and vascular plants 85%). The Australian terrestrial mollusc fauna, the land and freshwater molluscan fauna of Lord medium to large-sized Camaenidae is by far the Howe Island (69 terrestrial, 16 freshwater most speciose family, with 408 currently valid, species-group taxa) and Norfolk Islands (68 ter- named species-group taxa comprising 51.3% of restrial and 1 freshwater) is also almost entirely the named terrestrial land snails. The small- endemic to each island. Since Smith (1992) an sized Charopidae are next (129 named, valid additional 1 1 9 species-group taxa have been species-group taxa) with 16.2% of the named named (to Jun 1995) (Table 1). Smith (1992) native terrestrial fauna. Both of these groups, estimated that 30-40% of the non-marine fauna and some other families, contain a large number remained to be described, but a more likely esti- of unnamed taxa (e.g. Solem, 1991; Stanisic, mate is that the total non-marine fauna is 1994). In all, nearly half (48%) of the land mol- around 2000 species. luscs of eastern Australia are unnamed (Stanisic, There are 1 78 species of Australian (excluding 1994). Other large families include Pupillidae Lord Howe and Norfolk Islands) freshwater (41 taxa), Bulimulidae (31 taxa), Punctidae (23 molluscs recorded by Smith (1992) and 743 taxa), Pupinidae (19 taxa) and Helicarionidae species of terrestrial molluscs. Since then (to late (60 taxa). 1995) 119 additional species-group taxa [66 Smith (1992) lists 69 native terrestrial mol- freshwater (representing a 37% increase) and 53 luscs in 10 families on Lord Howe Island. The terrestrial (7% increase)] have been described most diverse are the Helicarionidae (18 species- bringing the totals to 244 and 796 respectively group taxa), Charopidae ( 1 7 species-group taxa),

Table Species-group 1. taxa of Australian freshwater and terrestrial molluscs listed by Smith (1992) with the number of species-group taxa named since added to the totals in brackets. The number in square brackets is the number of species-group taxa restricted to the state or territory, those in the totals columns being the sum of these numbers.

NSW Qld Vic. SA WA NT Tas. Total Aus.

Freshwater 42 56(58) 37(45) 34 31 27 42(98) 178(244) [11] [24] [16] 7 [11] [20] [8] [86] [1 6] Terrestrial 140 234(236) 50 72(99) 226(230) 71(94) 44 743(796) [79] [172] [20] [83] [201] [52] [35] [642] Total 182 290(294) 87(95) 106(133) 258(262) 98(121) 86(141) 901(1020) [90] [196] [36] [94] [221] [60] [121] [818] AUSTRALIAN TERRESTRIAL AND FRESHWATER MOLLUSCS 423

Diplomatinidae (14 species-group taxa) and (26T, 13FW). Victoria (13T, 5FW), South Aus-

Punctidae ( 1 1 species, with several undescribed, tralia (19T, 9FW), Western Australia (63T,

Climo, 1981). The last comprehensive review of 1 FW), Northern Territorv (2 IT), Tasmania (7T. the Lord Howe terrestrial molluscan fauna was 60FW) and Norfolk Island (2T, 1 FW). by Iredale (1944 — see also Stanisic, 1981). The current listing of only 6 terrestrial species from both Lord Howe and Norfolk islands does not reflect the true nature of the situation. For Freshwater molluscan fauna example Placostylus bivaricosus from Lord Australia does not possess a spectacularly Howe is certainly threatened, being restricted to diverse freshwater molluscan fauna, having only a few very small colonies which suffer high levels on the 1 8 species of freshwater mussels (Unionoidea: of rat predation. Data is urgently needed Hyriidae) and very few large-sized gastropods. current distributions and status of terrestrial Apart from the hyriids, corbiculids (2 species) taxa on both islands. and small numbers of viviparids (6 species) and thiarids species), most of the rest of the fauna (9 Extinctions (9 families) are of small size. Amongst these, the hydrobiids stand out as being particularly Many of the recorded extinctions of terrestrial diverse (117 named species-group taxa), many molluscs are those on oceanic islands. There land snails with very restricted distributions (Ponder, 1 994) have been some extinctions of large and many remain undescribed. The next largest on Lord Howe Island, probably mostly due to family, the Planorbidae, has about 40 species. rats (Smithers et al., 1977) and, possibly, pigs, Most families require major revision and more rather than habitat destruction. A large species, new taxa are known in Assimineidae, Bithynii- Epiglypta howinsulae (Cox), is almost certainly dae and Glacidorbidae than are currently extinct, as is a 'form' (or 'subspecies') of Placos- described. Although the Sphaeriidae was rela- tylus bivaricosus (Gaskoin), P. bivaricosus ether- on the tively recently revised (Kuiper, 1983), further idgei (Hedley), both of which occurred part of the island. Habitat work on this group is also needed. well forested southern was For an oceanic island. Lord Howe Island has destruction caused by introduced animals of an exceptional freshwater molluscan fauna. probably responsible for the extinction Apart from one record of an ancylid (Ponder, another 'form' of Placostylus, P. cuniculinsulae, (Smithers et al., 1977; Hut- 1981a), there is a remarkable hydrobiid fauna from Rabbit Island endemic (Ponder, 1982) comprising 15 named species- ton, 1986), as well as other supposed island (see Iredale, group taxa in three genera. Hemistomia minu- land snails from that tissima and another unnamed species live inters- 1944). Norfolk Island, the only recorded fresh- titially in sediments in stream beds. On water mollusc, the hydrobiid Posticobia norfol- kensis (Sykes), is now presumed to be extinct taxa Threatened (Ponder, 1981b). The environmental degra- introduction of feral Although 255 threatened Australian taxa are dation of this island and the extinction of 1 animals which has resulted in the listed in the 1 994 Red List (Groombridge, 993) birds and plants has probably affected more will be added as details of their distri- several snail fauna. Iredale (1945) butions and status become available. Other the terrestrial land extinction of the land snails of the species belong in groups which require taxo- records the Island species) which was com- nomic revision but by far the largest group of nearby Phillip (3 stripped of vegetation by feral animals by unlisted taxa are species which await descrip- pletely are the 1 940s and at least 4 species of land snails tion. Many undescribed, narrow range camae- presumed extinct on Norfolk Island itself nids are known, especially from Western Aus- now (R.V.J. Varman, in litt.). tralia, the Northern Territory and Queensland. are no published records of confirmed Charopids are speciose in eastern Australia and There non-marine molluscs on the Aus- many undescribed species have very limited extinctions of tralian mainland to date. However, this is prob- ranges, especially those on limestone outcrops. in part because there was a general lack of At present the following numbers of threatened ably systematic and/or well documented early col- taxa are listed by IUCN (Groombridge, 1993) lecting of many of the areas now entirely rural for each state and territory (T=terrestrial, and urban. It is probably also a reflection of the FW = freshwater) — NSW (mainland 18T, of the taxonomy of many of the 1FW; Lord Howe Island 4T, 8FW), Queensland poor state 424 W.F. PONDER

groups, particularly the lack of modern compre- 10 sites, 42% were undescribed (Colman and hensive reviews and revisions resulting in poor Burch, 1977). information on the current status of many Limestone areas in particular, because of their species. often highly localised cave and surface faunas, Given the considerable localization and high are particularly vulnerable. In the main, these vulnerability in permanent, mainly lowland habitats have only begun to be systematically

streams, many hydrobiids have undoubtedly collected in the last 1 years by J. Stanisic. Clear- become extinct in the last 200+ years in areas ing of trees from limestone areas for rural activi- where this family is known to be diverse (e.g. ties has probably had a serious impact in some north coast of Tasmania). The detailed work of areas, as have mining activities. Ponder et al. (1993) enables the reporting of at Forty-five introduced taxa (Smith, 1992)

least four suspected extinctions of Tasmanian make up 5. 1 % of the Australian mainland fauna. hydrobiids. Beddomeia tumidafrom Great Lake There are 13 families of terrestrial molluscs and has not been found since the flooding of that lake one freshwater family introduced to Australia, in the 1920s. An unnamed species of Beddomeia the majority of these taxa being found in rural with a distinctive shell morphology from Table and urban areas, mainly in the southern, south- Cape' collected prior to 1900 was not located in eastern and south-western coastal areas and a search of virtually all the streams in this Tasmania (Smith, 1981, 1992; Kershaw, 1991). heavily impacted area in 1989. An undescribed Some helicids (see Smith and Kershaw, 1981; species of Phrantela was collected from the Baker, 1989) and limacid slugs (see Altena and Serpentine River, SW Tasmania prior to the Smith, 1975) are economic pests. Nothing is flooding of this area as part of the new Lake Ped- known about the interactions between native der in 1972. This species has not been found in and introduced taxa. any of the extant streams and rivers in the area. A population attributed to Beddomeia lodderae Habitat conservation by Petterd ( 1 889) from Deep Creek, Duck River, but which probably represents a distinct The belief that habitats conserved for other species, cannot now be located (Ponder et al.. organisms (namely vertebrates and higher 1993: 603), this area having been cleared for plants), or on a representative ecosystem basis, agriculture. will provide effective conservation for invert- Drawdown from extraction of artesian water ebrates is well entrenched. Whereas this may be since the turn of the century has resulted in the effective for many widely distributed species

extinction of many of the artesian (mound) (which are typically at low risk) it is generally springs associated with the Great Artesian Basin inappropriate for many of the taxa of high con- (Habermehl, 1982; Ponder, 1986). Given that servation significance such as narrow range the considerable endemic aquatic invertebrate endemics, including many relict taxa. However, faunas associated with these springs (Ponder, there are some areas where general conservation 1986) were not recorded until the 1980s, there interests and those of molluscan conservation have probably also been extinctions of signifi- coincide, either because of general concerns for cant undocumented faunas. important habitats such as rainforests, or Near extinctions of viviparids have been because of happy accidents, where areas set reported in the Murray River, with the only aside for conservation include areas that are surviving population of one species living in a important for narrow range endemic invert- few irrigation pipelines (Sheldon and Walker, ebrate taxa, typically without this having being 1993a,b). Another species, extinct in the realised (e.g., limestone cave reserves). Murray, is known from a few additional popu- The conservation of rainforests is critical for lations. At the time of writing the viviparid in molluscan conservation. They occupy only the pipelines are in imminent danger of being about 4.5% of Australian forests but contain a exterminated because they pose a nuisance large proportion of the molluscan taxa. Existing through clogging the spray nozzles (K. Walker, rainforests are essentially refugia, occupying pers. eomm.). 2 only about 20,000 km , mainly along the east About two thirds of Australia's rainforests coast of Australia (Webb and Tracey, 1981) with have been cleared in the last 200 years but the a few small areas of monsoonal rainforest first detailed survey for molluscs in east coast patches from Cape York Peninsula in the north rainforests was undertaken in 1975 in mid-east east to the Kimberley in the north west. Much of Queensland. Of the 92 species collected in only the lowland rainforest has been cleared for AUSTRALIAN TERRESTRIAL AND FRESHWATER MOLLUSCS 425

agriculture. Rainforest remnants are still being However, dry forests can be important habitats cleared and are sometimes considered of little or when local topography or geology results in no conservation value if they are of insufficient establishing suitable conditions for refugia. size to maintain large vertebrates. However, Other terrestrial areas of high conservation such areas may be vital for the continuing exist- value arc largely circumscribed by geomorpho- ence of locally endemic invertebrates, small logical features resulting in their being very long- plants and even small vertebrates (e.g., Meave et term suitable habitats. Of particular importance of al., 1991). arc limestone outcrops in which endemicity The tropical rainforests ofNE Queensland are terrestrial species is usually very high. This is a very important habitat for land snails especially true in eastern Queensland and New

1 and in north West- (Stanisic, 1 994) some locations with in excess of South Wales (Stanisic, 994) 40 species of land molluscs. Dry rainforest (vine ern Australia (Solem, 1981a,b, 1984a, 1985, thickets) inland from the coast in Queensland 1988, 1991). When limestone areas occur in have reduced diversity (averaging about 10 rainforests diversity is particularly high (Stan- pro- species) although inland areas of moist refugia isic, 1994). Limestone habitats are often or limestone have higher diversities (Stanisic, tected, at least in part, if they are associated with are not 1994). The subtropical rainforests of south east caves. However, many other outcrops Queensland and New South Wales also contain protected and are often cleared or mined. diverse molluscan faunas (Colman and Burch, Arid zone refugia, particularly deep gorges, associated 1977; Stanisic, 1994). Although generally a can be very important, especially if fauna lower diversity than those further north with limestone. There is a rich camaenid 1984a, 1985, (Stanisic, 1994), areas associated with lime- in the Kimberley (Solem, 1981a,b, (Solem, 1992a,b), stones in the Macleay Valley peak in diversity at 1988, 1991), Flinders Ranges (Solem, 1993), 36 species but in most sites in northern NSW and the gorges of the Red Centre even the coastal areas along the Great Aus- 1 0-20 species is usual (Stanisic, 1 994). There are and species (Solem, few locations in the world in which the land mol- tralian Bight have a few endemic In important review of arid zone lusc diversity exceeds 30 species (Solem, 1 984b). I992a,b). an published data on Monsoonal rainforests occur in small patches refugia (Morton et al., 1995), molluscs were taken into across northern Australia and are mostly poorly terrestrial and aquatic assessing the relative documented for molluscs but the Kimberley account. However, when of particular habitats rainforest patches have been surveyed in some conservation importance vertebrates (ANZECC listed detail (McKenzie et al., 1991). These contain a only threatened considered. rich molluscan fauna (Solem, 1991; Solem and species) were of Australian aquatic habi- McKenzie, 1991) with 115 species, all but 22 The conservation by Lake ( 1 980) and McComb being camaenids. There is an average 12.5 tats is reviewed

1 The only national review of the species (average 3.7 species of camaenids) per and Lake ( 988). conservation of Australian inland aquatic fauna rainforest patch (size < 1 ha to > 20), the mean Michaelis (1986), but few molluscs species range for camaenids was about 20 km in is that of Australian lakes are very diameter and about 64% of camaenids were were noted. Most contain endemic molluscan recorded from only one or two patches. Temper- recent and do not faunas. few Tasmanian lakes, particularly ate rainforests, typically dominated by Nothoja- A Lake Sorell, Lake Crescent and Great Lake are gus, are found in Victoria and Tasmania, contain a few endemics (Fulton, although the great majority (456,000 ha) is in older and including some molluscan taxa. The few Tasmania. These forests also contain a distinc- 1983), seen in large rivers in Australia have a rather low diver- tive fauna, but it is less diverse than that sity of freshwater molluscs when compared with northern rainforests. Asia and North Amer- The remaining 95.5% of Australian forests are some rivers in south east ica. The most conspicuous of the freshwater mainly dry forests, much of this dominated by molluscs, the hyriids, are typically widely dis- eucalypts (about 67.4%). Such forests are gener- molluscs, tributed, although Hyridella xlenelgensis (south ally poor habitats for terrestrial western Victoria) and H. moretonicus (Tas- although wet sclerophyll (mixed eucalypt and faunas. mania) are exceptions (McMichael and Hiscock, rainforest species) can contain diverse 1958). Rainfall can be high in some areas with eucalypt faunas Small streams and, to a lesser extent, local forests but even in these areas molluscan of gener- ground-water seepages and springs, are the are typically sparse, probably because regular fires. major habitats of the majority of hydrobiid ally low water retention in soils and 428 W.F. PONDER

there is virtually no information on the biology conservation. Memoirs of the Queensland and ecology ofmost Australian non-marine mol- Museum 36: 47-53. DEST, 1994. Australia's biodiversity, an overview of luscs. Such basic data gathering is vital to our selected significant components. Biodiversity understanding of how we might best facilitate Series, Paper 2, Biodiversity Unit, Department of the long term survival of many taxa. Coupled the Environment, Sport and Territories: with this, work on the threats to native molluscs Canberra. 87 pp. needs to be encouraged and facilitated. For Evans, J.W., 1942. The food of trout in Tasmania. example, there is virtually no data on the long Salmon and Freshwater Fisheries Commission, term affects of fire, damming, salination and Hobart. pollution on Australian molluscs. Fulton, W.. 1983. Macrobcnthic fauna of Great Lake, Arthurs Lake and Lake Sorell, Tasmania. Aus- tralian Journal of Marine and Freshwater Acknowlegements Research 34: 775-785. Groombridge, B. (ed.), 1993. 1994 IUCN Red List of I thank Ian Loch, Greg Shirley and Alison Miller threatened animals. IUCN, Gland,. Switzerland for comments on the manuscript. The data for and Cambridge. UK. Ivi +286 pp. the 1994 IUCN listings was provided by several Habermehl, M.A.. 1982. Springs in the Great Artesian colleagues, mainly Vince Ressner, Brian Smith, Basin, Australia — their origin and nature. Aus- tralian Mineral Shirley Slack-Smith, John Stanisic, Phil Colman Bureau of Resources, Geology and Geophysics, Report No. 235. and Bronwyn Scott. Hart, B., 1993. A national approach to river manage- ment. Search 24: 125-130. Hill, A., Johnson, M.S. and Merriticld, H., 1983. An References electrophorctic and morphological examination Altena. CO. van R. and Smith. B.J.. 1975. Notes on of Uothriembrion kendricki (Pulmonata: Bulimu- introduced slugs of the families Limacidae and lidae), a new species previously considered eon- Milaeidae in Australia, with two new records. specific with 11. bulla (Menkc). Australian Journal Journal of the Malacologieal Society of Australia of'Zoology'31: 227-242. 3: 63-80. Hutton. I., 1986. Lord Howe Island. Conservation Baker. G.H., 1989. Damage, population dynamics, Press: Canberra. 157 pp. movement and control of pest helicid snails in Invertebrate Advisory Committee, 1994. Interim list native soul hern Australia, pp. 175-185 in Henderson, I. of invertebrates which are threatened or rare (cd.) BCPC Monogr. No. 41. Slugs and Snails in in Tasmania. Parks and Wildlife Service: Hobart. II arid Agriculture. 44 pp. Butcher. R.J., P.E. Clunie, and A.L. Yen, 1994. The Iredale, T„ 1944. The land Mollusca of Lord Howe Victorian Flora and Fauna Guarantee Act: flag- Island. Australian Zoologist 10: 299-334. ship legislation for invertebrate conservation. Iredale, T, 1 945. The land Mollusca of Norfolk Island.

Memoirs of the Queensland Museum 36: 13-19. Australian Zoologist I I: 46-71. Butcher, R. and T.J. Doeg, 1995. Conservation of Jackson, P.D., 1981. Trout introduced into south- freshwater invertebrates. Victorian Naturalist eastern Australia: their interaction with native 112: 15-19. fishes. Victorian Naturalist 98: 18-24.

1 Campbell, I.C. and T.J. Doeg, 1989. Impact of timber Kershaw, R.C., 99 1 . Snails and slugs introduced to or harvesting and production on streams: a review. pests in Tasmania. Queen Victoria Museum and Australian Journal of Marine and Freshwater Art Gallery. Launceston. 67 pp. Research 40: 519-539. Kuipcr. J.G.J., 1983. The Sphaeriidae of Australia. Climo, F.M., 1981. Punctidae — minute land snails. Rasleria 47: 3-52.

P. I 1 in: Recher, H.F. and Ponder, W.F., (eds), Lake, P.S.. 1980. Conservation. Pp. 163-173 in: Lord Howe Island, a summary of current and pro- Williams, W.D. (ed.). An ecological basisfor water jected scientific and environmental activities. resource management. Australian National Uni- Occassional Reports of the Australian Museum versity Press: Canberra.

I. Martin, P., and Lockie. S., 1 993. Environmental infor- Colman, P.H. and Burch, J. B., 1977. Molluscs. Pp. 54- mation for total catchment management: incor- 58 in: Goldstein, W. (ed.), Rainforests. Reprinted porating local knowledge. Australian Geographer 24: 75-85. from Parks and Wildlife. 2( I ). National Parks and Wildlife Service, NSW. McComb. A.J., and Lake, P.S. (eds), 1988. The con- Cooling, M.P. and Boullon, A.J., 1993. Aspects of the servation of Australian wetlands. Surrey Beatty hyporheic zone below the terminus of a South and Sons: Chipping Norton. 196 pp. Australian arid-zone stream. Australian Journal McKenzie, N.L., Johnston, R.B. and Kendrick. P. G. oi Marine and Freshwater Research 44: 411- (eds), 1991. Kimberley rainforests of Australia. 426. Surrcv Beatty and Sons: Chipping Norton, xiv + Daniell. A.. 1994. Genetics and terrestrial mollusc 490 pp. AUSTRALIAN TERRESTRIAL AND FRESHWATER MOLLUSCS 429

McMichael, D.F., and Hiscock, I.D., 1958. A mono- Proceedings ol the Royal Satiety ofTasmania 126: graph of the freshwater mussels (Mollusca: Pcle- 23-28. cypoda) of the Australian region. Australian Ponder, W.F., 1994. Australian freshwater Mollusca: Journal of Marine and Freshwater Research 9: Conservation priorities and indicator species. 372-508. Memoirs ot the Queensland Museum 36: 191- Meave. J., Kellman. M., McDougall, A. and Rosales, 196. J., 1991. Riparian habitats as tropieal forest ref- Ponder. W.F. and Clark, G.A., 1990. A radiation of

ugia. Global Ecology and Biogeographv Letters 1: hydrobiid snails in threatened artesian springs in 69-76. western Queensland. Records of the Australian Michaelis, F.B., 1984. Possible effects of forestry on Museum 42: 301-363. inland waters of Tasmania: a review. Environ- Ponder, W.F.. Clark, G.A., Miller, A.C. and Toluz/i. mental Conservation 11: 331-344. A., 1993. On a major radiation of freshwater Miehaelis, F.B., 1986. Conservation of Australian snails in Tasmania and eastern Victoria: A pre- group aquatic fauna. Pp. 599-6 1 3 in: DeDeckkcr, P. and liminary overview of the Beddomeia Williams, W.D. (eds), Limnology in Australia. (Mollusca: Gastropoda: Hydrobiidae). Invert- CS1RO: Melbourne. ebrate Taxonomy 7: 501-750. Morton, S.R., Short, J. and Barker, R.D.. 1995. Ref- Ponder. W.F.. Colgan. D.J.. Clark. G.A., Miller. A.C ugia for biological diversity in arid and semi-and and Terzis, T., 1994. Microgeographic. genetic Australia. Biodiversity Series, Paper 4, Biodiver- and geographical differentiation of freshwater sity Unit, Department of Environment, Sport and snails — the Hydrobiidae of Wilsons Promon- Territories. 171 pp. tory, Victoria, south eastern Australia. Australian New, T.R., 1995. Introduction to invertebrate conser- Journal of Zoology 42: 557-678. vation biology. Oxford University Press: Oxford, Ponder, W.F., Colgan, D.J., Terzis, T., Clark. S.A. and morphologically xxi + 194 pp. Miller, A.C, 1996. Three new Norris, R.H., Lake, P.S. and Swan, R., 1982. Ecologi- and genetically determined species of hydrobiid northern cal effects of mine effluents on the South Esk gastropods from Dalhousie Springs, new River, North-eastern Tasmania III. Bcnthie South Australia, with the description of a macroin vertebrates. Australian Journal ofMarine genus. Molluscan Research, 17: 49-109. and Freshwater Research 33: 789-809. Ponder, W.F., Eggler, P. and Colgan, D.J., 1995. Gen- (Gastropoda: Pctterd, W.F., 1889. Contributions for a systematic etic differentiation of aquatic snails Aus- catalogue of the aquatic shells of Tasmania. Hydrobiidae) from artesian springs in arid Papers and Proceedings of the Royal Society of tralia. Biological Journal of the Ltnnean Society Tasmania 1888: 60-82. pis 1-4. 56: 553-596. Jenkins, B., 1989. An Pollard, D.A., and Burchmore, J.J., 1986. A possible Ponder, W.F., Hcrshler, R. and hydrobiid snails from scenario for the future of Australia's freshwater endemic radiation of in northern South Australia: their fish fauna. Pp. 615-636 in: DeDcckker, P. and artesian springs and anatomy. Williams, W.D. (eds), Limnology in Australia. taxonomy, physiology, distribution CSIRO: Melbourne. Malacologia 31: 1-140. how environmental Ponder, W.F., 1981a. Freshwater snails. P. 11 in: Ridge, S.. 1995. A study of on the disturbance and Recher. H.F. and Ponder, W.F. (eds). Lord Howe parameters impinge freshwater molluscs in the coastal Island, a summary of current and projected scien- abundance of drainages between Jervis Bay and Sydney. tific and environmental activities. Occassional Honours Thesis, University of Wollongong. Reports of the Australian Museum I. Walker. K.F.. 1993a. Pipelines as a Ponder, W.F.', 1981b. Posticobia norfolkensis (Sykes). Sheldon. F. and snails. Regulated Rivers: an apparently-extinct, freshwater snail from refuge for freshwater Management 8: 295-299. Norfolk Island (Gastropoda: Hydrobiidae). Pro- Research and 7- Sheldon, F. and Walker, K.F.. 1993b. Shell variation ceedings of the Ltnnean Society of NSW 105: 1 in Australian Notopala (Gastropoda: Prosobran- 21. Viviparidae). Journal of the Malacological Ponder, W.F., 1982. Hydrobiidae of Lord Howe- chia: Australia 1993: 59-71. Island (Mollusca: Gastropoda: Prosobracnhia). Society of in B.J., 1 98 1 . Introduced non-marine molluscs Australian Journal of Marine and Freshwater Smith, Victorian Naturalist 98: 24-27. Research 33:89-159. Australia. 1992. Non-marine Mollusca. Zoological Ponder, W.F.. 1986. Mound springs of the Great Smith, B.J., Catalogue of Australia 8: 1-398. Artesian Basin. Pp. 403-420 in: DeDeckkcr. P. B.J., and Kershaw, R.C., 1979. A field guide to and Williams, W.D. (eds). Limnology in Smith, the non-marine molluscs of south eastern Aus- Australia. CSIRO: Melbourne. Australian National University Press: Ponder, W.F., 1992a. Bias and biodiversity. tralia. Canberra, x +285 Australian Zoologist 28: 47-51. pp. Smith, B.J. and Kershaw. R.C., 1 98 1 . Tasmanian land Ponder, W.F., 1992b. A new genus and species of and freshwater molluscs. Fauna of Tasmania aquatic cave-living snail from Tasmania Handbook, 5. 148 (Mollusca: Gastropoda: Hydrobiidae). Papersand pp. 430 W.F. PONDER

Smithers. C, McAlpine, D.. Colman, P. and Gray, M., Johnston, R.B. and Kendrick, P.G., (eds). 1977. Island invertebrates. Lord Howe Island. Kimberley rainforests of Australia. Surrey Beatty Australian Museum: Sydney. and Sons: Chipping Norton..

Solem. A., 1 98 la. Camaenid land snails from Western Stanisic. J., 1981. Land Mollusca. Pp. 12-13 in: and central Australia (Mollusca: Pulmonata: Recher, H.F. and Ponder, W.F. (eds). Lord Howe Camaenidac). II. Taxa from the Kimberley, Island, a summary of current and projected scien- Amplirhagada Iredale. 1933. Records ofthe West- tific and environmental activities. Occassional

ern Australian Museum, Supplement 11: 147- Reports of the Australian Museum 1. 320. Stanisic. J., 1990. Systematica and biogeography of Solem, A., 1 98 1 b. Camaenid land snails from Western eastern Australian Charopidae (Mollusca, Pul- and central Australia (Mollusca: Pulmonata: monata) from subtropical rainforests. Memoirs of Camaenidae). III. Taxa from Ningbing Ranges the Queensland Museum 30: I -24 1. and nearby areas. Records of the Western Aus- Stanisic, J., 1994. The distribution and patterns of tralian Museum, Supplement 17: 427-705. species diversity of land snails in eastern Solem, A., 1 984a. Camaenid land snails from Western Australia. Memoirs of the Queensland Museum and central Australia (Mollusca: Pulmonata: 36: 207-214. Camaenidae). IV. Taxa from the Kimberley, Wes- Stoddart, J. A.. 1983. The accumulation of genetic traltrachia Iredale, 1933. Records of the Western variation in a parthenogenetic snail. Evolution 37:

Australian Museum, Supplement 1 i: 147-320. 546-554. Solem, A., 1984b. A world model of land snail diver- Stoddart, J.A., 1985. Analysis of species lineages of sity and abundance. Pp. 6-22 in: Solem, A. and some Australian thiarids (Thiaridae. Prosobran- Van Bruggen, A.C. (eds). World-wide snails. Bio- chia. Gastropoda) using the evolutionary species geographical studies on non-marine Mollusca. concept. Journal of the Malaeological Society of E.J. Brill/Dr W. Backhuys, Leiden. Australia 7: 7-16. Solem, A., 1985. Camaenid land snails from Western Tilzey, R.D.J., 1980. Introduced fish. Pp. 271-279 in: and central Australia (Mollusca: Pulmonata: Williams, W.D. (ed), An ecological basis for water Camaenidae). V. Remaining Kimberley taxa and resource management. Australian National addenda to the Kimberley. Records ofthe Western University Press: Canberra. Australian Museum, Supplement 20: 707-981. Walker, K.F., 1985. A review of the ecological effects Solem. A., 1988. New camaenid land snails from the of river regulation in Australia. Hvdrobiologia northeast Kimberley, Western Australia. Journal 125: 111-129. the Malaeological of Society of Australia 9: 27- Walker, K.F., Thorns, M.C. and Sheldon, F., 1992. 58. Effects of weirs on the littoral environment of the Solem, A., 1991. Land snails of Kimberley rainforest River Murray, South Australia. Pp. 270-293in: patches and biogeography of all Kimberley land Boon, P.J., Calow. P.A. and Petts G.E. (eds). River snails. Pp. 145-246 in: McKenzie, N.L., John- Conservation and Management. Wiley: Chiches- ston, R.B. and Kendrick, P.G., (edsj, Kimberley ter. rainforests of Australia. Surrey Beatty and Sons: Webb, L.J. and Tracey, J.G.,1981. Rainforests: pat- Chipping Norton. terns and change. Pp. 606-694 in: Keast. A. (ed). Solem, A., 1992a. Camaenid land snails from southern Ecological biogeography of Australia. W. Junk, and eastern South Australia, excluding Kangaroo The Hague.

Island. Part 1 . Systematica, distribution, and vari- Williams, W.D., 1980. Catchment management. Pp. ation. Records the of South Australian Museum. 1 00- in: 1 14 Williams, W.D. (ed). An ecological Monograph Series 2: 1-338. basis for water resource management. Australian Solem, A„ 1 992b. Camaenid land snails from southern National University Press: Canberra. and eastern South Australia, excluding Kangaroo Wilson, J.H., 1966. The food of trout in Great Lake. Island. Part 2. Biogeography and covariation. Australian Societv of Limnology, Newsletter 5- Records of the South Australian Museum, 21-23. Monograph Series 2: 339-395. Woodruff, D.S. and Solem, A., 1990. Allozyme vari- Solem, A., 1993. Camaenid land snails from Western ation in the Australian camaenid land snail and central Australia (Mollusca: Pulmonata: Cristilabrum primum: a prolegomenon for a Camaenidae). VI. Taxa from the Red Centre. molecular phylogeny of an extraordinary radi- Records of the Western Australian Museum, ation in an isolated habitat. The Veliger 33- 129- Supplement 43: 983-1459. 139. Solem. A. and McKenzie, N.L., 1991. The compo- Yen, A.L. and Butcher, R.J., 1992. Practical conser- sition of land snail assemblages in Kimberley vation of non-marine invertebrates. Search 23- rainforests. Pp. 247-263 in: McKenzie, N.L., 103-105.