Zoosyst. Evol. 85 (2) 2009, 199–275 / DOI 10.1002/zoos.200900004

Diversity and disparity ‘down under’: Systematics, biogeography and reproductive modes of the ‘marsupial’ freshwater Thiaridae (Caenogastropoda, Cerithioidea) in Australia

Matthias Glaubrecht*,1, Nora Brinkmann2 and Judith Pppe1

1 Museum fr Naturkunde Berlin, Department of Malacozoology, Invalidenstraße 43, 10115 Berlin, Germany 2 University of Copenhagen, Institute of Biology, Research Group for Comparative Zoology, Universitetsparken 15, 2100 Copenhagen, Denmark

Abstract

Received 11 May 2009 We systematically revise here the Australian taxa of the Thiaridae, a group of freshwater Accepted 15 June 2009 Cerithioidea with pantropical distribution and “marsupial” (i.e. viviparous) reproductive Published 24 September 2009 modes. On this long isolated continent, the naming of several monotypic genera and a plethora of species have clouded both the phylogenetical and biogeographical relation- ships with other thiarids, in particular in Southeast Asia, thus hampering insight into the evolution of Australian taxa and their natural history. Based on own collections during five expeditions to various regions in Australia between 2002 and 2007, the study of rele- vant type material and the comparison with (mostly shell) material from major Australian museum collections, we describe and document here the morphology (of adults and juve- niles) and radulae of all relevant thiarid taxa, discussing the taxonomical implications and nomenclatural consequences. Presenting comprehensive compilations of the occurrences for all Australian thiarid species, we document their geographical distribution (based on over 900 records) with references ranging from continent-wide to drainage-based pat- terns. We morphologically identify a total of eleven distinct species (also corroborated as distinct clades by molecular genetic data, to be reported elsewhere), of which six species are endemic to Australia, viz. “Thiara australis”, Plotiopsis balonnensis, and “Stenomela- nia” denisoniensis with wide distribution and Melasma onca, Sermyla venustula, and Ri- palania queenslandica with more restricted ranges. In contrast, Thiara amarula and Ste- nomelania cf. aspirans as well as Melanoides tuberculata, Plotia scabra, and Sermyla riqueti are widely distributed also outside Australia, in particular in the Malay Archipela- go and the Indo-West Pacific, respectively. The occurrences especially of the latter three species are discussed, concluding on quite distinct historical contingencies. Three thiarids species, viz. Stenomelania cf. aspirans, Sermyla riqueti and Plotia scabra, are recorded here for the first time for Australia. Based on a new taxonomic framework for the Thiar- idae we point out some of the pertinent problems with naming and artificial delineation Key Words of species, revealing why typology and earlier practice of splitting was misleading in case of these truly “Darwinian snails”. We also report on finding two clearly distinct vivipar- endemism ous modes in Australian thiarids, discussing their distribution in certain fluvifaunal pro- drainage division vinces and major drainage systems in concert with these reproductive features of life his- fluvifaunal provinces tory tactics. While the live-bearing T. amarula, S. cf. aspirans, and R. queenslandica, that colonization release veligers (ovo-viviparity), are found to have very restricted occurrences in rivers marine intrusion and streams in the Jardinian province of NE Queensland only, the five more widely dis- invasion tributed Australian endemics Plotiopsis balonnensis and “Stenomelania” denisoniensis as cryptic species well as “Thiara australis”, Sermyla venustula and Melasma onca in the Leichhardtian (ovo-)viviparity province all brood and release shelled juveniles (eu-viviparity). Finally, we hypothesize (eu-)viviparity on the evolutionary history and colonization of Australia by different lineages of these veliger larvae freshwater Cerithioidea.

* Corresponding author, e-mail: [email protected]

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 200 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Introduction “Earlier in the evening I had been lying on a sunny bank & was reflecting on the strange character of the Animals of the country as compared to the rest of the World. A Disbeliever in everything beyond his own reason, might exclaim, ‘Surely two distinct Creators must have been at work’”. (Charles Darwin, January 1836 at Wallerawang, New South Wales)

Australia has an eminently curious and peculiar fauna as being only recent invaders from the north, as it was and flora, distinct in a multitude of ways from that in held by pre-continental drift views e.g. for birds (Mayr other parts of the world, and immediately evident to the 1944), amphibians (Darlington 1965) or other faunal first naturalists. When Charles Darwin visited the area members in particular of inland waters (Williams & Al- of Sydney and the Blue Mountains early in 1836 during len 1987). Also the tropical rainforests dominating his world spanning voyage on board the “Beagle”, northeastern Queensland and New Guinea today were known for its pivotal influence not only on his career long conceived of as unimportant appendage of forests but also on the origin of evolutionary theory, he soon in SE Asia, due to the earlier prevailing palaeobotanical recognized this “strange character” of the fauna, as it view that rather the widespread eucalypt forest of Aus- appears in the above given passage of his “Beagle” tralia was the ancestral habitat, thus also for animal di- diary (Keynes 1988; Nicholas & Nicholas 1989). versification (for discussion see e.g. White 1986, 1994; Long predicted only as “terra australis incognita” the Archer et al. 1991; Flannery 1994; Heinsohn 2008). fifth continent is historically “new” land; however, in Accordingly, it remained the standard scenario that this fact this strange dry land with its weird and wonderful long isolated continent certainly must have been colo- assortment of animals and plants is one of the world’s nized from abroad, in particular by invaders from the most ancient landmasses which harbours one of the Indo-Malay Archipelago, once Australia has been in richest biota with a high number of endemisms. Austra- reach for several taxa from the Oriental region. lia’s fauna and flora was only very slowly discovered The fallacy of such a perception of Australia’s “colo- and studied, albeit its many unique elements – from nial” history became obvious only recently when it was kangaroos and cockatoos, lungfishes and lyrebirds to established by use of molecular phylogeny that, for ex- the platypus – have ever since puzzled biogeographers, ample, the most speciose and widely distributed oscine systematists and other naturalists. Next to endemic and passerines (songbirds, comprising more than half of all unique forms evolution has also produced, for example, extant avian species) apparently originated in the Aus- some native marsupials strikingly similar to Old World tralo-Papuan region as part of (eastern) Gondwana and species, such as the bandicoot and the Tasmanian wolf only subsequently colonized the Eurasian continent resembling placental rabbits and carnivores, respec- (Barker et al. 2004). Instead of Australian passerines tively. Even among the fossil fauna a most spectacular being confamilial Northern Hemisphere derivatives and convergence of anatomical specializations of a xylopha- relatively recent invaders, a phylogenetic reconstruction gus (wood-boring) Australian metatherian from the based on two single-copy nuclear gene sequences sug- Oligo-Miocene to an extant marsupial of Australia as gested multiple waves of emigration from Australia well as to the primate Daubentonia of Madagascar has with dispersal into Eurasia, Africa, and the New World, been reported lately (Beck 2009). Australia’s rich native commencing as early as the Eocene. Further evidence avifauna exhibits similar cases of rampant convergent for Australia actually having been the birthplace of evolution; however, being less conspicuous, these were songbirds as part of a large-scale authochthonous radia- not recognized as such until most recently (see below). tion that far exceeded in scope that of the marsupials Australia’s biota was long not only considered peculiar came from passerine fossils in Early Eocene strata (c. but “primitive”, as it soon became evident that this con- 54.6 mya) at Murgon in southeastern Queensland tinent harbours many apparently ancient floral and faunal (Boles 1995), as well as from other molecular studies elements that only survived there, such as e.g. cycads, on birds resulting in a renaissance of higher level sys- lungfishes, monotremes and the many marsupials. In ad- tematics of this vertebrate clade (Edwards & Boles dition to the ancestry of these lineages an exceptionally 2002). Essentially turning the phylogeny of passerine high degree of endemism with many speciose taxa gave birds and significant parts of Australia’s avifauna up- witness to the long isolation of the continent. On the side down, this new viewpoint not only reverses the other hand, from very early on the Australian fauna was classical scenario of oscine biogeography and the struc- considered, albeit erroneously, “impoverished” and “less- ture of their radiation, but also foreshadows similar perfected”, as even Darwin noted (see Ospovat 1981: evolutionary trajectories for other biotic elements in 219). Thus, many naturalists way into the last century Australia. For example, the crown Corvida among the thought of Australia as a backward land populated by the Australian birds, that according to recent evidence have world’s most inferior creatures, as it has eloquently been their origin in Australia, began to colonize Africa and described by Flannery (1994). the New World via Africa some 35 mya when terranes In concert with these mixed concepts of individuality of Australian origin approached Asian landmasses and inferiority many biotic elements found today in (Jonsson et al. 2008). On the other hand, using molecu- Australia were, nevertheless, conventionally considered lar phylogenetic data from other bird groups that were

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 201 once instrumental in the development of classical spe- that there are “a few remarkable genera which are not ciation and biogeographic theory, Filardi & Moyle found in other parts of the world. These contain but (2005) hypothesized a reversal of the simple one-way, few species for the most part”. He was also one of the downstream flow of colonists, in this case from conti- first to comment on Australia “possessing living repre- nents to island archipelagos in the tropical Pacific. sentatives of extinct fossil forms of Molluscan life”. In Evidently, the Australian fauna is composed of sev- particular, he pointed out that the malacofauna along eral major biogeographical components reflecting quite the northern coasts “is that of the Indian Archipelago, distinct spatial and temporal histories. Recently, this into which the Australian element enters very slightly”. was also shown for Gondwanan elements (as being in- Nearly a century later, and irrespective of the fact dicative of ancient origin and vicariance) versus Asian that our knowledge on the systematics and biogeogra- elements (indicating a more recent origin in and disper- phy of Australia’s molluscan taxa in comparison with sal from Asia), for example, for pierid lepidoptera Asian counterparts hasn’t considerably improved, among the Australian butterflies (Braby et al. 2007). In McMichael (1967: 139) commented on the evolutionary contrast, in case of the agamid lizards, with its 70 spe- relationships of the Australian gastropods of the family cies being one of the most diverse components of the Thiaridae, stating that “they can only be with their reptilian fauna of the continent, an ancient Gondwanan Asian counterparts and they must be regarded as rela- origin was excluded, as phylogenetic data were found to tively recent arrivals in Australia from the north.“ The be more consistent with a more recent, around 30 mya same recent origin and dispersal from the north has old, i.e. Miocene over-water dispersal from SE Asia been assumed earlier for unionid bivalves by McMi- and across the Indonesian Archipelago with subsequent chael & Hiscock (1958), thus founding – albeit not immigration into Australia (Hugall & Lee 2004; Hugall substantiating – what can be called the traditional bio- et al. 2008). For the aquatic fauna of Australia a brief geographical view of the freshwater fauna of Australia. overview was earlier provided by Williams & Allen These authors depicted the northern hemisphere and SE (1987: 189–193), discussing several sources from Asia as the cradle of all limnic bivalves in the Austra- which individual taxa might have originated (ancient lian region, although they also noted the resemblance Gondwana, SE Asia, marine or even terrestrial ances- with the South American freshwater bivalve fauna, only tors, cosmopolitan Northern Hemisphere origin, acci- to relate this to a shared northern origin. dential introductions), but at the same time revealing Undoubtedly, in an arid continent like Australia the many of the open questions that remained until today. freshwater fauna deserves particular attention; however, As theories regarding the origin and evolution of hitherto only few investigations have focussed on its many other Australian faunal elements, the influence of most important and prominent constituent components vicariance versus dispersal events, and the directionality such as, for example, fishes (see e.g. Unmark 2001; and timing of colonizations are still highly controver- Thacker et al. 2007, 2008) or Crustacea and Mollusca. sial, with no confidence or clear consensus yet, it Only most recently Australian freshwater crayfish be- remains the challenge to unravel and test alternative came the focus of several phylogenetic studies, such as, explanations in each individual case, as “they rest more for example, the giant prawn Macrobrachium with on plausible speculation than firm evidence” (Williams some of the biggest but mostly widely distributed spe- & Allen 1987: 189). Sound judgements concerning the cies ocurring also outside Australia (Murphy & Austin ancestry of lineages, the direction and colonization his- 2004a, 2004b; de Bruyn et al. 2004; Carini et al. 2006), tory, as well as the question of endemism is, unfortu- the yabby Cherax destructor from inland Australia nately, currently hampered by insufficient systematic (Nguyen et al. 2004), species of the widespread atyid and biogeographical data for many Australian taxa, in genus Caridina (Page et al. 2007), or the burrowing particular for those groups with potentially high de- freshwater crayfish Engaeus from southeastern Austra- grees of endemisms and low dispersal abilities such as lia as an important area for evolution (Schultz et al. e.g. among the aquatic fauna. While this holds true for 2009). However, questions on the origin and affinity of the prominent vertebrate fauna, it is even more the case these taxa under study in the wider Australian context for the many invertebrates hardly ever studied until to- were rarely addressed and, thus, remained largely un- day. Therefore, to date it has never been tested in gen- clear yet. eral or for a particular taxonomic group whether the Another conspicuous freshwater element is the vivi- limnic molluscs of Australia is either such an “impover- parous snail Notopala of the architaenioglossan gastro- ished appendage” of the Southeast Asian biota, or pods Viviparidae, that were also most recently subject unique to and originated on the Australian continent to few studies, albeit restricted in its focus to the inland since ancient times. waters of Western Queensland (e.g. Carini & Hughes 2006; Carini et al. 2006). To date, though, a systematic and more comprehensive account on viviparid phylo- On the origin of Australia’s freshwater fauna geny, phylogeography and biogeography is lacking. The same is basically true for the most intensively studied Very early on Tenison-Woods (1888) has recognized group of freshwater snails, viz. the Hydrobiidae, which some peculiar features of Australian invertebrate fauna have been intensively monographed, though, with its and noted, albeit without special focus on limnic taxa, various and often locally endemic radiations, for exam-

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 202 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae ple, in the springs of the Great Artesian Basin in South members an enigmatic (and still in detail unresolved) Australia and long term permanent streams in the biogeographical phenomenon becomes apparent. coastal drainages of SE Australia and Tasmania. These snails exhibit not only diverse morphologies on the var- ious taxonomic levels (species and genera), but also re- Thiaridae versus Pachychilidae present many examples of endemisms, partly on a very restricted geographical scale (Ponder et al. 1989, 1993, According to our ongoing phylogenetic analyses of Ceri- 1994, 1995, 1996; Ponder & Clark 1990; Miller et al. thioidea, using morphological data and molecular mark- 1999; Colgan & Ponder 2000; Ponder & Colgan 2002). ers (Glaubrecht 1996, 1999, 2006; Lydeard et al. 2002; While the Hydrobiidae are known to exhibit clear di- Glaubrecht et al. unpubl. data), we found that several versifications with some remarkable endemic radiations lineages have independently colonized limnic habitats in several parts of Australia, the systematic knowledge of worldwide. In contrast to the view of earlier authors (e.g. other limnic gastropod groups is still poor, in particular last revised in Smith 1992; see also Smith 1996) who as to the most prominent and widely distributed Thiari- have traditionally subsumed all relevant Australian taxa dae. According to Smith (1996) thiarids are in terms of under the “melaniids” or Thiaridae, we proposed that number of native genera and species third in Australia two quite distinct, instead of only one of these limnic after the Hydrobiidae (with a total of 12 genera and 114 Cerithioidean lineages are present in Australia. As we species) and the Planorbidae (with 10 genera and 41 spe- have recently shown using morphology and molecular cies). On the other hand, it has been shown of all things genetics, Pseudopotamis Martens, 1894 with two species for the limnic lineages among the otherwise largely mar- endemic to the Torres Strait Islands between Australia ine Cerithioidea that they constitute exceptionally well and New Guinea, is actually a member of the SE Asian suited models for evolutionary biology and historical Pachychilidae (Glaubrecht & Rintelen 2003). Sharing biogeography studies. Limnic Cerithioidean snails have distinct features of the shell, radula and reproductive several times independently colonized freshwater envir- anatomy (i.e. a uterine brood pouch), and supported by a onments, thus repeatedly entering new adaptive zones, molecular phylogeny based on mitochondrial gene frag- and they exhibit an array of evolutionary strategies with ments (COI and 16S), Pseudopotamis was found to be in respect to their reproductive biology (see review, discus- fact the adelphotaxon to species of the pachychilid Tylo- sion and further references in Glaubrecht 1996, 1999, melania endemic to the Indonesian island of Sulawesi 2006). However, a prerequisite for the thorough evalua- (see e.g. Rintelen & Glaubrecht 2005). tion of these different evolutionary as well as biogeogra- Linking vicariance and dispersal events with Earth his- phical aspects are robust, well-corroborated phylogenies tory, and based on recently available detailed tectonic for each of these lineages and a detailed knowledge of mapping of the origin of land masses in the region around the occurrences of their constituent members. Wallace’s line that revealed complex movements of ter- In order to provide these basic systematic and zoogeo- ranes over the past 20–30 mya, we have suggested a novel graphic data, a revision of the Thiaridae is in need. There- biogeographical scenario (Glaubrecht & Rintelen 2003). fore, it is the aim of this paper to present a systematic In order to account for the highly disjunct occurrences of evaluation of the Australian thiarids, their individual oc- the two pachychilid adelphotaxa we anticipated that when currences and distributional ranges on the continent, Australian terranes (of Gondwanan origin) once have ap- based on morphological (including anatomical) investiga- proached Asian land masses (of Laurasian origin) the Ty- tions of all relevant taxa, and to present these data on the lomelania lineage managed to colonize islands in the zoogeography in comparison with congeneric and, if ap- Wallacea, subsequently resulting in some very successful plicable, conspecific thiarids from adjacent areas in the and speciose lacustrine radiations (see for review Glau- Indo-West Pacific studied at the same time (Glaubrecht brecht & Rintelen 2008), whereas Pseudopotamis, with unpubl. data). Ultimately, we want to test whether the its two species restricted to two isolated islands in the Australian continent supports a unique and diverse fresh- Torres Strait, remained relictual at the very northern mar- water (malaco)fauna, here focussing on the thiarids. gin of what is today the Australian continent.

The Thiaridae Troschel, 1857 of Australia Thiaridae Hitherto, confusion existed as to the systematics of Aus- tralian freshwater Cerithioidea, being traditionally treat- Thiaridae, as conceived of now, forms a monophyletic ed as Thiaridae, or even earlier as “Melaniidae”. In the taxon with its constituent species being pantropically following we briefly review the fundamental systematic distributed, including autochthonous occurrences in distinction among these “thiarids” of earlier authors, ac- Central and South America, on some Caribbean Is- cording to our most recent morphological and molecular lands, Africa and in particular Southeast Asia reaching studies on the phylogenetic systematics of these fresh- far out on islands of the western Pacific and into Aus- water lineages, as its lack has previously misled ac- tralia. Its species are found both in lotic and lentic counts on the Australian molluscan fauna. However, freshwater environments, with some taxa (e.g. Stenome- only by way of this crucial systematic distinction be- lania, Sermyla and species conventionally assigned to tween thiarid versus pachychilid affinity of some of its Thiara) tolerating brackish water conditions and occur-

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 203

Figure 1. Shells of Australian Thiaridae, with type species and in alphabetical order as given in the text; species endemic to the Australian continent are listed here as (e). a. Thiara amarula; QLD, Mowbray River (ZMB 106353); b. “Thiara” australis (e); NT, Victoria River (ZMB 106619); c. Melanoides tuberculata; WA, Kununurra, Lilly Lagoon (ZMB 106690); d. Melasma onca (e); NT, Waterhouse River, Stevie’s Hole (ZMB 106681); e. Plotia scabra; NT, Little Roper River (ZMB 106679); f. Plotiopsis balon- nensis (e); NSW, Gwyder River, Bingara (AMS 322678); g. Ripalania queenslandica (e); QLD, Cardwell, Saltwater Creek (syntype BMNH 1879.403-1). h. Sermyla riqueti; QLD, billabong at Norman River (ZMB 107209); j. Sermyla venustula (e); QLD, Bynoe River (ZMB 106712); k. Stenomelania cf. aspirans; QLD, Mowbray River (ZMB 106171); m. “Stenomelania” denisoniensis (e); QLD, Meelele River (106341). Scale = 1 cm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 204 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae ring in the lower courses and estuaries of smaller and McMichael not given 6 genera larger rivers (see for review Glaubrecht 1996). (1967) In contrast to the highly endemic (and endangered) B. J. Smith 10 7 genera and subgenera pachychilid lineage of Pseudopotamis, the Thiaridae (1992) (þ 6 species) (þ 6 incertae sedis are widely distributed in all major regions of continen- species) tal Australia, with the exception of the arid central and most southern parts as well as the island of Tasmania, Although the poor state of knowledge on these and as revealed herein. For a long time the number of spe- other freshwater taxa in Australia has repeatedly been cies and genera of Australian thiarids as well as their stated, starting from Hubendick (1955) and McMichael genealogical relationships remained at best tentative. (1967) to Smith (1992, 1996), with respect to Thiaridae Several taxa have been described and named over the only Stoddart (1983, 1985) presented additional data. last two centuries and rich material deposited under His biochemical approach, though, was restricted to the various and varying specific names in museum collec- question of species delineation in the two species he tions, in particular in the major repositories in Sydney, identified as “Thiara”[=Plotiopsis] balonnensis (Con- Brisbane, Darwin and Perth. However, since E. A. rad, 1850) and “Thiara”[=Stenomelania] denisoniensis Smith’s (1882) survey of Australian freshwater shells (Brot, 1877). Consequently, since no comparative or systematic work on these particularly interesting ele- comprehensive attempt has been made to survey the ments of the malacofauna has for over a century never available material of Australian thiarids with the aim of gone beyond the point of compiling faunal lists with a systematic evaluation of these taxa, it is to date still even more names, but only few synonymies suggested. true what McMichael (1967) once stated: “Unfortu- There are essentially three studies to be mentioned that nately no one has yet correlated the Australian genera have dealt in some more detail with the taxonomy of listed by Iredale (based on shell characters) with those thiarids in the past, among which is first of all the literally of Morrison [1954]”; note, however, that the latter ac- “basic list of freshwater Mollusca of Australia” compiled count is now also long outdated for several reasons (see by Iredale (1943). He essentially presented a list of names Houbrick 1988; Glaubrecht 1996, 1999, 2006). and references, but his suggestions of new taxonomic Research on thiarids is further hampered by their in- names for Australian thiarids implied the evolutionary in- credible variable shells that are also known from other dependence and own biological identity of these particu- freshwater Cerithioidea which all exhibit large phenoty- lar taxa, albeit then without any thorough evaluation. Irre- pic plasticity. This conchological variability might lead spective of the title of his contribution, also McMichael to overestimating the number of evolutionary meaning- (1967) has not presented more than a review of the then ful units, i.e. biological species, as discussed in general accumulated systematic knowledge, also based essen- e.g. in Glaubrecht (2004, 2009) and shown specifically tially on non-Australian Thiaridae. The most recent cata- in the case of other lineages of limnic Cerithioidea, such logue of freshwater taxa with some taxonomic decisions as Melanopsidae (Glaubrecht 1993, 1996) or Pachychili- has been thankfully compiled by B. J. Smith (1992). The dae (Khler & Glaubrecht 2001, 2003, 2006; Glau- following list gives the number of thiarid genera and spe- brecht & Khler 2004). In addition to the difficult cies as differentiated in these above mentioned accounts assessment of large individual, ecological and/or geo- (note that the later all included the pachychilid genus graphical variability, an isolated treatment of only the Pseudopotamis (see above) with its two species unknown Australian thiarids or in particular of singular specimens only to E. A. Smith (1882) at that time): from widely separated localities on this vast continent is always in danger of misinterpreting taxonomic diversity E. A. Smith 12 species 1 genus: “Melania” by underestimating conchological disparity, resulting in (1882) Lamarck, 1799 systematic-taxonomic uncertainties and unresolved phy- [= Thiara Rding, 1798] logenetic affinities. Thus poorly known, the Australian Iredale 16 species 6 genera (with 2 genera thiarids would be of only limited use for biogeographi- (1943) and 4 species new) cal evaluations along the lines given above.

" Figure 2. Characteristic habitats where thiarid species are found (often syntopically) in the various freshwater bodies of Australia. a. Daly River (NT), at causeway crossing (Oct. 2007); with occurrence of “Thiara” australis, Melasma onca and “Stenomelania” denisoniensis; b. Middle course of the Roper River (NT), at Jalmurark (Oct. 2007); with occurrence of “T.” australis, M. onca and Sermyla venustula; c. Flooded area of the upper Roper drainage, Little Roper River (NT), at Homestead Road crossing (June 2004); with occurrence of “T.” australis, M. onca, Sermyla venustula, and “Stenomelania” denisoniensis (note that also Plotia scabra is reported from here); d. Main course of Salt Creek (NT), a tributary of the Roper River, at crossing of Roper Highway (June 2004); with occurrences of “T.” australis, Sermyla venustula, and “Stenomelania” denisoniensis (note that also Plotia scabra is reported from here); e. Dry parts of river bed of Salt Creek (NT), with isolated pools (June 2004); f. Cox River, draining into the Gulf of Carpentaria (NT section; Oct. 2007); with occurrence of “T.” australis and “Stenomelania” denisoniensis; g. Stagnant pools of the upper Mowbray River (QLD), draining into the Coral Sea (May 2002); habitat of Thiara amarula and Stenomelania cf. aspirans; h. Lower course of the North Johnstone River (QLD), at highway crossing (May 2002); with occurrences of Thiara amarula and Ripalania queenslandica (note that also Plotia scabra is reported from here). [All photographs by M. Glaubrecht]

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 205

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 206 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Based on the data presented in this paper, we here tions for biogeographical studies of its freshwater fau- suggest to differentiate among the Australian Thiaridae na, as Unmack (2001) pointed out for fishes. Much 8 genera now (also corroborated by molecular genetic more than vagil terrestrial animals, limnic organisms data, to be reported elsewhere), with a total of 11 spe- are immediately dependent on their habitat and are thus cies distinguishable essentially on the basis of their directly prone to any alterations of this environment. shell morphology. They are depicted by representative Dispersal and range expansions are in general restricted shells in Figure 1 for comparison, and here stringently to rare events like change of river courses, temporary given with their valid generic and specific names as hydrological connections (e.g. through flooding) or evaluated below in the paper. Of these 11 species more more permanent relationships across drainage limits than half of them (n = 6 species) are endemic to Aus- (such as river capture), as well as global changes such tralia only, viz. “Thiara australis”, Plotiopsis balonnen- as sea-level fluctations or tectonics resulting in drifting sis and “Stenomelania” denisoniensis with wider distri- continents and terranes. In light of these drastic bution as well as with more restricted ranges Melasma changes the elimination of individual taxa often seems onca, Sermyla venustula and Ripalania queenslandica. a more likely event than the survival of species and the In contrast to these endemics, Thiara amarula and Ste- colonization of new habitats. In the evolution of the nomelania cf. aspirans as well as Melanoides tubercu- Australian inland water fauna in particular several en- lata, Plotia scabra, and Sermyla riqueti are widely dis- vironmental factors have been influential, among which tributed also outside Australia, in particular in the feature most prominently (i) the continent’s geological Malay Archipelago and the Indo-West Pacific, respec- history with its long geographical isolation, (ii) its phy- tively. Interestingly, these latter five species all have siography dominated by its flat topography (with a either restricted or very patchy occurrences in Austra- post-Oligocene uplift of less than 300 meters and a late lia, some being non-authochthonous, as is argued here Cenozoic relief not much greater than today), and (see Discussion). Three of the taxa, viz. Stenomelania (iii) its past climate with increasing overall aridity (Wil- cf. aspirans, Sermyla riqueti and Plotia scabra, are re- liams & Allen 1987: 184–186). corded here for the first time. To discuss this in concert with the most impressive While only two of the thiarids, viz. Plotiopsis balon- and instructive geological and climatic history of the nensis and “Stenomelania” denisoniensis, are widely Australian continent, as illustrated and highlighted for distributed and occurr in many streams and rivers or plants (e.g. by White 1986, 1994; Markgraf et al. 1995; other water-bodies throughout Australia the highest spe- Martin 1998) or examplified by the mammal fauna (Ar- cies diversity is found in the coastal rivers and inland cher et al. 1991), is way beyond the scope of the pre- streams and creeks of the wet-dry tropical northern sent account. Nevertheless, it is important to note here parts of Australia, where some taxa are endemic. In that rifting of Gondwana and subsequent drifting of particular interesting here are the two major drainage Australia with its correlated climatic changes over the systems of the Daly and Roper Rivers. Some of the ty- past 35 million years since the Miocene transformed it pical thiarid habitats of this region are depicted in Fig- from a wet, green continent to one that is largely dry ure 2, in addition to habitats along the Gulf of Carpen- and brown. This not only resulted in, for example, the taria and the eastern coast of Queensland. retreat, contraction and fragmentation of rainforests re- placed by eucalytus-acacia dominated vegetation and by semi-arid to arid conditions in regions comprising Zoogeography and hydrology of Australia today about 70 % of the continents surface area (e.g. The present distribution of living organisms over the Martin 1978; Keast 1981; White 1994; Hesse et al. world’s surface is the result of two closely interwoven 2004). The dramatic geographic and climatic chance processes: the evolution of organisms and the evolution also has, of course, played a major role in the diversifi- of their habitats. In reconstructing the history of taxa and cation of Australia’s biota, as the increasing aridifica- areas, historical biogeography aims at synthesizing sys- tion interacted with evolutionary processes such as tematic and geological patterns (Glaubrecht 2000). How- adaptation and specialization, as well as speciation and ever, to date these efforts have been primarily restricted radiation. On the other hand, it has resulted in extinc- to a few organismal groups, such as e.g. birds, mammals, tions or at least the lack of particular taxa, as it has fishes, butterflies (e.g. Unmack 2001; Beheregaray 2008; been noted, for example, for the ‘primary’ freshwater Glaubrecht 2009). Although long largely neglected in fishes in Australia (see Bishop & Forbes 1991; Unmack this context, limnic gastropods – with their restricted 2001). As a consequence this continent has been con- habitat preference and high habitat fidelity, limited dis- sidered to possess only an impoverished freshwater fish persal faculty in concert often with their ancestry (and fauna, as compared to other large continental areas, sometimes even rich palaeontological record) – have the composed essentially of ‘secondary’ freshwater fishes potential to serve as ideal and important models for the with close affinities to marine taxa. As pointed out study of historical biogeography and phylogeography, as above, despite the fact of the presence of apparently an- argued e.g. in Glaubrecht (1996, 2000). cient lungfishes (Neoceratodus forsteri) in Queensland At the same time, the geological ancestry as well as and of the presumably relictual salamanderfish (Lepido- aridity of Australia provides almost perfect precondi- galaxias salamandroides) of Gondwanan (or even Pan-

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 207 gaean) ancestry, most other Australian fishes are gener- ble whether they actually represent natural objects or ally assumed to be of relatively recent origin and distri- just artificial constructs (Mackey et al. 2008). In the bution, with about 70 % of the inland species having course of biogeographical studies in Australia several strong affinities with tropical Indo-Pacific marine attempts have been made to regionalize this vast conti- fishes (Whitley 1959; Merrick & Schmida 1984; Wil- nent. Unfortunately, most of the published studies have liams & Allen 1987; Unmack 2001). referred separately to either different taxa (terrestrial, Most relevant here in context of the freshwater gastro- aquatic and marine) and/or to differently delineated re- pods of Australia we can assume that the general topo- gions of Australia on the basis of the composition and graphy and even drainage systems were more or less si- distribution of the organisms under study. The impor- milar to those of the present since the Miocene, while the tance, but also the problems for reconciling these ap- climate and precipitation pattern underwent dramatic proaches to biogeographical regionalization has only re- change, not only in the course of Australia’s northward cently been described and discussed by Mackey et al. drift (from the landmass being situated mainly between (2008). Here no attempt will be made to review the latitudes 40 and 70 S in the middle Paleocene as com- various arrangements for the freshwater fauna. How- pared to its present position between 10 and 40 S), but ever, it seems necessary for the characterization of the also due to oscillations between cool, dry glacial periods geographical ranges of the thiarid snail fauna and for and warmer, moister interglacial periods since the Early comparative purposes to briefly look into some of the Pliocene (around 4 mya) that continued into the late most important regionalizations suggested on the one Pleistocene and Holocene (Williams & Allen 1987; see hand for the Australian biota in general and for differ- also Haynes et al. 1991). These climatic events forced ent drainage systems on the other hand. the aquatic biota to adopt to the increasingly more arid For example, Iredale & Whitley (1938) have divided conditions, i.e. to seasonal and high unpredictable preci- the continent and New Guinea in nine zoogeographical pitation with often strong monsoonal rainfall, thus to the regions, called ‘fluvifaunulae’, with distinct occur- ephemeral nature of many water bodies with high sali- rences of mammals, birds, fishes and molluscs. Later, nities in temporally standing waters and marked hydrolo- Whitley (1947) modified the limits of these original gical fluctuations of many rivers and streams. ‘fluvifaunulae’, naming region like Greyian, Leichhard- At the same time the geographical and climatic tian and Jardinian in honor of early explorers and natur- chances brought about other significant, albeit less ob- alists associated with them (see Figure 3a). This system vious evolutionary consequences, such as fragmentation has been adopted and modified by McMichael & His- of species’ ranges with separation of populations poten- cock (1958) for the Australian freshwater bivalves, se- tially leading to speciation and/or fusion and secondary perating nine fluvifaunal provinces for the Australian contact resulting in hybridization or not. For example, continent and Tasmania plus two for New Guinea (see as Williams & Allen (1987: 194) pointed out, the flucta- Figure 3c). Lake (1971) employed a regionalization tions in climate and consequently precipitation patterns based on a total of 15 separate drainage systems, of caused gross changes in the distribution of areas where which four drain along the wet tropical northern conti- predictable filled or flowing waters have occurred, thus nental margin into the sea (Figure 3b). Based on the bringing about geographical separation of populations biogeographical study of fishes, Unmack (2001) sug- as the usual prerequisite for speciation. Only briefly gested again a slighty modified zonal structure of hy- mentioning potential evolutionary pathways, they as- drogeographical provinces, using a mixture of geogra- sumed that speciation seems likely to have occurred phical and river system names (Figure 3d). Using only in those water-bodies which predictably (regularly) regional occurrences, he described a ‘line of ende- contained water. However, to date the details of these mism’, dividing Australia into the northeastern region historical processes and their evolutionary consequences with lower frequency of endemic fishes (along the since the mid-Miocene remain to be studied for most of Northern and Eastern Province) and the remaining part Australian’s less conspicuous fauna, in particular for the of the continent to the southwest with many ende- aquatic invertebrates such as freshwater molluscs in misms. It remains to be seen whether this regionaliza- general and Australian Thiaridae in particular. tion is also reflected in the distribution of other fresh- water organisms, such as among the molluscs the thiarids. Although covering but a smaller part of Aus- Regionalization of Australia: fluvifaunal provinces tralia, the volume of water deluge (estimated at 350 cu- and major drainages bic kilometer per year) from these fluvial areas along the northern margin of the continent comprise 65 %. Instrumental in faunal studies are diverse concepts of Most of the rivers in the Leichhardtian province are major regional subdivisions. However, as there is no considered seasonal, i.e. they are often not permanently agreed theoretical basis or general correspondence of running, with their water regime thus being highly in- the various approaches, the concept of biogeographical fluenced by the monsoonal rainfalls and frequently re- regions on whatever scale remains vexed. Although sulting in flooding of extensive areas. these regions are generated to reflect a range of biolo- Many zoogeographical accounts on Australia refer to gical, physical and ecological phenomena, it is debata- the most widely used subdivision of surface hydrologi-

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 208 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 3. Biogeographical regionalization for the Australian continent. a. Faunal re- gions (= fluvifaunulae), suggested by Whit- ley (1947); b. Drainage systems, according to Lake (1971); c. Biogeographical pro- vinces for the freshwater fish fauna, after Unmack (2001); d. Fluvifaunal provinces of Australia and New Guinea, after McMi- chael & Hiscock (1858); e. Major drainage divisions of Australia, modified from Wil- liams & Allen (1987). a–b, modified from Bishop & Forbes (1991).

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 209 cal units based on the continent’s major drainage sys- ious collections as cited and with details given under the individual tems as given e.g. in Williams & Allen (1987); see Fig- species. Types were on loan or studied from the following collections: ure 3e. As is evident from the comparison of the sys- Academy of Natural Sciences Philadelphia (ANSP), British Museum of Natural History (BMNH), Muse d’Histoire Naturelle, Genve tems compiled now in Figure 3 various names in this (MHNG), National Museum of Natural History, Smithsonian Institu- context are used redundantly, as e.g. the Indian Ocean tion, Washington, D.C. (USNM), Museum fr Naturkunde, Berlin (or Pilbara) division corresponds to the Greyian pro- (ZMB) and the Zoologisk Museum University Copenhagen (ZMUC). vince or fluvifaunulae, the Southwest Coast to the Vla- It should be noted that all available soft body material of Australian minghian and the Sturtian fluvifaunal province largely thiarids is subject to ongoing molecular genetic studies by the authors match the Lake Eyre division. On the other hand, on the phylogeny of Thiaridae. The results of these studies will soon be others combine certain drainages, as e.g. the Timor Sea published elsewhere, but have been consulted already for the present documentation to the extend that they are in accordance with the (shell) (including the Kimberley region) and Gulf division morphology based evidence as discussed here. This is in particular true within the Leichhardtian province. for the more cryptic species identities of the two (potentially invasive) Therefore, for comparative purpose we will in the fol- species Melanoides tuberculata and Plotia scabra, which are to be com- lowing refer to these fluvifaunal provinces as used earlier pared to and verified against “Stenomelania” denisoniensis, Plotiopsis by McMichael & Hiscock (1958) in combination with balonnensis and “Thiara” australis, respectively. those of the major drainage division as given in Williams & Allen (1987); see Figures 3d–e. By this way of inte- gration we are able to differentiate, for example, within Methods the Leichhardtian fluvifaunal province occurrences re- This work only includes material actually seen and clearly identified stricted to either the Timor Sea or the Gulf of Carpentar- to the species by the first and second author. A first analysis of the ia drainage, or refer within the Northeast Coast division material was done for an unpublished Master Thesis at the Humboldt separately to the Jardinian and Krefftian provinces. In University in Berlin by N. B. in 2005–2006. It is supplemented here the course of our study on thiarid biogeography these dif- by the 2007 field collection by the first two authors, and a collection ferentiated references proved to be important, as both the of thiarids in Arnhem Land also in 2007 plus some additional sam- provinces as well as the drainage system divisions corre- ples from other collectors. For each species the type specimens as well as representative specimens were photographed using a digital sponding to major rivers were found most relevant. It will camera (Canon EOS 350D), with pictures subsequently assembled also allow to place the present study in the wider frame- with Adobe Photoshop CS2 (version 9.0 for Windows). work of investigations into the evolution and biogeogra- Distribution maps. All known localities of material from the con- phy of Australia’s freshwater fauna. sulted collections or collected thiarids from species-specifically iden- tified samples were comprehensively compiled for the material exam- ined sections, comprising a total of over 900 individual records, and Materials and methods the distribution maps given under each species. In case of earlier mu- seum samples usually no geographic coordinates were available, re- Material sulting in time consuming “georeferencing” of these localities in question, as we attempted to cover the species’ range as completely This study is based essentially on samples collected during five field as possible. During this work, started a decade ago at the AMS by trips to Queensland (May–June 2002, October 2007), the Northern Ter- M. G., we used over the years various sources for finding localities ritory (June–July 2004, September–October 2005, October 2007), and assembling distribution maps, e.g. “Meyers Grosser Weltatlas” Western Australia (July 2004) and Central Australia (October 2005) by (Brockhaus Bibliographisches Institut, Mannheim, 1997; 5th edition; the first two authors, partially with help of Thomas von Rintelen (2002, scale 1 : 800,000) and “Encarta” as implemented under Windows, 2004) and Vince Kessner (2004, 2005). In addition, all relevant museum most recently Google Earth, Google Maps and different gazeteers, material, mostly thiarid shells, was studied in particular from the AMS, and in particular the detailed maps for parts of the Australian conti- NTM, WAM and QM collections, supplemented by the private collec- nent, e.g. the Hema state maps (for the “Red Centre”, “Northern Ter- tion of Vince Kessner (Adelaide River) and samples from other mu- ritory”, “Western Australia”, and “Queensland” (scale 1 : 2,500,000; seums (as listed below under the species). For this purpose, in particular various editions) as well as the map “Australia” (scale 1 : 2,800,000; the rich shell (and in addition also more recent wet) material in the Karto & Graphik Verlagsgesellschaft, Frankfurt a. M., 1995) with its AMS was sorted and identified to the species in 1996, with additions locality index. covering collections in the subsequent years in particular done by Win- Mapped localities were finally transferred on a dot-by-dot basis to a ston Ponder. Additional samples preserved in ethanol, available also for digitally reduced version of the master drainage pattern map of Australia; our accompanying molecular genetics study, were collected in particu- thus, data in each distribution map given herein correspond to the local- lar during their Arnhem Land expedition by Winston Ponder and Vince ities as listed under each material examined section. In order to avoid Kessner in 2007, next to few samples from other areas (see under ac- redundancies in these lists locality information has been summarized by knowledgments), while older material stored routineously in formalin the first author in cases of repeated collections done at the same spots (all earlier collections of the AMS) proved to be unsuitable. and represented by the same material in more than one museum collec- Living snails were fixed in 70–96 % ethanol in the field, with some tion (which is often the case given the limited accessibility to freshwater shells from each sample cracked the same day in the field to allow the in remote areas of Australia). Were applicable, we gave priority to the preservative to penetrate the soft bodies. Ethanol was changed once or lat./long. references as determined for our own ZMB samples. Note that twice in the field, transpored to and shipped mostly by the Museum & for readability of the maps figured here in cases of multiple samples ana- Art Gallery of the Northern Territory in Darwin to the ZMB. All sam- lysed from very closely adjacent localities only one or few dots are shown ples were finally stored in 96 % ethanol, with voucher material depos- to represent occurrences; thus out of the total 900 records about 270 are ited now in the NTM in Darwin and most complete at the ZMB. shown in the maps (i.e. a reduction ratio of 1 : 3 on average). Identifications are based, in addition to the original accounts of the Few questionable records from museum collections, with obviously types, on comparative studies of the type material itself from the var- erroneous label information (such as e.g. “N Gulf of Carpentaria,

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 210 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

1258.50 S 139300 E” given for a sample of Plotiopsis balonnensis, Embryos and juvenile shells. Ontogenetic stages were removed from which lays way out in the sea) were excluded from the biogeographi- the brood pouch, counted according to selected size classes, with cal analysis. Some other samples were also excluded when identifica- some mounted on stubs and spattered with gold-palladium for SEM tions could not be verified so far, as it was in particular the case in studies and documentation. Some stages were treated with Hexa- P.balonnensis being easily misidentified as “Thiara australis” (see methyldisilazan following the procedure described by Nation (1983) for more details under that species). before SEM study. For the fluvifaunal provinces of Australia we refer to the names Early ontogenetic stages in caenogastropods are characterized by used in McMichael & Hiscock (1958), in combination with those of the initial whorl of the shell lacking growth lines. Compared to mar- the major drainage systems as given in Williams & Allen (1987); see ine taxa, freshwater gastropods have a less distinct sculptural transi- above under hydrology and Figure 3. tion from the primary (or embryonic) shell without growth lines and Ecology. Only rarely ecological remarks were found for samples in the secondary (= larval) shell to the adult or tertiary shell (= teleo- museum collections; in those few cases these informations are in- conch). Due to the abbreviation or loss of early ontogenetic stages cluded with the material listed for each species. For the ecology we particularly in viviparous freshwater Cerithioidea a distinct transition here restrict ourselves to few general notes, but refrain from a de- is lacking; see Glaubrecht (1996). Therefore, the term ‘protoconch’ as tailed ecological description of Australian thiarids, as this will be sub- generally applied to the first two ontogenetical stages is not compar- ject to a separate account elsewhere. able to the conditions found among viviparous gastropods. Conse- quently, here the more general term ‘embryonic shell’ is prefered for Anatomy. Examinations and dissections were made with the aid of a all shelled stages in the brood pouch. It may differ very little from stereo microscop (Leica Wild MZ 9.5) and documented by camera luci- the shell formed by the crawling benthic animal after hatching which da drawings at the ZMB. Radula as well as embryos and juveniles in the is called the teleoconch. females’ brood pouch were removed, at the same time a piece of foot Measurements of the juvenile shells (see Figure 4b) follow standard tissue was taken for molecular genetic study (to be reported elsewhere). methods and terminology as described in Glaubrecht (1996). Height, Pictures of juvenile shells, radulae and opercula were made with the width and maximum diameter at one whorl were obtained from the ZMB Leo 1450 VP and CamScan DV 4 scanning electronic microscope SEM integrated software and/or scanning electron micrographs (preci- at 10 KVafter probes being spattered with gold-palladium. sion to 1 mm), with the h/w ration calculated from this. Radula. Radulae were extracted during dissection, processed using Proteinase K as described by Holznagel (1998) and cleaned ultrasoni- cally for SEM study. Number of rows and the absolute length of the Museum codens radula (with a precision to 0.1 mm) were taken under the SEM using the integrated software. The number of cusps of radula teeth are re- AMS Australian Museum, Sydney ported as follows: rachidian (or central tooth, number of cusps on the ANSP Academy of Natural Sciences, Philadelphia left side/median cusp/cusps on the right side; lateral teeth: inner BMNH The Natural History Museum, London (formerly British cusps/pronounced median cusp/outer cusps; marginal teeth: number of Museum Natural History) cusps on inner plus on outer tooth (see Glaubrecht 1996). CAS California Academy of Sciences, San Francisco Adult shell morphometry. Standard shell measurements have been essen- MCZ Museum of Comparative Zoology, Harvard University, tially taken from the dry material collections plus additional wet samples Cambridge, Mass. after a first species assignment. The following biometrical parameters of MHNG Muse d’Histoire Naturelle, Genve the adult shells were studied for a total of about 1600 individuals, using NHMB Naturhistorisches Museum, Basel an electronic calliper (precision 0.1 mm); see Figure 4a: NTM Northern Territory Museum, Darwin QM Queensland Museum, Brisbane height of shell, width of shell, length of aperture, width of aperture, SMF Senckenbergmuseum, Frankfurt am Main height of last body whorl, height of penultimate three whorls, number USNM National Museum of Natural History, Smithsonian of whorls. Institution, Washington, D.C. Statistical analyses. Analyses of shell and radula parameters as well VK private collection of Vince Kessner, Adelaide River as some of their ratios were done using the SPSS for Windows pack- WAM Western Australian Museum, Perth age (version 11.0). We are not documenting here the results for all ZMB Museum fr Naturkunde, Berlin parameters taken, as only few were found to be informative and rele- (formerly Zoologisches Museum Berlin) vant in the present context. ZMUC Zoologisk Museum University, Copenhagen

Figure 4. Shell measurements. a. Shell parameters of the adult shell. Abbrevia- tions: B – width of shell; BW – height of last body whorl; H – height of shell; L3W – height of penultimate three whorls; LA – length of aperture; WA – width of aperture. b. Shell parameters of the juvenile shell. Abbreviations: h – height; w – width, d – maximum diameter at one whorl.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 211

Abbreviations of Australian Federal States from other nomenclatural acts, in particular from for- AUS Australia mally new naming of taxa. Instead of introducing new WA Western Australia ones, we have whereever possible in the following NT Northern Territory made use of generic and species names already avail- QLD Queensland able from the literature. Thus, for example, we consist- NSW New South Wales ently use throughout the present paper the two generic SA South Australia allocations of “Thiara” (in case of australis) and “Ste- nomelania” (in case of denisoniensis) in quotation marks only, in order to indicate that our results denote Results a phylogenetic placement other than with these two par- ticular lineages traditionally used and characterized by Part I: Systematic account the type species amarula and aspirans, respectively. We have transferred and allocated species to genera for In the absence yet of a robust systematic phylogeny of which names are available in those cases when our phy- Thiaridae including the Australian taxa and based on logenetic, morphological and geographical data, ideally combined molecular analyses of mitochondrial as well as in combination, provide sufficient and consistent evi- nuclear genetic information (Glaubrecht et al. unpubl. dence for the existence of such an independent lineage, data), we here use primarily morphological features to as is the case, in addition to the above mentioned (yet characterize individual taxa, in combination with the geo- “unnamed”) two lineages, in Melasma onca, Plotia sca- graphical information assembled. The resulting systema- bra, Plotiopsis balonnensis, and in Sermyla venustula. tization suggested here, however, already takes into ac- These are here presented in combinations already sug- count our ongoing effort of molecular analyses, to the gested and/or used by earlier authors (see e.g. extend that the findings of both approaches are consistent; B. J. Smith 1992), with the notable exception of only in cases of remaining conflict we clearly have notified the latter species. these. Other than by forestalling some of the findings of our phylogenetic studies (to be later presented elsewhere) Note on the morphological descriptions. For the follow- it would not be possible to describe the results of the pre- ing characterization of the genus and species of Austra- sent morphological investigations in a consistent and con- lian Thiaridae only the most distinct features will be giv- sequent way; instead we would fall behind the knowledge en, with diagnosis and description being as brief as already accumulated, that way perpetuating again out- possible, since the figures depicting shell variability bet- dated taxonomic concepts for the Australian thiarids. ter illustrate relevant shell features than any verbal at- In the systematic part we have arranged the taxa tempts. Also, a detailed anatomical description of the starting with the type genus of the family Thiaridae, Thiaridae is beyond the purpose and scope of the present viz. Thiara Rding, 1798, followed by its type species. paper. As most features studied so far are common in all It then presents subsequent taxa in the same manner, thiarids they proved to be hardly of any use for the differ- with both genera and their type species and constituent species in alphabethical order (see also Figure 1). It ential description of individual Australian taxa. There- should be noted that we here for the first time docu- fore, we will under each species briefly outline only ment three additional thiarids, viz. Plotia scabra, Ser- those anatomical features that are typical and distin- myla riqueti and Stenomelania cf. aspirans, as taxa guishable for the species in question, if applicable, but with (more or less regular) occurrences in Australia not give here a very general diagnosis of thiarid anatomy. known to or listed by B. J. Smith (1992, 1996) in the All constituent thiarid taxa studied so far are charac- last available survey. terized by a series of anatomical features listed e.g. in Houbrick (1988) and Glaubrecht (1996). Most promi- Note on the taxonomy. We comment on and synony- nent are a pigmented head foot with broad snout, an mize within the known taxa several thiarids on the gen- oval and paucispiral operculum with excentric nucleus eric and specific level of formerly uncertain status, (instead of a round multispiral operculum in Pachychi- while elevating other names from being so far treated lidae), a mantle edge with ventrally originating papillae as mere synonyms to represent valid biological species, of which usually 3–4 are most pronounced on the aba- as will be evident from comparison with the compila- nal side of the mantle (in contrast to a smooth mantle tion presented last by B. J. Smith (1992). We restrict the edge e.g. in pachychilids). The taenioglossate radula is list of synonyms to the most important ones and in par- generally very small (< 0.3 cm) and narrow, with about ticular to those used by authors for the Australian taxa 90 to 140 rows and a central or rachidian tooth that is in those cases where the species are not endemic. Also about three times as wide as high, with a lateral dent- the literature references in the synonymy lists are only like protrusion visible under the SEM, and flanked on given in abbreviated form, but can be found in the both sides of the mesocone by three to four smaller, compilation by B. J. Smith (1992) in complete fashion pointed denticles. The lateralia exhibit very long, slen- not to be repeated here. der lateral protrusions, with one most pronounced and However, due to the preliminary character of our usually well-rounded mesocone flanked by two smaller phylogenetic analyses (see above) we refrain herein inner and mostly three to four outer denticles. In most

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 212 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae thiarids there is a wing-shaped plate on the inner lateral nomic history, as there is no doubt about the identity side, bearing these smaller inner denticles. The long and allocation of the Australian amarula within Thiara and slender marginalia are broad at their distal ends sensu stricto. However, it is important to note that we with a series of fine comb-like denticles, usually vary- here suggest to take up again Brot’s approach separating ing in number between 9–11. The organisation of the scabra and balonnensis as genetically distinct lineages inner mantle organs are that of most Cerithioidea, with from that denoted as Thiara by its type amarula. We will the exception that in all Thiaridae the female gonoduct refer to these two other thiarid taxa by re-erecting and in its pallial part (in earlier description often erro- utilizing Brot’s original generic names Plotia and Plo- neously termed “uterus”) is tube-like and completely tiopsis; see under those generic entries below. closed along its longitudinal axis, with the exception of As the concept of this genus is still diffuse, presum- a vaginal opening at the distal end, which lays a consid- ably comprising quite different species with not only a erable length behind the anal opening of the rectum. wide spectrum of shell morphologies, it is important to As a most characteristic feature all Thiaridae are vi- note that there are also distinct reproductive strategies viparous with a special incubatory structure, or brood found. On the one hand, species like Thiara amarula pos- pouch, located in the neck region of the head foot (i.e. sess brood pouches filled with embryonic stages without subhemocoelic). This feature sets the Thiaridae clearly shells only (see below), while on the other hand species apart from other caenogastropods, even those that are with shelled juveniles found in the brood pouch are cur- also viviparous, as these mostly brood eggs, embryos rently also attributed to Thiara. We here have attempted and/or juveniles in their oviduct (this time truly func- to already allocate these latter taxa, like e.g. australis, tioning as a “uterus”), like in some (but not all) SE balonnensis and scabra under separate generic names, as Asian Pachychilidae and Oriental Paludomidae within our molecular genetic studies (Glaubrecht et al. unpubl. the Cerithioidea, or the Viviparidae; for review see e.g. data) indicate that all three represent distinct phyloge- Glaubrecht (1996, 1999, 2006, and literature cited netic lineages and do not form a monophylum with typi- therein). The thiarid brood pouch, which has a pore or cal Thiara as represented by the type species. opening immediately opposite to the distal end and va- Diagnosis. Shells with pronouncedly stepped shape and gina of the female gonoduct, extends far into the right only slightly rounded but enlarged, smooth last whorl. propodium, as shown for several of its constituent taxa There are 2 9 steplike descending whorls, of which the (Glaubrecht 1996; Schtt & Glaubrecht 1999). We will – last one is always the widest and tallest (comprising here report on a variety of reproductive modes among the viviparous Australian Thiaridae and will comment about a third of the entire height); shells often eroded on these alternative strategies under each species; see and thus decollated. Axial elements are characteristi- also Part III for a summary description. cally formed as apically pronounced ribs only, some- times appearing like knots or keels, only rarely as api- cally projecting spines. Thiara Rding, 1798 Description Thiara Rding, 1798: Mus. Bolt. Cat. Cimeliorum Etribus Reg. Nat. Hamb.: 109. Shell. Moderate to large size; black; some spiral, but Melania Lamarck, 1799: Mem. Soc. Hist. Nat. Paris 1: 75. Tiaropsis Brot, 1877: in Martini & Chemnitz, Conch. Cab. 1(24): 301. more prominently axial sculpture exists. The whorls are Melacantha Swainson, 1840: Treatise on Malacology: 199. in a step-wise fashion, making the shell to appear stair- Amarula Sowerby, 1842: Conchological Manual: 61. (or pagodula-)like. The apical whorls and the tip of the shells are mostly corroded. The aperture is oval and en- Type species. Helix amarula Linnaeus, 1758; by monotypy. tirely rounded (i.e. holostome, with no sipho), but with Taxonomy. “Helix” amarula, as described 1758 by Carl a slightly flaring lip anteriorly; the umbilicus is closed, Linnaeus, became by monotypy the type species of the callus not thickened. Thiara Rding, 1798 as well as of Melania Lamarck, External morphology. The colour of the body in live 1799, the latter being hence an objective junior synonym animals is variable, mostly dark or black, but irides- of the former (ICZN, opinion 96, direction 48). The cend. The head-foot is short and squarish, usually with name Thiara (meaning “Papstkrone” in German) relates light transversal stripes. On the edge of the mantle to the stepped, high-spired shape of the shell of amarula. there are 15–20 papillae, in which the anal papilla is Due to the considerable variation in shell shape and three times larger than the others. sculpture in Thiara, there are various strikingly differ- ent concepts of this genus. Brot (1877) attempted to se- Radula. The radula consists of about 130 rows and is parate distinct conchological lineages when first sug- highly variable. The rachidian or central tooth is clearly gesting, next to Tiaropsis, the subgenera Plotia (with wider than high, with the edge bulged out downwards scabra as type species) and Plotiopsis (type species: of the basis. On the lateral edge short dentations are balonnensis). Subsequent authors have made use of this visible. The number of denticles of the rachidian varies classification scheme in different ways and by subsum- between 7–10. The inner smaller denticles of the later- ing the many named taxa quite differently. No attempt alia appear to be either isolated from each other or will be made here to unravel this complicated taxo- forming a wing-like plate. The slender, long marginals

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 213 are broader distally, with a varying number of comb- parts of the Indo-West Pacific, including e.g. the Solo- like denticles, mostly between 5–12. mons, Hebrides, New Caledonia and Fiji. Reproduction. The brood pouch of the type species Thiara Ecology. Thiara lives in lotic and lentic freshwater ha- amarula was always found to contain only early embryonic bitats, preferring, however, the headwaters and middle stages, mostly having a diameter of 0.1 mm, lacking any reaches of rivers, although it is also found often in the shell. However, in other species described traditionally un- lower courses of rivers as far down as to the brackish der this genus other reproductive modes are also found. water influence.

Distribution. Thiara exhibits a wide distribution in the Thiara amarula (Linnaeus, 1758) tropic and subtropic regions, ranging from eastern Afri- Figures 1a, 2g, 5, 6, 7 ca to India, the south of China and the Phillipines, north to Taiwan and across Southeast Asia and the Ma- Helix amarula Linnaeus, 1758: Syst. Nat. X: 774. lay Archipelago to New Guinea, the Bismarck Archipe- Buccinum amarula – Mller, O.F. 1774: Vermium terr. et fluv. Hist. lago and Australia, as well as to islands in the western 2: 137.

Figure 5. Shells of Thiara amarula (Linnaeus, 1758) in Australia, QLD. a. Woobadda River (ZMB 106348); b. Cooper Creek (ZMB 106350); c. Mowbray River (ZMB 106353); d. North Johnstone River (ZMB 106349). Scale ¼ 1 cm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 214 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Bulimus amarula – Bruguire, 1791: Encycl. Mthod.: 458. Radula (Figure 6). The taenioglossate radula corre- Melania amarula – Lamarck, 1822: Hist. nat. Ani. s. Vert. 6: 166. sponds to that of typical Thiaridae; however, as this is Paludina (Melania) amarula – Frussac, 1827: Bull. univ. sci. nat. the largest species, the radula T. amarula is much long- 10: 410. er and stronger in comparison to other thiarids in Aus- Tiara amarula – Adams, H. & Adams, A., 1854: Genera of recent Mollusca: 295. tralia. The rachidian tooth (Figure 6c) is three times Thiara amaruloidea Iredale, 1943: Aust. Zool. 10: 207. (1 syntype longer than wide, with the typical lateral tooth-like ex- AMS C.100605, 2 syntypes AMS C.109657; from Cardwell, tension clearly visible. On both sides of the enlarged, QLD). [vidi] rounded mesocon there are three to four smaller and Type locality. “Asiae fluviis”, as given by Linnaeus; i.e. more pointed denticles. The denticle number of this in Asian rivers. central tooth varies between the specimens and even within one radula. The lateralia have a distinct, very Type material. Whereabouts not known. long and thin lateral extension. Inside of the widely overhanging, rounded mesocon there are two smaller Taxonomy. The variability of the shell, with considerable teeth, on the outside there are three to four; which can differences between adult and juvenile forms, in concert vary between and within the specimens studied. with the widely adopted typological practice of naming (morpho-)species has led to a plethora of species and Reproduction. The brood pouch of the females of subspecies names in Thiara amarula (see e.g. Brot 1877; Thiara amarula is located dorsally in the neck region Riech 1937; Starmhlner 1976). In naming the speci- above the oesophagus, extending on the right side into mens from Cardwell as “amaruloidea”, Iredale (1943) the foot. Inside there are well over thousand early em- delievered at last an example of this practise. However, bryonic stages, densely packed in the entire pouch and amarula shells from Australia (Figures 1a, 5) cannot be embedded in thin compartimental tissue, as depicted distinguished by any conchological feature from those and described in detail by Schtt & Glaubrecht (1999). long known from other parts of its distributional range. Ecology. The preferred habitat of Thiara amarula in Diagnosis. Characteristic for this species is the large, Australia appears to be clearly above the influence of heavy, stair-like, turriforme shape of the thick shell, brackish water. In the Mowbray (Figure 2g) and the with three to four whorls in the adults. The largest part Bloomfield River the species was found about 4–6km of the shell, about 2/3, comprises the last whorl, with away from the coast in clear freshwater reaches of the the moderately high spire being often corroded. The ax- tributaries to larger streams, well above the intersection ial ribs, most pronounced apically, project at the outer with the lower courses of these rivers and therefore out- edge of the whorls as small spines; however, they are side the reach of direct marine influence (see detailed often missing due to corrosion. The aperture is oval map in Schtt & Glaubrecht 1999). and holostom. Distribution. The distribution of Thiara amarula ex- tends from the south and east coast of Africa in the Description west of the Indian Ocean to the Malay Archipelago, the Philippines and further out into the Indo-West Pacific Shell (Figure 5). The shell of Australian Thiara amarula reaching the Solomon and Fiji Islands as well as Sa- is dark brown to black, or partially of faint olive-green moa, as documented in Schtt & Glaubrecht (1999). colour. The overall shape is oval, with step-like whorls In Australia T. amarula is restricted within the Jardi- in both adults and juveniles. In the former the spines on nian fluvifaunal province to a small region in the north- the outer edge of the whorls are mostly degraded, in ju- east of Queensland, with mostly relatively short streams veniles they can be of about half a centimeter in lenght. and rivers draining to the Coral Sea, ranging from the The height of the complete shell reaches over 5 cm (see Boomfield River in the north to Cardwell in the south Table 1), with often more than 2/3 of this height taken (see Figure 7). Note that the geographic range given in by the last whorl. The aperture is oval and holostom, B. J. Smith (1992: 78) as to include “N Gulf, N coastal, with an angular protrusion at the upper (= posterior) WA, NT” is unsubstantiated. margin due to the spined ribs forming there. External morphology. The snails have a black pigmen- Material examined ted body with a compact shape. The edge of the mantle Queensland: Bloomfield River, 2 km N of Wujal-Wujal, Granite has about 18 to 21 papillae, including the anal papilla Creek, 200 m upstream from causeway, 6 km from coast on the right side of the mantle and body, next to the (ZMB 210026); Granite Creek, W of Bloomsfield (1555.99 S anus and the opening of the brood pouch. On the left 14519.54 E) (ZMB 107217); Woobadda River, tributary of Bloomfield hand side there are ventrally just behind the mantle’s River (1557.35 S 14521.11 E) (ZMB 106348); Douglas Creek, near edge four papillae which emerge in the living animal Daintree at crossing (16 16.194 S 145 58.60 E) (ZMB 107218); Cooper Creek (1610.43 S 14525.18 E) (ZMB 106350); Mossman, Daintree more than the other; note that Abbott (1948: 288) men- River (AMS C.109658) (ZSM 12430); Mossman River (AMS C.93924) tioned 10–14 of these mantle papillae, so that their (CAS 46935); Mowbray River (AMS C.158117, C.158275, C.317842) number apparently varies. See for a detailed description (QM 16571, 64459) (USNM 854006) (1633.87 S 14527.83 E) Schtt & Glaubrecht (1999). (ZMB 106353, 210027, 107219); Barron River, below 150 m Lake

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 215

Figure 6. Radula of Thiara amarula (Linnaeus, 1758), QLD, Woobadda River (ZMB 103648). a. Radula ribbon; scale bar = 200 mm; b. Lateral and central teeth; scale bar = 50 mm; c. Rachidian; scale bar = 10 mm; d. Marginal teeth; scale bar = 25 mm.

Placid (1652.170 S 14540.405 E) (ZMB 107220); Barron River Clump Point (AMS C.109435); Tully River (AMS C.9285); Rocking- (BMNH 1922.3.24.9) (MCZ 198983); N of Cairns (AMS C.117626, ham Bay, saltwater creek (BMNH 1879.5.21.405); Cardwell C.117617), from a creek (AMS C.158127); near Cairns (AMS C.109657; syntypes of “amaruloides”). (AMS C.109659) (MCZ 31304, 183317) (SMF 108247/2); North Johnstone River (1730.34 S 14559.55 E) (ZMB 106349, 106354); “Thiara” australis (Lea & Lea, 1851) Figures 1b, 2a–f, 8, 9, 10, 11, 48

Melania australis Lea, I. & Lea, H. C. 1851: Proc. Zool. Soc. Lond. 1850: 185. Plotiopsis australis – Iredale, 1943: Aus. Zool. 10 (2): 208. Thiara australis – Stoddart, 1985: J. Mal. Soc. Australia 7 (1–2): 9. Melania cerea Brot, 1860: Rev. Zool. 1860: 266. (holotype MHNG; unknown type locality) [not seen] Melania decussata Brot, 1862: Matriaux I: 55. (unnecessary nom. nov. for Melania australis) Melania elseyi Smith, 1882: J. Linn. Soc. Lond. Zool. 16: 261. (4 syn- types BMNH 1857.9.30.18; “Australia”, leg. J. R. Elsey, Esq.) [vidi] Sermylasma retracta Iredale, 1943: Austr. Zool. 10: 208. (types lost?; Lennard River, WA) Type locality. “Victoria River, North Australia“. Type material. Holotype (USNM 119569; coll. Lea) (Figure 8a). Taxonomy. This form, described by Lea & Lea (1851) Figure 7. Geographic range of Thiara amarula (Linnaeus, as distinct species, was allocated first by Iredale (1943) 1758) in Queensland, Australia. Note the restriction within the within Plotiopsis, created by Brot (1877) as subgenus Jardinian fluvifaunal province. but elevated by the former to generic rank. Thus, Ire-

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 216 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 217 dale suggested first to separate australis from those conical shape with a large last whorl, taking up about taxa which are more closely related to Thiara amarula. 1/3 of the entire length. The maximum of nine whorls At the same time Iredale followed Brot in allocating are stepped and stair-like, often with pronounced and balonnensis also to Plotiopsis, according to similar widely spaced axial ribs. These strong ribs are not ex- shell morphology. Stoddart (1985) retransferred austra- tended into spines or knots, but spaced relatively far lis back to Thiara Rding, 1798, revoking again the away from each other. Spiral ridges are also present, in (useful, as we think) distinction between Plotiopsis and particular at the lower part of the whorls, but are less Thiara. Apparently by mistake, B. J. Smith (1992) listed pronounced than e.g. in balonnensis. The colour is light australis merely as synonym of Melanoides tuberculata or dark brown, to reddish-brown or sometimes even Mller, 1774, to which it shows no resemblance, black. The aperture is oval to drop-shape, with the ba- though. sal margin slightly projecting. For the time being we refrain from attributing a new Juvenile shell (Figure 9). The first whorl of the em- generic name for australis which is here recognized as bryonic shell has an irregular sculpture with a clearly a clearly distinct species, distinguishable by conchologi- visible wrinkled pattern, as described for other euvivi- cal features and its exclusive geographical range (see parous thiarids, such as e.g. Tarebia granifera and Me- below). Our molecular investigations (Glaubrecht et al. lanoides tuberculata (see Glaubrecht 1996). About the unpubl. data) strongly indicate a classification separate last third of the second whorl there are growth lines both from Thiara (with amarula) and Plotiopsis (with visible, which become increasingly stronger on the balonnensis). In case of verification and thus being ne- third whorl with the appearance of pronounced axial cessary later we suggest a new generic affiliation. ribs, which are crossed by spiral ribs from there on. “Melania” elseyi, of which Smith (1882) gave a de- These axial and spiral elements are distinct; they vary, tailed description based on the four syntypes in the however, with very different patterns developing even BMNH, is probably a more elongated form, with shell in the juveniles of the same brood pouch. The shell di- sizes up to 3110.5 mm, which is well within the mor- mensions found (see Table 2) are within the range phological space of australis, according to shell meas- known for other not only Australian Thiaridae. urements of this species (Table 1). Our material, partly sampled along the historical route of the expedition External morphology. We found no clear difference in J. R. Elsey once took part in, did not produce suffi- the anatomy of australis to those of other Australian ciently distinct specimens to allow for a separate spe- thiarids under study. cies diagnosis of this taxon. Based on shells from the Lennard River from the Kimberley region in WA, Ire- Radula (Figure 10). The radula is typical for thiarids, dale (1943) also named a new species, viz. retracta (al- without allowing for specific differentiations, neither in located with his new genus Sermylasma, which is char- comparison with other closely allied taxa nor across the acterized by the type species venustula). However, the range from which we collected. In addition, we found former also resembles australis and is well located that form and number of the denticles of individual within its geographic range, therefore here tentatively teeth vary within and between populations studied. included; unfortunately, the whereabouts of the types is Reproduction. In the subhemocoelic brood pouch of unknown (Smith 1992: 79). “Thiara” australis we found a number of up to 30, but Diagnosis. The conical shells are similar to those of most regularly less than 10 juveniles with shell. In our Plotiopsis balonnensis, except for the last whorls which anatomical study, restricted so far to the material from are clearly less subsuturally angulated and stepped, but Roper River, in three quarters of all the specimens in- instead more rounded. The axial ribs are widely spaced vestigated females with a brood pouch were found. and clearly more pronounced, as are the spiral grooves Thus, we anticipate a high degree of parthenogenesis in and ridges. The anterior margin of the holostome aper- this euviviparous species to occur. ture is slightly flaring, expanding the oval opening at Ecology.“Thiara” australis is a detritus feeder and a eu- its basis. ryoecic species in the limnic habitats of northern Austra- lia. It is found in different water bodies, ranging from Description small creeks and the headwaters of rivers to their lower courses well above brackish water influence. Although Shell (Figure 8). The highly variable shells reach living both in lentic as well as lotic environments, it was heights of over 30 mm (Table 1). They are of an overall most abundant in flowing water; however, it also with-

3 Figure 8. Shell morphology of “Thiara” australis (Lea & Lea, 1851). a. Holotype (USNM 11969); NT, “Victoria River”; b. NT, Viktoria River (BMNH 1844.12.27.1.11); c. NT, Victoria River (ZMB 106619); d. WA, Fitzroy River, Fitzroy Crossing (ZMB 106693); e. NT, Howard River Crossing (ZMB 106598); f. NT, Little Roper River (ZMB 106630); g. NT, Salt Creek (ZMB 106633); h. NT, Roper River, Roper Bar (ZMB 106635); j. NT, Towns River (ZMB 106640); k. QLD, Gregory River, Riv- ersleigh (ZMB 106706). Scale bar = 1 cm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 218 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 9. Juvenil shells from brood pouch of “Thiara” australis (Lea & Lea, 1851). a–d. NT, Salt Creek; specimen 1 (ZMB 106633). a. Lateral view; b. Apical whorls, lateral; c. Apical view; d. Details of the protoconch. e–h. NT, Salt Creek; specimen 2 (ZMB 106633); e. Lateral view; f. Apical whorls, lateral; g. Apical view; h. Details of the protoconch. j–m. NT, Finnis River (ZMB 106648); j. Lateral view; k. Apical whorls, lateral; l. Apical view; m. Details of the protoconch. Scale bar = 200 mm. stands arid conditions by retreating in stagnant water sion to other thiarids further east. Note the apparently holes and even small pools with high temperatures. completely exclusive, thus vicariant (allopatric), distribu- tion of “T. ” australis in comparison to Plotiopsis balon- Distribution. The geographic range of “Thiara” austra- nensis, as revealed in Figure 48 (see Discussion). lis is limited to the tropical north of the Australian con- tinent, ranging from the Kimberley drainages with the Material examined Fitzroy River in the north of Western Australia through most rivers in the hot-wet zone of Northern Territory to Western Austalia: “Kimberley” (BMNH); Fitzroy River (BMNH the Gulf of Carpentaria drainages in northwestern 1841.11.74.93.103) (AMS); Geike Gorge, Fitzroy River (18 05 S 12543 E) (AMS C.324388) (VK 0951); Geiki Gorge (186.521 S Queensland, thus occurring in the entire Leichhardtian 12541.891 E) (ZMB 106696); Fitzroy River at Fitzroy Crossing town- fluvifaunal province (Figure 11). We found the eastern ship (AMS C.427355); bank of Fitzroy River at Sheep Yard Camp most locality so far at the lower Gilbert River, assuming (WAM 459–80); Fitzroy Crossing (1812.653 S 12534.74 E) the region of the Gregory Range to mark the faunal divi- (ZMB 106693); Isdell River (AMS); Charley River, 25.3 km WSW of

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 219

Figure 10. Radula of “Thiara” australis (Lea & Lea, 1851). a–b NT, Douglas River (ZMB 106615). a. Lateral and central teeth; scale bar = 25 mm; b. Marginal teeth; scale bar = 10 mm; c–d. NT, Salt Creek (ZMB 106633); c. Lateral and central teeth; scale bar = 50 mm; d. Marginal teeth; scale bar = 10 mm. e–f. NT, Finnis River (ZMB 106648); e. Lateral and central teeth, specimen 1; scale bar = 25 mm; f. Lateral and central teeth, specimen 2; scale bar = 25 mm.

Mt. Blythe (1622.35 S 12512.35 E) (VK 12.386); billabong on side of Victoria River (BMNH 1844.12.27.1.11, 1857.11.18.130, Weber Plains Road (1540.35 S 12844.39 E) (AMS); Fowl Yard, pool 1857.9.30.10); Victoria River, on Victoria Highway (1536.890 S in Osmand Ck., Bungle Bungle (1716 S 12830 E) (AMS); Ord River, 13107.820 E) (AMS C.427957, C.427657); Victoria River, bridge 100 m below dam, on W side of river (1547.480 S 12841.580 E) (WAM 458-50); Victoria River Crossing, just off the highway, (AMS C.427964, C.114695); Ord River, Ivanhoe Crossing (1541.22 S (1445.46 S 13135.68 E) (AMS C.22645) (VK 24.182); Victoria 12841.23 E) (AMS) (VK 24.180); lake behind Diversion Dam, Kunu- River, Old Victoria River Crossing (1534.862 S 13106.142 E) nurra (1547.51 S 12841.94 E) (AMS C.324387); Ord Irrigation area, (ZMB 106619); Victoria River, at Victoria River Gorge (1537.79 S Kununurra (WAM 728–77); Farber Beach, Ord River, Kununurra 13108.099 E) (ZMB 106621); Victoria River, 195 km West of Kather- (AMS); Lake Kununurra, 250 m along canal at pump station ine (AMS C.324389); Bulita Station, 55 km W of Victoria River (1547.340 S 12843.005 E) (ZMB 106692). Research Station, Victoria River Downs (1607 S 13025 E) Northern Territory: “North Australia” (BMNH 1857.9.30.9); (NTM P7843); Victoria River at Dashwood Crossing (1620.02 S “N.T.” (AMS C.26019, C.26017); East Baines River, at crossing of 1316.86 E) (AMS); Victoria River on Top Spring, Timber Road Victoria Highway (1545.737 S 1301.75 E) (ZMB 106697); (AMS); “Port Essington” (BMNH); Port Darwin (AMS); Howard

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 220 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 11. Geographic range of “Thiara” australis (Lea & Lea, 1851) in northern Australia. Note the occurrence in the entire Leichhardtian fluvifaunal province, including Timor Sea and Gulf of Carpentaria drainages.

Springs, 66 km E of Stuart Highway (weir, clear water, muddy bottom) Bridge (1347.36 S 13121.185 E) (ZMB 106615); Douglas River, at (1227.5 S 13130 E) (AMS C.324390, C.110490, ex C.68505) crossing to Tipperary Station, Douglas Daly Research Farm (NTM P27474); margins of Howards Springs (1227.5 S 13103.0 E) (1350.22 S 13109.71 E) (VK 24.375); 1 km upstream of junction of (NTM 27451); Howard Springs Creek (1227.268 S 1313.108 E) Douglas and Daly Rivers (on sandy bank, besides rapids) (1350.26 S (ZMB 106594); Howard Ck., Koolpinyah Stn, 30 miles from Darwin 13108.49 E) (NTM P27450); Daly River Crossing (1346.026 S (AMS); Howard River, crossing (1227.752 S 1315.008 E) 13042.688 E) (ZMB 106670, 106710, 106715, 107264); Daly River, (ZMB 106596, 106598, 106701, 106702); Berry Springs, S of Darwin Oolloo crossing (144.24 S 13115.056 E) (ZMB 106666, 107289); (1242.111 S 13059.854 E) (ZMB 106704, 107290) (AMS); Darwin Daly River at Kathleen Falls (1445.46 S 13135.65 E) (AMS C.21440, River, 46 road miles from Darwin, off Stuart Highway (1244.56 S C.324384); Kathleen Falls, Flora River NP (1445.412 S 13135.791 E) 13057.88 E) (NMT P27467) (AMS); Darwin River, Weed Quad. 1, 2, (ZMB 106618) (VK 24.179, 24.181); Katherine River, at Katherine, 4 (NMT P27469, P27472, P27468, P27470, P27473); Coomalie Low Level crossing (1429.441 S 13214.991 E) (ZMB 107288); Creek, at rest area on Stuart Highway (130.88 S 13107.04 E) Katherine River, 500 m downstream from Lower Land Bridge at (AMS C.324138); along Stuart Highway (13010 S 13107.50 E) (NTM Springvale Homestead (1429.49 S 13214.73 E) (ZMB 106698); P6467); Finnis River, NW of Batchelor (131.3160 S 13057.0930 E) Lagoon, King River, S Katherine (AMS); creek, SW of Katherine, fossil (ZMB 106648, 106649, 106664); Crater Lake, S Batchelor, NE of Ade- (AMS); creek just before turnoff from Roper Highway to Elsie Falls, laide River (132.76 S 1315.445 E) (ZMB 106659); Rum Jungle at SE of Katherine, fossil (AMS); Roper River (AMS C.109742); Elsey Litchfield Road (132.604 S 13059.862 E) (ZMB 106661); Adelaide Falls, Roper River on Elsey Station, in sandy mud, shallow running River (BMNH 1891.11.21.153-166, 1892.1.29.194); Adelaide River, water (VK 0957); Elsey Cementery, 11 km S of Mataranka Springs north, c. 18 km downstream from crossing (138.742 S 13113.14 E) (1505.15 S 13307.44 E) (AMS C.339837); Elsey River, at Elsey (ZMB 106610); Adelaide River, south, at crossing (1328.975 S Cementery (ZMB 106652); Warloch Ponds on Elsey Creek, near old 1315.853 E) (ZMB 106611); Scott’s Creek, 9 km E of Adelaide River Stuart Hwy, Elsey Station, Mataranka area (165.042 S 1337.258 E) (VK 0969); Glass Water Swamp, Litchfield Station (1319.52 S (ZMB 192014); Waterhouse River, Stevie’s Hole (1455.782 S 13032.57 E) (VK 24.370); Bamboo Creek, 3–10 m from Daly River 1338.732 E) (ZMB 106680, 107287); Little Roper River, at crossing (1340.118 S 13039.501 E) (ZMB ZMB 106612, 106672, 106674, (1455.581 S 1337.176 E) (ZMB 106627, 106630, 106677, 107285, 107263) (VK 24.394); Douglas River Crossing (1340.09 S 107286) (AMS C.317321); Mataranka, 1 km of Kowai Roper Ck. near 13039.59 E) (AMS) (VK 24.369); Douglas River crossing, Bond junction with Waterhouse Creek (1455.740 S 1337.060 E) (AMS

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 221

C.5776, C.324383); Roper River at 4 Mile Point (1456.137 S Taxonomy. This genus, albeit maybe best known among 13310.033 E) (ZMB 106625, 107284); Mulurark Rapids at Elsie Park, all thiarids, is highly polymorphic in its shell and vari- Roper River (14 56.68 S 133 12.38 E) (AMS C.324385) (VK 24.384– able in all characters studied. It is also widely distrib- 386) (ZMB 106624); Roper River, at Jalmurark Campground (1457.158 S 13313.29 E) (ZMB 106675, 107265); Roper River, uted in the Old World, with introductions now in the Roper Falls, 4 km E of Jarmurak (1457.401 S 13315.018 E) New World, and with a plethora of specific names de- (ZMB 107283); Salt Creek nr Elsey Creek, at crossing of Roper High- scribed and subgenera attributed (see Glaubrecht 1996; way (150.703 S 13314.417 E) (ZMB 106631, 106633, 106683, Facon et al. 2003). A comprehensive monograph is still 106684, 107266); Elsey Creek on Roper Highway (150.627 S missing, but highly wanted and timely. In addition to 133 15.096 E) (ZMB 106707, 107267); Roper River at Roper Bar the enormeous conchological variability and recently (14 42.802 S 134 30.474 E) (ZMB 106635, 106708, 107268) (AMS) worldwide distribution, typical Melanoides is also (VK 25873, 25852) (NTM P27475); East Alligator River, Ubir, at Cahills Crossing to Arnhem Land (1225.542 S 13257.882 E) known to have parthenogenetic reproduction and thus (ZMB 106642); Goyder River, Arnhem Land (AMS); eastern Goyder produce clonal lines, which renders the application of River crossing, Central Arnhem Land (1302.7 S 13497.7 E) the biospecies concept even more problematic. For ex- (AMS C.461368); Rose River catchment, southern end of Parsons ample, in Africa there are at least 28 species named Ranges, Numbulwar District, 150 km N of Roper Bar, eastern Arnhem within this genus but their systematics is unresolved; in Land (13 43.40 S 135 06.2 E) (NTM P8703); Mumpumapu waterhole, Southeast Asia there are usually 9 species considered; Phelp River drainage, Numbulwar-Roper rd., Mumpumumpu Outsta- tion, Arnhem Land (1438.3 S 13532.6 E) (AMS C.461355); Wandoo see review and references in Glaubrecht (1996). This River, Numbulwar-Roper River rd., Arnhem Land (1414.02 S situation makes it highly problematic to establish firm 13536.11 E) (AMS C.461357); track to Arthur rd. and Numbulwar, ground for the samples found in Australia. Thus, we Koolatong rd. crossing, Arnhem Land (1310.5 S 13572.6 E) here refrain for the time being from a more formal (AMS C.461365); Wilton River, Arnhem Land (VK 0956); ca 8 km NE treatment of this taxon. of Towns River Crossing, Gulf of Carpentaria (1459.82 S 13516.28 E) (ZMB 192015); ca 3.8 km NE of Towns River Crossing, Gulf of Carpentaria (151.199 S 13514.136 E) (ZMB 190010); Towns River, at crossing (152.57 S 13512.718 E) (ZMB 106640, 107269); Melanoides tuberculata (O. F. Mller, 1774) Cox River (in stagnant waterhole N of causeway) (1519.394 S Figures 1c, 12, 40 13520.699 E) (ZMB 107270); Cox River crossing, billabong 2 km SE (1520.30 S 13521.15 E) (VK 25.912); Limmen Bight River, at road Nerita tuberculata Mller, 1774: in Verm. terr. et fluv. hist. 2: 191. crossing (1528.865 S 13524.054 E) (ZMB 107271) (AMS); Melanoides fasciolata Olivier, 1804: Voy. l’Emp. Othoman 2: 69. McArthur River crossing, at Booroloola (1604.866 S 13619.026 E) (whereabouts of types unknown; from Alexandria, Egypt) (ZMB 107272) (VK 24.374); junction of rocky creek and McArthur River at Borroloola (1605.00 S 13618.50 E) (VK 24.381); Wearyan Type locality. “In littore Coromandel”; i.e. India, Coro- River, along beach at crossing (1610.02 S 13645.481 E) mandel Coast. (ZMB 107273) (AMS C.324386) (VK 13.864, 25.834); Foelsche River Type material. A total of 35 syntypes (3 þ 32 speci- (16 12.628 S 136 53.034 E) (ZMB 107274); Robinson River, at road mens in two lots) were found by M. G. to be extant crossing (1628.27 S 13702.932 E) (ZMB 107275); Calvert River just below junction with Bluey Ck., on Savannah Way (1656.066 S in the ZMUC, marked as “type” and with label “Ex 13721.578 E) (ZMB 106709, 107276) (AMS) (VK 25.835, 26.348). org[inal] Mulleri et legit Fbr. Ind. Oc.” (see Fig- Queensland: Gregory River, SE of Burketown at Savannah High- ure 40a for three of these syntypes). Note that the way crossing (1753.517 S 13917.209 E) (ZMB 107277); Gregory claim in B. J. Smith (1992: 76) of type material and River at Gregory Downs (1838.695 S 13914.875 E) (ZMB 107278); locality being unknown is erroneous. Lawn Hill Creek at Adels’ Grove, Gregory River catchment (1841.365 S 13831.81 E) (ZMB 106705); Lawn Hill Creek, nr the Taxonomy. The species is highly polymorphic, with Cascades, Lawn Hill NP (1842.00 S 13829.00 E) (VK 26.353, shells being very variable (see above). Several species 26354); Lawn Hill, Boodjamulla Creek, downstream of Indarri Falls were described and lists of synonyms published (e.g. (1842.051 S 13829.196 E) (ZMB 107281); Gregory River at Rivers- leigh (1901.116 S 13843.529 E) (ZMB 107279); Gregory River Germain 1921; Pilsbry & Bequaer 1927; Rensch 1934; crossing, Riversleigh Station (1901.25 S 13843.22 E) (VK 26.357) Riech 1937; Benthem Jutting 1956, 1959; Brandt 1974; (ZMB 106706); O’Shanassy River (1901.354 S 13845.741 E) Starmhlner 1969, 1976, 1983, 1984, 1993; Brown (ZMB 107280); Judy Lagoon, Armraybald Station, SE of Burketown 1994); for a description of the general anatomy, radula (1757.37 S 13945.12 E) (VK 26.360); small stream, 6.5 km N of Al- and juveniles see Glaubrecht (1996). mora (18 14.27 S 139 15.38 E) (VK 26.355); Bynoe River, at cross- Note that Melanoides tuberculata, found only re- ing (17 51.53 S 140 47.58 E) (VK 26.350); Gilbert River, at Burke cently in Australia, is presumably introduced to various Road crossing, NE of Normanton (1710.117 S 14145.999 E) (ZMB 107282) (VK 26.361). localities on the continent (Glaubrecht 2000). It is very similar in its overall shell shape, size and sculpture to the native “Stenomelania” denisoniensis (as depicted in Melanoides Olivier, 1804 Figure 40 in comparison with that species; see also Ta- ble 1), from which we were unable to clearly separate it Melanoides Olivier, 1804: Voy. l’Emp. Othoman 2: 69. based alone on conchological characters of adult or ju- Type species. By monotypy, the type species of the venile shells and radula. However, our preliminary mo- genus is Melanoides fasciolata Olivier, 1804, which is lecular studies (Glaubrecht et al. unpubl. data) revealed a junior synonym of “Nerita” tuberculata Mller, 1774 populations of M. tuberculata as genetically clearly dis- (see Morrison 1954: 32). tinct evolutionary entities and at the same time dis-

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 222 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae closed its presence as non-native element to the Austra- anthropologically influenced water bodies, like irrigation lian fauna. canals and artificial lakes; see list of locations below. Distribution. Melanoides tuberculata exhibits outside Description Australia a pantropical distribution with many introduc- tions (Glaubrecht 1996, 2000; Facon et al. 2003). Its Shell (Figures 1c, 40a). The shell is slender, high- spotty occurrences in Australia (Figure 12) is verified spired, with 10–15 whorls; it is thin and coloured for several samples, among them e.g. from the north of brown to black. The suture is sunken with the whorls Western Australia (Lilly Lagoon, nr. Kununurra; being only slightly shouldered, but instead rounded. ZMB 106690), by molecular data (Glaubrecht & Brink- The highly variable sculpture is dominated by pro- mann, unpubl. data). However, in other cases its iden- nounced axial and spiral elements resulting in a more tity at other localities is based on shell morphology or less reticulate pattern, often also with tubercles. This only yet. Note that the known samples listed here are reticulate sculpture is already found in the juvenile from the immediate vicinity of larger cities, indicating shell (see Glaubrecht 1996). The spire is about five possible introduction(s). times higher than the aperture, but often corroded. The aperture is holostom. Material examined

Ecology. Found in virtually all limnic water bodies of the Western Australia: Lilly Lagoon, Kununurra (ZMB 106690) tropics, from the headwaters to the estuaries, as it also (1546.825 S 12844.477 E). tolerates brackish water conditions (Glaubrecht 1996). Northern Territory: Darwin, George Brown Botanic Garden As is the case mostly in Australia, it can also be found in (1226.739 S 13050.179 E) (ZMB 103694, 106592); One Mile Railway

Figure 12. Occurrences of Melanoides tuberculata (O. F. Mller, 1774) in Australia. While the identity of a sample from the north of Western Australia (Lilly Lagoon, nr. Kununurra; ZMB 106690) is verified to be as of M. tuberculata by molecular data (Glau- brecht, unpubl. data), evidence of occurrence at the other localities is based on shell morphology only yet. Note that the samples are from the immediate vicinity of larger cities, indicating possible introduction(s).

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 223

Dam, Darwin (1227.39 S 13050.44 E) (AMS C.322948); Darwin, cation is given as if in “Arnhem Land (as Arnheim’s Ludmilla Creek (1225 S 13050.5 E) (AMS C.307840); Berry Springs, Land)”; however, it is clearly to the west of this terri- S of Darwin (12 42.111 S 130 59.854 E) (ZMB 107210); Manton River tory, with no tributary draining into it from Arnhem (1250.282 S 1317.998 E) (ZMB 106603); Blacksoil Plan, Adelaide Land. River, Arnhem Highway (1239.7 S 13120.3 E) (AMS C.322949); Springvale Hmst, Katherine (1430.09 S 13213.72 E) (AMS C.322950); Type material. 5 syntypes (BMNH 1870.10.26.124) river, E of Maturanka (14 42.94 S 34 30.44 E) (AMS C.317344). (Figures 13a–b). Queensland: Buchans Pt., N Cairns (AMS); Crystal Creek, Paluma, N Townsville (ZMB 104144); Alice River, 25 km W Townsville (1920 S Taxonomy. In the BMNH collection there are actually 146 40 E) (ZMB 104145); Airlie Beach (AMS); Frenchman Creek, two samples (BMNH 1857.9.30.11, leg. J. R. Elsey, Rockhampton (23 21 S 150 34 E) (AMS C.414973); Brisbane (AMS). Esq.; and BMNH 1870.10.26.124) which are both sup- New South Wales: Cudgen Lake, south end (2819.66 S 15333.47 E) (AMS C.337978); Yamba, nr. Grafton, in fish tank posed to be type material of Melasma onca. For both (AMS); Sydney, St. Peters, Brown Street, in fish pond (AMS). the location “North Australia” is given; however, they were inventorized in this museum at quite different times, as indicated by their accession numbers (with Melasma Adams & Angas, 1864 the first digits recording the year). Actually, only five Melasma Adams, A. & Angas, G. F. 1864: Proc. Zool. Soc. Lond. of these specimens (viz. BMNH 1870.10.26.124) were 1863: 415. sampled by the famous Stuart Expedition, mentioned by A. Adams and G. F. Angus as source of their shell Type species. Melania (Melasma) onca Adams & An- material on which they originally based their species gas, 1864; by monotypy. description. The size of these five syntypes vary be- Taxonomy. Adams & Angas (1864) created Melasma as tween 19.18 and 26.32 mm, but correspond to that gi- subgenus within “Melania” to suit the then newly found ven for the shell (with one inch = 2.54 cm) in the origi- species onca. Later authors have, however, essentially nal diagnosis of M. onca. Note also, that B. J. Smith ignored both their generic as well as specific name. (1992) by mistake listed Melasma onca as synonym of Most recently, B. J. Smith (1992: 76) subsumed Melas- Melanoides tuberculata (see above). ma with its type species onca under Melanoides tuber- culata, based on his study of type material as he expli- Description citly stated. However, M. onca is quite different from the latter species and highly distinct on the basis of Shell (Figure 13). Conical shell with an average height morphology, molecular genetics and geographic range, of about 20 mm and nine whorls (Table 1). Most re- as shown below. markable are, next to the slender, straight, evenly Brot (1877) postulated a relationship of onca and spaced axial ribs, the perl-like chain of subsutural “Melania” venustula by combining both within Sermyla grain-like nodules formed apically by these ribs. The Adams & Adams (1854). Iredale (1943) shared this basal part of the whorls have spiral grooves, albeit not viewpoint, but decided to create in addition, albeit very prominent. The shell, in particular the last whorl, superfluously, the generic name Sermylasma. Our stu- is very variably coloured, most often black to yellow- dies do not support a closer relationship of these two brownish, and with dark spots. The aperture shape is species or lineages, though (Glaubrecht et al. unpubl. drop-like to oval. data). Therefore, we here suggest to use the available Juvenile shell (Figure 14). The shells of the juveniles in and valid generic name Melasma for the discrete evolu- the brood pouch were found to have a maximum of tionary lineage represented only by onca. three whorls. Their sculpture is of wrinkled ornamenta- Diagnosis. The olive-shaped shell is distinctly charac- tion on the initial cap, subsequently followed by a faint terized by a perl-like chain of subsutural grain-like reticulate pattern with densily packed straight axial ele- nodules which are built apically by closely and ments to dominate, though, over the spiral ridges. The evenly spaced axial ribs. These ribs are largely size (Table 2) is within the usual range found for thiar- orthoclinal. ids. External morphology. No differences of M. onca were Melasma onca (Adams & Angus, 1864) found in comparison with the external morphology of other Australian thiarids. Figures 1d, 2a–c, 13, 14, 15, 16, 46–47 Radula (Figure 15). The size and general appearance of Melania (Melasma) onca Adams, A. & Angas, G. F. 1864: Proc. Zool. this species’ radula is similar to other thiarids and does Soc. Lond. 1863: 415. not allow for distinctive differentiation just based on it, Melania onca – Brot, 1877: in Martini & Chemnitz, Conch. Cab. 1 given the commonly found interspecific and intraspeci- (24): 330. fic variability. The number and shape of denticles in Sermylasma onca – Iredale, 1943: Aust. Zool. 10: 209. the central and lateral teeth of onca vary, with the num- Type locality. “North Australia, tributary of Adelaide ber of denticles of both marginalia to be mostly be- River”. Note that in B. J. Smith (1992: 76) this river lo- tween 7–9.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 224 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 13. Shell morphology of Melasma onca (Adams & Angas, 1864). a. “N. Australia” (syntypes BMNH 1857.9.30.11); b. “Adelaide River”, NT (syntypes BMNH 1870.10.26.124); c. NT, Daly River, Bamboo Creek (ZMB 106673); d. NT, Daly River, Oolloo Crossing (ZMB 106667); e. NT, Waterhouse River, Stevie’s Hole (ZMB 106681); f. NT, Roper River, Roper Bar (ZMB 106636). Scale bar = 1 cm.

Reproduction. This species is clearly exceptional, as in Ecology. Melasma onca lives on sandy to stony bottom the brood pouch of Melasma onca we found a consider- of typical flood-drought rivers and streams in the NT. It ably high number (> 150) of shelled juveniles in var- is found from the headwater region to the middle and ious stages of development. For further details see Part III lower reaches of these rivers, as e.g. in the Daly and and Figure 47 there. the Roper River (Figures 2a–c). Their diet consists es-

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 225

Figure 14. Juvenil shells from the brood pouch of Melasma onca (Adams & An- gas, 1864); NT, Rover River, Roper Bar (ZMB 106636). a. Lateral view; b. Api- cal whorls, lateral; c. Apical view; d. Details of the protoconch. Scale bar = 100 mm. sentially of detritus and algae. B. J. Smith (1992: 76) Distribution. Melasma onca is endemic to the tropical noted that Melasma onca is necrophagous, nocturnal wet northern part of the Northern Territory of Australia. and estuarine; we found it also be very active during Its geographic range is restricted essentially to the west- the day, but have no prove for its occurrence under es- ern Leichhardtian fluvifaunal province, including Timor tuarine conditions. Sea, but also Gulf of Carpentaria drainages, where this

Figure 15. Radula of Melasma onca (Adams & Adams, 1864). a–b. NT, Roper River (ZMB 106636). a. Lateral and central teeth; scale bar = 50 mm; b. Marginal teeth; scale bar = 10 mm; c–d. NT, Roper River, 4 Mile Point (ZMB 106626); c. Lateral and central teeth; scale bar = 50 mm; d. Marginal teeth; scale bar = 10 mm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 226 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 16. Geographic range of Melas- ma onca (Adams & Adams, 1864) in northern Australia. Note the occurrence in the western Leichhardtian fluvifaunal province, including Timor Sea, but also Gulf of Carpentaria drainages. species is found in the river systems of the Daly and the per River, Mulurark (1456.771 S 13312.609 E) (ZMB 106622); Elsey Roper River drainage, from the Adelaide River and Falls, on Roper River, Elsey Station (145701500 S 1331504500 E) South Alligator River eastwards to rivers in Arnhem (VK 25.851, 25841); Roper River, at Jalmurark Camp Ground (1457.158 S 13313.29 E) (ZMB 107222); Roper River, Roper Falls, Land (Figure 16). It is apparently restricted to the inland 4 km E of Jarmurak (1457.401 S 13315.018 E) (ZMB 107224) areas (where it is not under marine influence). The (VK 24.387); Elsey Park, near junction of Salt Creek and Roper River slightly isolated occurrence at Robinson River, south of (145704500 S 1331500200 E) (VK 24.388); Roper River at Roper Bar the Gulf of Carpentaria (as indicated by shells in the (1442.795 S 13430.575 E) (ZMB 106636, 107223) (VK 701, 26.351, AMS), could not be verified by own collections in 2007. 25.842); Goyder River, Arnhem Land (1301.68 S 13458.60 E) Note that in his brief account on this species (NTM P24903) (VK 10.126); western Goyder River crossing, Central B. J. Smith (1992) mentioned also other regions of Aus- Arnhem Land (14 14.02 S 135 36.11 E) (AMS C.461364); eastern Goyder River crossing, Central Arnhem Land (1301.19 S tralia than the Northern Territory, e.g. „WA, N Gulf 13458.34 E) (AMS C.461366); eastern Goyder River crossing, Central and N costal“. However, this is not verified by our sur- Arnhem Land (1301.37 S 13458.37 E) (AMS C.461370); track to vey, as our own collections and the material of several Numbulwar, Walker rd. crossing, Arnhem Land (1335.10 S museums analysed provide no indication for a distribu- 13542.18 E) (AMS C.461352); Ngukurr-Roper Bar rd., at Wilton rd. tion outside NT. Also note that his comment “intro- crossing, Arnhem Land (1440.55 S 13434.19 E) (AMS C.461351); duced from unknown country” (Smith 1992) is ob- Robinson River, at road crossing (1645.50 S 136590 E) (AMS). viously incorrect. Plotia Rding, 1798 Material examined Plotia Rding, 1798: Mus. Cat. Cimeliorum Etribus Reg. Nat. Hamb.: Northern Territory: Adelaide River (ZMB 9068); Glass Water 109. Swamp, Litchfield Station (131905200 S 1303205700 E) (VK 24.389); Daly River, road crossing (1346.02 S 13042.61 E) (AMS C.317339); Type species. Buccinum scabrum O. F. Mller, 1774; by Daly River Crossing (1346.026 S 13042.688 E) (ZMB 106671, subsequent designation (Brot 1877). 106711, 106714, 107221); Bamboo Creek, 3–10 m from Daly River (1340.118 S 13039.501 E) (ZMB 106614, 106673); 1 km upstream of Taxonomic remarks. In former taxonomic accounts or junction of Douglas and Daly rivers (1350.26 S 13108.49 E) other studies on Australian thiarids this taxon was not (NTM P8579) (VK 24.390); Oolloo Crossing, Daly River (1404.24 S mentioned. We here present the first evidence for Plo- 13115.056 E) (ZMB 106667, 107227); Kathleen Falls, Flora River tia scabra to occur among the fauna of Australia (see (144503300 S 1313503600 E) (VK 24.183); Katherine River (13430 S below under distribution). 132580 E) (NTM P15835); Katherine River, at Katherine, low level crossing, downstream from bridge (1429.441 S 13214.991 E) Diagnosis. Relatively small shell (of 10–20 mm), with (ZMB 106617, 106699); South Alligator River, Coronation Hill pronouncedly stair-like whorls and subsutural angulated (13360 S 132360 E) (AMS C.323838); Roper River (AMS C.109742); last whorl; spire mostly elongated, often decollated. Arnhem Land, Waterhouse (ZMB 19084); Waterhouse River, 1 km E of With distinct axial ribs that apically often run out in Mataranka, in stagnant pool (14560 S 13370 E) (AMS C.317348); Ste- short spines. Shells of juveniles exhibit at the end of vie’s Hole at Waterhouse River (1455.782 S 13308.732 E) (ZMB 106681, 106682, 107226) (AMS C.317333); Little Roper River, the fifth whorl slightly flexed, axial ridges or ribs at crossing (1455.581 S 1337.176 E) (ZMB 106628); Roper River, at which project outwards, as later visible also in the adult 4 Mile Point (1456.137 S 13310.033 E) (ZMB 106626, 107225); Ro- shells.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 227

Plotia scabra (O. F. Mller, 1774) mhlner 1983). Later, Abbott (1948: 291) transferred Figures 1e, 17, 18, 19, 20 scabra to Plotiopsis (there treated as a subgenus of Thiara), which Brot (1877) had earlier suggested for Buccinum scabrum Mller, O. F. 1774: Verm. terr. et fluv. hist. 2: 136. some Australian thiarids. Benthem-Jutting (1956) re-ar- Helix scabra – Chemnitz, 1786: Syst. Conch. Cab. 9: 188. ranged scabra and several other species within Thiara. Melania scabra – Ferussac, 1807: Essai m. Conch.: 73 (non Reeve, According to our new systematization of Thiaridae 1859). (Glaubrecht et al. unpubl. data), we here assign scabra Melania (Plotiopsis) scabra – Brot, 1877: in Martini-Chemnitz, Conch. Cab. 1 (24): 7. to Plotia instead of Thiara (with type species amarula) Thiara (Plotia) scabra – Preston, 1915: Fauna British India. Moll. I: 35. or Plotiopsis (with type species balonnensis). At the Thiara (Plotiopsis) scabra – Abbott, 1948: Bull. Mus. Comp. same time, our molecular studies allowed in several Zool. 100: 191. cases, where samples were at first identified as either Thiara scabra – Benthem-Jutting, 1956: Treubia 23 (2): 393. australis or balonnensis, to recognize these as belong- Type locality. “In paludosis littoris Coromandel Tran- ing to scabra due to clustering of those samples with quebari Danorum maxime vulgare; centena & ultra Indonesian populations of the latter species. benevoleutia D. Spengler”; i.e. India: Tranquebar, Coro- mandel Coast. Description Type material. 3 syntypes (ZMUC, acc. no. 329) (Fig- Shell (Figure 17). The whorls of the small shell of 10– ure 17a). 20 mm length (Table 1) is mostly well rounded, with Taxonomy. The highly polymorphic shells of scabra eight step-like whorls of which the last one is the lar- tempted conchologists to create many synonyms, of gest, comrpising about 1/3 of the height of the shell. which here only some of the most important ones are The spiral ridges often form distinct ribbons; the axial given. Within his concept of “Melania”, Brot (1877) ribs form variable, short spines, which are flexed to the treated scabra as type species of his subgenus Plotia exterior. The shell is mostly thin, of overall brown to (see also Thiele 1929; Rensch 1934; Wenz 1938; Star- black colour, with red to brown dots or flames. The

Figure 17. Shell morphology of Plotia scabra (O. F. Mller, 1774) in comparison with type material of Plotiopsis balonnensis (Conrad, 1850) and “Thiara” australis (Lea & Lea, 1851). a. Syntypes (ZMUC 329) of Plotia scabra; India, Coromandel Coast, Tranquebar; b. Holotype (ANSP 26514a) of Plotiopsis balonnensis; QLD, “Balonne River”; c. Holotype (USNM 119569) of “Thiara” australis; NT, “Victoria River”; d–f. Plotia scabra in Australia; d. QLD, Eungella (ZMB 103714); e. NT, Little Roper River (ZMB 106679); f. NT, Salt Creek (ZMB 106634). Scale bar = 1 cm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 228 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 18. Juvenil shells of Plotia sca- bra (O. F. Mller, 1774). a–d. India, Ta- mil Nadu (ZMB 200312). a. Lateral view; b. Apical whorls, lateral; c. Apical view; d. Details of the protoconch; e–h. QLD, Eungella (ZMB 103714); e. Lateral view; f. Apical whorls, lateral; g. Apical view; h. Details of the proto- conch. Scale bar = 200 mm. shape of the aperture is oval, slightly narrowing at the Reproduction. We regularly found a series of embryonic upper margin, with the basal lip slightly flaring. developmental stages, from eggs to juveniles with shells of up to five whorls, in the euviviparous females Juvenile shell (Figure 18). The shell of the juvenile is of P.scabra. On the other hand, while in some popula- characterised by distinct spiral and axial elements, tions studied females were mostly carrying eggs in their starting from the second whorl on, while the initial cap brood pouch at one time, others were found to contain exhibits the typical wrinkled sculpture found in other up to 29 juveniles in different developmental stages. euviviparous thiarids. With increasing number of whorls The details of the life-history of a given population and the step-like appearance of the shell also markedly in- any correlation of the reproductive biology of this spe- creases, due to subsutural angulation. After the fifth cies with seasons or other environmental factors re- whorl the axial ribs become most pronounced and from mains to be studied; so far a pattern could not be knobs where they are crossed by spiral ridges found. Among 50 females studied we found only eight Radula. The radula corresponds to those typical for animals without brood pouch, determined as males and other thiarids (Glaubrecht 1996; Glaubrecht & Zorn un- concluded that this species is parthenogenetic (Glau- publ. data). brecht & Zorn unpubl. data).

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 229

Figure 19. Distribution of Plotia scabra (O. F. Mller, 1774), with occurrences outside Australia; localities from Glaubrecht & Zorn (unpubl. data).

Figure 20. Occurrences of Plotia scabra (O. F. Mller, 1774) in Australia. Note the isolated locations known so far in the Daly and Roper River drainages in western Leichhardtian and those in the Jardinian fluvifaunal provinces; for details and discussion see text under the species.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 230 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Ecology. Plotia scabra can be found in stagnant as well results of the molecular genetic analyses (Glaubrecht as running water, ranging from headwaters down to et al. unpubl. data) and, therefore, here suggest to dis- coastal areas (see e.g. Rensch 1934; Benthem Jutting tinguish also taxonomically a lineage Plotiopsis, to ac- 1953, 1956; Glaubrecht 1996). It is mostly limited to the commodate balonnensis as taxon quite distinct and ge- middle courses of rivers, but is also known to be found in netically different from all other Australia Thiaridae, in brackish waters under estuarine conditions, revealing an particular the very similar australis. euryhaline tolerance (Starmhlner 1983; Glaubrecht & Zorn unpubl. data). Plotia scabra feeds on detritus and Diagnosis. The slender elongated shell is highly tur- algae. In Australia we found specimens of this species reted, with stair-like shape; often thin. The last whorl is mostly in mixed samples, for example, with “Thiara” well rounded and of stepped appearance as the suture is australis or misidentified earlier as P.balonnensis, so its sunken; also all other whorls are subsuturally angled. ecology needs to be studied in more detail. The structure of the spiral grooves is typically of a wave-like pattern. The axial ribs are raised and often Distribution. Plotia scabra has a wide distribution from pronounced as nodules, apically sometimes even with the east coast of South Africa to India and Sri Lanka knob-like protrusions. across the Southeast Asian mainland and the islands of the western Indo-West Pacific, where scabra can be Plotiopsis balonnensis (Conrad, 1850) found from the Sunda Islands and the Philippines to Figures 1f, 21, 23, 24, 48 New Guinea including the Bismark Archipelago, the Solomons, the New Hebrides and Fiji (Glaubrecht & Melania lirata Menke, 1843: Moll. Nov. Holl. Spec.: 9. (junior pri- Zorn unpubl. data); see Figure 19. mary homonym of Melania lirata Benson, 1843) (whereabouts of In Australia scabra occurs with some scattered, types unknown; from Avon River, WA) hitherto unrecognized occurrences, here recorded and Melania balonnesis Conrad, 1850: Proc. Acad. Nat. Sci. Philad. 5: 11. verified for the first time (as listed under material ex- Melania (Plotiopsis) balonnensis – Brot, 1874: in Martini & Chem- amined). Accordingly, the species is found in Berry nitz, Conch. Cab. 1 (24): 7. Springs, in the Daly and the Roper River systems in Plotiopsis balonnensis – Iredale 1943: Aus. Nat. 10 (2): 207. Melania tetrica Conrad, 1850: Proc. Acad. Nat. Sci. Philad. 5: 11 Northern Territory, as well as in Queensland in at least (non Melania tetrica Gould, 1847) (holotype ANSP 26515; from three highly separated locations along the Coral Sea Murray River) coast, thus occurring in the western Leichhardtian and Plotiopsis tetrica – Iredale 1943: Aus. Nat. 10 (2): 207. in the Jardinian fluvifaunal provinces (Figure 20). How- Melania incerta Brot, 1862: Matriaux I: 52. (nom. nov. for Melania ever, it remains open for further study whether the Aus- lirata Menke, 1843) tralian populations of this species are introduced, or re- Plotiopsis incerta – Iredale 1943: Aus. Nat. 10 (2): 208. present a hitherto cryptic species for the continent. Melania oncoides Tenison-Woods, 1878: Proc. Linn. Soc. N.S.W. 3: 5. (5 syntypes AMS C.100624; creeks nr Bourke, Darling River, NSW) Material examined Melania tatei Brazier, 1881: Proc. Linn. Soc. N.S.W. 6: 551. (nom. Northern Territory: Berry Springs (1242.153 S 13059.875 E) nov. for Melania tetrica Conrad, 1850) (ZMB 106599); Oolloo Crossing, Daly River (1404.24 S Melania subsimilis E. A. Smith, 1882: Zool. J. Linn. Soc. Lond. 16: 13115.056 E) (ZMB 107216); Bamboo Creek, 3–10 m from Daly Ri- 262 (8 syntypes BMNH 1840.10.20.35-42; from Lower Murray ver (1340.118 S 13039.501 E) (ZMB 107215); Little Roper River, River, SA) south bank at old crossing (1455.589 S 13307.137 E) (ZMB 106679): Plotiopsis centralia Cotton, 1943: Trans. Roy. Soc. S. Aust. 67 (1): Salt Creek, nr Elsey Creek (150.703 S 13314.417 E) (ZMB 106634). 145 (holotype SAMA D14133; from Innamincka, Coopers Creek, Queensland: Broken River, Eungella (ZMB 103714); North John- SA) stone River (ZMB 106343); Endeavour River (ZMB 106351). Plotiopsis subornata Iredale 1943: Aus. Nat. 10(2): 208 (11 syntypes AMS C.51804; from Burdekin River, QLD). Plotiopsis flata Iredale, 1944: Aus. Nat. 11(5): 117 (5 syntypes AMS Plotiopsis Brot, 1874 C.139444; from Breeza, on the Mooki River, NSW). Plotiopsis sociana Iredale, 1944: Aus. Nat. 11(5): 117 (whereabouts Plotiopsis Brot, 1874: in Martini & Chemnitz, Conch. Cab. 1 (24): 7. of types unknown; from Clarence River, NSW). Type species. Melania balonnensis Conrad, 1850; by Plotiopsis thrascia Iredale, 1944: Aus. Nat. 11(5): 117 (15 syntypes AMS C.139443; Tibooburra, on the Bulloo River, NSW). original designation. Taxonomy. Followig Brot (1874), who originally sug- Type locality. “Balonne River, Australia”; this river is gested Plotiopsis, Iredale (1943) included next to ba- to the west of Brisbane, QLD. lonnensis four other species, viz. P.tetrica, P.australis, Type material. Holotype (ANSP 26514a); 6 paratypes P.incerta und P.subornata. Iredale’s opinion that the (ANSP 26514) (Figure 21a). genus Plotiopsis is limited to Australia was followed by McMichael (1967). In contrast, Abbott (1948) sub- Taxonomy. As the shells of Plotiopsis balonnensis are sumed the widely distributed Plotia scabra (see above) highly variable, as is evident from Figure 21 (see also under this genus. B. J. Smith (1992, 1996) considered e.g. McMichael 1967; Stoddart 1985), many popula- Plotiopsis as subgenus of Thiara. However, we found tions have erroneously been elevated to the status of this affiliation to be in conflict with our preliminary species and named accordingly. Due to its polymorphic

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 231

Figure 21. Shell morphology of Plotiopsis balonnensis (Conrad, 1850). a. Holotype; QLD, “Balonne River” (ANSP 26514a); b. WA, Murchinson River (ZMB 106583); c. WA, Walyunga N.P. (ZMB 106658); d. QLD, S of Rockhampton (AMS 414039); e. NSW, Gwyder River, Bingara (AMS 322678); f. NSW, Dungog (AMS 421943); g. NT, Three Mile Point, Finke River at crossing of Stuart Highway (ZMB 106689); h. NT, Boggy Hole, Finke River (ZMB 106688). Scale bar = 1 cm. shell in concert with its widespread occurrences in the structed here, we follow earlier suggestions by most distant regions of Australia it is not astonishing B. J. Smith (1992: 78) in treating the above as syno- that many of the samples of this species collected and nyms of this widespread species. inventorized in various museums in and outside of Aus- E. A. Smith (1882) remarked on the resemblance of tralia have been named under different species names, P.balonnensis from the Burdekin River and “Melania” such as, for example, “Melania” lirata Menke, 1843 scabra known at that time only from localities in SE from the Avon River area in WA, “Melania” subsimilis Asia. He also assumed that “Melania” scabrella Phi- Smith, 1882 and “Melania” centralia Cotton, 1943 lippi from Java might be conspecific with balonnensis. from South Australia, as well as from several locations Later, Hubendick (1955) mentioned the conspecifity of the Murray Darling drainage in NSW. These shells, of balonnensis with thiarids that occur ouside of Aus- however, as far as is evident from the extant type mate- tralia, in particular in East India and the West-Pacific. rial and/or original descriptions all resemble more or However, the species is apparently restricted to the less the typical balonnensis. Since they also occur well Australian continent, as is evident from our collecting within the geographic range of this species as recon- and survey of all available thiarid material so far.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 232 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Description third of the entire shell length. The rim of the aperture is thin, slightly flaring at the basis. Shell (Figure 21). The shell is elongate, sometimes compact and conical and comparatively thin, with a Juvenile shell (Figure 22). The shells of the juveniles can size of about 30 mm and the height mostly being two reach a size of 5 mm and up to 5 whorls. Most of them times larger than the width (Table 1). Shell shape is were found to vary between 1–4 mm when inside the stepped; the colour light brown to black-brown, with brood pouch, where the smaller ones have about three occasional transversally crossing rows of orange-red complete whorls. Size and sculpture of the embryonic dots. There are 6–9 whorls, of which the last one is the shell is not clearly different from that of other species largest, comprising about half of the size of the shell. (Table 2 and Figure 22), with wrinkled initial cap and The axial and spiral structure is distinct, with axial ribs growth lines starting on the second whorl. On the third terminating apically as rounded knobs. The aperture is whorl more pronounced sculpture with crossing spiral oval and pointed at the upper margin, comprising one ribs and axial elements become visible that increase in

Figure 22. Juvenil shells of Plotiopsis balonnensis (Conrad, 1850). a–d. NT, Boggy Hole, Finke River (ZMB 106688). a. Lateral view; b. Apical whorls, lateral; c. Apical view; d. Details of the protoconch; e–h. WA, Murchinson River (ZMB 106583); e. Lat- eral view; f. Apical whorls, lateral; g. Apical view; h. Details of the protoconch; j–m. WA, Lake Leschenaultia (ZMB 106725); j. Lateral view; k. Apical whorls, lateral; l. Apical view; m. Details of the protoconch. Scale bar = 200 mm.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 233

Figure 23. Radula of Plotiopsis balonnensis (Conrad, 1850). a–d. WA, Lake Leschenaultia (ZMB 106725). a. Radula ribbon; scale bar = 100 mm; b. Lateral and central teeth; scale bar = 25 mm; c. Rachidian; scale bar = 10 mm; d. Marginal teeth; scale bar =10 mm; e–h. NT, Three Mile Point, Finke River (ZMB 106689); e. Radula ribbon; scale bar = 100 mm; f. Lateral and central teeth; scale bar = 25 mm; g. Rachidian; scale bar = 10 mm; h. Marginal teeth; scale bar = 25 mm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 234 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae number on the subsequent whorls. The fifth whorl in as it was also found and figured for Melanoides tuber- most of the older juveniles is much larger and about the culata (Glaubrecht 1996). The number of juveniles vary size of half of the complete shell, with the axial and spi- mainly between 3–7, in size between 1–4 mm. In fe- ral patterns already quite similar to that in the adults. males from the Finke River in central Australia we found the highest number of juveniles with n = 58; Radula (Figure 23). The radula shows the typical ap- shell size was here about 1–2.5 mm. As a consistent pearance of that found in other Thiaridae. The differ- trend we usually found that the more juveniles were in ences found in shape and number of denticles are with- the brood pouch the smaller they were in size. Appar- in the normal variation and no specific pattern was ently, P.balonnensis is a k-strategist, i.e. there are large found. For example, the more rectangular enlargement and well developed juveniles inside the pouch. The spe- of the central denticle of the rachidian (see Fig- cies was found to be apparently parthenogenetic, as in ures 23b–c in comparison to 23f–g) or in the lateralia our survey no male could be detected (Glaubrecht & as shown in Figures 23b and f, was neither consistently Zorn unpubl. data). found along the whole radula of those individuals nor in all the radulae only of members of the same popula- Ecology. The species is euryoecic, feeding on detritus; tion. Within the marginalia we found a varying number it lives in stagnant as well as in running waters. As of 6–9 denticles. P.balonnensis is also found in the very arid central Reproductive system. The snails are euviviparous and parts of Australia (Finke River drainage), it was sup- only completely developed crawling juveniles with posed to be resisting droughts (McMichael, 1967); shells comprising several whorls hatch from the brood however, we found the species there in 2005 to survive pouch, with the largest young laying to the most ante- in deep, cold permanent water holes and pools within rior part of the brood pouch, adjacent to the opening, the dry river beds only.

Figure 24. Geographic range of Plotiopsis balonnensis (Conrad, 1850) in Australia. Note the occurrences in the Greyian (Pilbara), Vlaminghian (Southwest Coast), Sturtian (Lake Eyre or Central Australian), and Mitchellian (Murray-Darling) as well as Northeast and Southeast Coast fluvifaunal provinces, but the lack of known localities in the entire Leichhardtian.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 235

Distribution. The geographic range of P.balonnensis in- bank of a dry creek (24030 S 132430 E) (NTM P4250); Finke cludes the Greyian (Pilbara), Vlaminghian (Southwest River at Finke Gorge (AMS NT 79–32); Palm Valley, Palm Creek, 0 0 Coast), Sturtian (Lake Eyre), and Mitchellian (Murray- Krichauff Ranges (24 03 S 132 44 E) (NTM P24424); Cycad Darling) as well as Northeast and Southeast Coast flu- Gorge, Palm Valley (AMS); Palm Valley, 2 km up into valley (2403.036 S 13241.680 E) (ZMB 106718); Finke River, Boggy vifaunal provinces, with locations widespread in West- Hole (shallow waterhole in dry river) (2408.113 S 13250.233 E) ern Australia, South Australia, Victoria and along the (ZMB 106688, 106719); Boggy Hole (24080 S 132530 E) (NTM coast of New South Wales (Figure 24). However, it no- P15861); Three Mile Point, Finke River at crossing of Stuart High- tably lacks from the entire Leichhardtian province, with way (2433.182 S 13314.355 E) (ZMB 106689); Walker River [i.e. the notable exception of locations in the extrem east of Palmer River tributary, Finke River drainage], nr Alice Springs this province in the Great Dividing Range, east and (AMS C.2149). south of the Gregory Range in the tributaries of the Ei- Queensland: “Central Queensland” (AMS C.118147); Georgina 0 nasleigh and thus Gilbert River (verified in 2007) and River (AMS); King Creek, 21 km S of Bedourie (24 31.93 S 13933.90 E) (AMS C.322677); Einasleigh River, 4 km E of Eina- reported from the tributary region also of the Flinders sleigh (1830.938 S 14406.682 E) (ZMB 107260) (AMS); Soda River that both, interestingly, drain to the NW into the Gorge Spring and Soda Valley Creek (2036.300 S 14402.000 E) Gulf of Carpentaria. Also most remarkable are the iso- (AMS); Innot Hot Springs (AMS); Endeavour River Falls (1522.270 lated occurrences of P.balonnensis in the southern part S 14501.770 E) (ZMB 106346, 107256); Bloomfield River of the Northern Territory (i.e. Sturtian or Lake Eyre (AMS C.488); Barron River at Gilies Highway, Atherton Tableland province in central Australia), where it is historically (AMS); Barron River Gorge, half way to hydro station at river access reported and still extant in the Finke River drainage (1651.170 S 14539.791 E) (ZMB 107259) (AMS); Barron River, at (verified 2005). In addition, the species is also reported Barron Gorge Power Station (AMS C.127066, C.322679); rapids be- from isolated localities along the Georgina and Dia- low Lake Placid, lower end of Barron River Gorge (16 52.17 S 14540.405 E) (ZMB 107257) (AMS); Barron Falls (AMS C.9284); mantina River drainages. Rocky Creek, Atherton Tableland (AMS C.158252); Lake Eacham Note that P.balonnensis lacks in the Leichhardtian (QM 14001); Lake Barrine (1715 S 14538 E) (AMS); Peterson province, where another thiarid, viz. “Thiara” australis, Creek, Gauging Stn. (QM 64462); Bellenden-Ker Range, nr Babinda clearly dominates (see Figure 48). Note also that un- (AMS); Fisher’s Creek, Palmeston Rock, at Palmerston Hwy named fossiles from the late Tertiary found in the (1734.167 S 14553.876 E) (ZMB 107259) (AMS C.126522); John- Northern Territory were recorded by McMichael (1967: stone River (QM 64490); Herbert River, N of Ingham (AMS); Water- 0 0 139). view Creek, Iourama Falls NP (18 52 S 146 07 E) (AMS); Ross River (1921.730 S 14643.930 E) (AMS C.338671); Ross River (19170 S 146490 E) (AMS C.338666) (BMNH 1884.12.27.8-18); Burdekin River (BMNH 1846.10.?.26-29); Burdekin River, nr Charter Material examined Towers (AMS); Alice River, Townsville (19190 S 146350 E) (AMS); Western Australia: De Grey River (20110 S 119110 E) (AMS); De Hervey Range, 100 km SW of Townsville, Fletcher Creek crossing Grey River, E of Pt. Hedland, at crossing of Gt. Nt. Highway (19460 S 146050 E) (AMS); Fletcher Creek, at Gt Basalt Wall, nr (20200 S 119120 E) (AMS); Fortescue River, at NW Coastal High- Charters Towers (QM 14196); Charter Towers, NW at Toomba Stn, way, SW of Dampier (21180 S 116080 E) (AMS); Pilbara Springs, Gt Basalt Wall (19580 S 145340 E) (QM 53804); Charter Towers, 51 Miaree Pool, S. of Dampier (2051.150 S 11636.500 E) (AMS); Mill- miles WNW (QM 64488); Lolworth Creek, nr Gt Basalt Wall (QM stream N.P., top end of Lily Pond/Crystal Pool (21350 4000 S 14211); Alligator Creek, Charters Towers to Wando Vale Rd. crossing 117040 E) (AMS); Hamersley Region, Karijni NP (2228.521 S (QM 64494); Bowen (AMS); Bowen, Port Denison, headland between 11833.080 E) (ZMB 106727); Murchinson River, Kilbarri N.P., at Kings and Queens beaches (AMS); Port Denison (BMNH Ross Graham Lookout (2748.770 S 11428.540 E) (ZMB 106583); 1879.5.21.489-90); Rolleston River, Mount Cooper (AMS); Gregory Murchinson River, 5 km upstream NW Coastal Highway (27490 S River, 10 miles N of Proserpine (AMS); Mrytle Creek, crossing 1 N 114440 E) (AMS); Ellendale Pool, Greennough River (WAM 4605- on Bruce Highway, 5 miles N of Proserpine (AMS C.322654); Salt- 80); “Perth” (BMNH); Swan River (BMNH); Avon River, NE of water Ck., Proserpine (QM 4412); Pioneer River, 16 miles from Perth, Walyunga (31440 S 11640 E) (AMS C.60411, C.322680, Mackay (AMS C.118146); Lethe Brook, 5 km S of Proserpine on C.322655); Walyunga Pool, NE corner (3144.080 S 11604.280 E) Bruce Highway (AMS); Cattle Creek W of Mackay, 2 km W of Finch (ZMB 106658); Avon River, NE of Perth, 1 km upstream from park- Hatton township (AMS); Broken River, Eungella Dam, 80 miles W of ing area, Walyunga N.P. (31440 S 11640 E) (WAM); Lake Lesche- Mackay (AMS); Cedar Creek, tributary of S. Pine River, on Daybora- naultia, Chidlow (WAM 471-80); Lake Leschenaultia, NE corner Samford Rd. (AMS); N. Pine River, 2 miles S of Dayboro on Day- (3151.020 S 11614.990 E) (ZMB 106725); Ashburton River, 500 m boro-Samford Rd. (AMS); Denison Creek, on Nebo-Mackay Highway E of NW coastal highway (21580 S 115010 E) (AMS). (AMS); 16 km S of Sarina, Sarina-Malborough Rd. (AMS); Boyne Northern Territory: Finke Creek (BMNH 1908.12.8.7-9); Finke River at Rosedale, S of Gladstone (2413.400 S 15115.300 E) (AMS); River, nr Glen Helen (23250 S 132160 E) (NTM P15965); Glen Barambah Ck., tributary of Boyne River, 8 km SE of Gayndah Helen Reserve (23250 S 132160 E) (NTM P15858); Finke River, (2619.9830 S 15211.9830 E) (AMS); Boyne River, Gayndah (AMS); at Glen Helen Gorge, nr resort (2341.322 S 13240.606 E) Twelve Mile Creek, Boyne River, nr Gladstone (AMS); Eastern Boyne (ZMB 106687, 106716); Ormiston Creek, Ormiston Reserve (23380 River, S of Gladstone, Many Peaks Range (2418.150 S 15123.300 E) S 132450 E) (NTM P15859) (AMS); Ormiston Gorge, outlet (AMS C.322660, C.274Q); Calliope River, S of Gladstone, Bruce (2337.704 S 13243.375 E) (ZMB 106686, 106717); Hermannsburg Highway (AMS); Bobs Creek, Fitzroy River, nr Rockhampton (AMS); (BMNH 1909.10.23.9-11); Finke River, Hermannsburg Mission Fairy Bower, 6 miles from Rockhampton (AMS); Rockhampton (26200 S 136000 E) (AMS C.322658); Finke River, at junction (AMS); Rockhampton, Yeppen Lagoon (AMS); Baffle Creek, 4 km E with Larapinta Drive and turnoff to Palm Valley, near Hermanns- Miriam Vale (QM 4274); Woolwash Lagoon, S of Rockhampton burg (on and under rocks in river) (AMS); Finke Gorge, SW of (23230 S 150290 E) (AMS C.414039); 15 km N of Miriam Vale, Alice Springs (dead, exposed on surface of red sand, on top of 2 km E of Bruce Highway (24160 S 151290 E) (AMS C.322656);

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 236 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Rockhampton, Frogmore Lagoon (AMS); creek nr Springsure (AMS); Roma (QM 64487); Kings Creek, Darling’s Down, Post-Tertiary fossil Port Curtis (AMS C.109649) (BMNH 1928.5.5.144-149); Montrose (AMS C.109774); N. Pine River, at Young’s River crossing, W. of Pet- Creek, 163 km S Sarina (22390 S 149330 E) (AMS); Miriam Vale rie (AMS); Howard Ck., 12 km from Mt Tambourine on Beenleigh (AMS C.109427); NW of Miriam Vale, Colossem Ck. (2424.200 S Road (AMS C.128695) (QM 10370). 15128.300 E) (AMS); 18 miles S of Biggenden on Biggenden-Gayn- New South Wales: Mole River, below Bonshaw Weir, tributary of dah Road (AMS); Kolan River, Bruce Highway (AMS); Prospect Ck., Richmond River, NW of Casino, on Edenville-Doubtful Creek Rd. Biloela (AMS); Prospect Creek, Dawson Valley (AMS); Granite (28440 S 152570 E) (AMS C.167652); Richmond River, Woodburn Creek, crossing Bruce Hway, N Gin Gin (24280 S 151350 E) (QM (AMS); Upper Richmond River (AMS); Richmond River, Casino, up- 43552); Gin Gin (AMS C.41983, C.42297); Granite Creek, 36 km S stream from Irving Bridge (28520 S 153030 E) (AMS C.31555, of Miriam Vale (AMS); Walkamin, Granite Creek (QM 64466); Daw- C.167443); Murray Swamps (AMS); Upper Clarence River (BMNH son River, 8 miles from Moura (AMS); Dawson River, Taroom 1879.5.21.482-4); Clarence River, nr Baryulgil crossing, S of Tabu- (2538.70 S 14947.690 E) (AMS C.327313); Taroom, banks of Daw- tum (29130 S 152330 E) (AMS); Little River, Bawdens Bridge, Graf- son River (25390 S 149470 E) (QM 56522); Taroom, NE at Croker ton (AMS); Wollombi River, Bulga, W of Singleston (AMS); Mac- Gully (25270 S 150090 E) (QM 56586); Burnett River, W of Child- quarie River (AMS); Lake Lidell, Hunter Valley, nr Musswelbrook ers, causeway on road WSW of Booyal (2513.450 S 15200.450 E) (AMS C.110529) (WAM 460-80); 25 miles S of Forbes, on Newell (AMS); Kalliwa Creek, tributary of Burnett River (25210 S 151520 Highway (small irrigation stream) (AMS); Peel River, Tamworth E) (QM 64489); Mingo Crossing, Burnett River (QM 64463); Isaac (AMS); Halls Creek, Goulburn River, Denman, Hunter Valley (AMS); and Burnett Rivers (BMNH 1885.6.12.48-60); Iris River, S of Child- Worondi Riverlet, nr Gouburn River, Hunter Valley (AMS); Dalwood, ers (25140 S 152220 E) (AMS); Burnett River, W of Childers, cause- Hunter River, below Wynwards estate winery (sandy stream) (AMS); way WSW of Booyal (2513.450 S 15200.450 E) (AMS); Eidsvold Hunter River (AMS); Lake Burragorang (3400 S 150260 E) (AMS C.33773); Mary’s Creek, nr Gympie (AMS); NW of Gympie, (AMS C.311883); Wah Wah Main, Murrumbidgee Irrigation Area Wide Bay Creek (26040 S 152140 E) (AMS); Mary River (AMS); (AMS C.110529); Hay, Murrumbidgee River (BMNH 1836.7.26.183- Borumba Dam, Mary River (QM 3028); Mary River, at Kenilworth 92) (AMS); Spring Creek, Backmead (via Casino) (AMS); Narrandera (AMS); Deacon’s Creek, on Bruce Highway, between Maryborough (irrigation canals) (AMS); Narrandera (inland Fisheries Pond) (AMS); and Gympie (AMS); Isis River, Bruce Highway, S of Childers Yanco, Agricultural High School (AMS C.57886); Lachlan River (AMS); Barambah Ck., Central Burnett Highway (26200 S 152120 (AMS C.109644); Gooloogong, Lachlan River (AMS C.57038); La- E) (AMS); Burnett River, on Mt. Perry Road, nr Gayndah (AMS); chlan River at bridge, 8 miles from Lake Cargelligo (AMS); Lake Burnett River at Trurich Creek, W of Childers (2518.400 S Cargelligo, Willow Dam, Barren Box Swamp, 25 miles from Griffith 15156.150 E) (AMS); Burnett River, Ceratodrus (28080 S 151040 (AMS); Leeton (dried canal and drainage canal) (AMS C.109646); 3 E); Burnett River (ZMB 46270, 104176); Coominga, NW of Ipswich, miles W of Hillston (small irrigation canal) (AMS); 25 miles S of northern corner of Atkinson’s Dam, Lower Lockyer Irrigation Project, Forbes, on Newell Highway (small irrigation stream) (AMS); Cudge- nr boat ramp (27370 S 152470 E) (NTM P27453); Atkinson’s Dam, gong River, nr Mudgee, Twelve Mile (stagnant river pools) (AMS); 50 km W of Brisbane (NTM P27458); Woody Point, Clontarf, Red- Bogan River, Brewarrine (AMS C.109865); Rochs, Barwon River, cliff Peninsula, Moreton Bay (AMS); Clontarf, Moreton Bay (AMS); Brewarrina (AMS); Macquarie River, nr Carinda (AMS); Marthaguy Sandgate, Moreton Bay (AMS); Brisbane (AMS C.109648 spinose Creek, tributary of Macquarie River, W Carinda (AMS C.100836); forms); Brisbane River, 60 miles from bay (AMS); Obi Creek, nr Macquarie River, nr Dubbo (AMS); Dubbo, Western Plains Zoo Maleny, Landsborough Shire, N Brisbane (26430 S 152530 E) (NTM (AMS); Crunningbar Creek, Warren (AMS); Gouburn River (32030 S P27456); Moggill Creek, Brisbane (in freshwater) (27290 S 152540 150100 E) (AMS); Namoi River, Narrabri (AMS C.263) (BMNH E) (NTM P27459) (AMS); Moggill Creek, Brookfield (27300 S 1894.6.5.194-197); junction of Bibba Creek and Namoi River, SE of 152300 E) (QM 64461); Kilcoy (AMS); Dalby (AMS); Camp Hill, Narrabri (AMS); Namoi River at Tarriaro Bridge on Narrabi-Turra- Brisbane (27300 S 153050 E) (QM 54747); Mt Crosby Weir & wan Rd. (AMS C.101257); Namoi River, nr Gunnedah (AMS); Namoi Pumping Station, Brisbane (27320 S 152480 E) (NTM P27455); River, 6 miles N of Boggabri (AMS); Poncaree, Darling River Candamine River, nr Cecil Plains (AMS); Brisbane River (AMS); (AMS); N bank of Darling River, Red Bank Station, 37 miles SW of Pullen Creek, Brisbane River (QM 64460); tributaries of the Brisbane Bourke (AMS); Bourke (AMS C.100624, syntypes of oncoides); Me- River (WAM 463-80); River Brisbane, nr Ipswich (BMNH nindee, Darling River, Krinchega NP (on riverbank) (32240 S 1886.4.26.168-77); Ipswich (AMS); Ipswich, Swanbank Powerstation 142230 E) (AMS C.322676); E Bank of Lake Menindee (32200 S (AMS); Brisbane, The Gap, Walton Bridge, Recreation Reserve 142200 E) (AMS); Barraba, Gizzard of Black Duck (AMS); Lowry (27260 S 152570 E) (QM 47184); stream flowing out of Enoggera Creek, Warrabah NP, 35.2 km NE Manilla (30330 S 150540 E) Reservoir, The Gap, W Brisbane (27270 S 152550 E) (NTM (AMS); Cobbadah Creek, nr Barraba (AMS); Myall Greek, Bingara P27457); Coomera River, nr Canungra (AMS C.109645); Tartar’s Ck., (AMS C.109781) (QM 64486); river nr Bundarra (AMS); Gywder Macpherson Ranges (28280 S 152500 E) (AMS C.129350); Hoff- River, Moree (AMS); Gwyder River, Anderson Creek, 2.5 km S of man’s Ck., tributary Hewtsons Hill, 14 km E of Legune (28260 S Upper Bingara, SE of Moree (AMS); Gwyder River, 400 m E of road, 152240 E) (AMS C.128691) (QM 10366); Balonne River (AMS) bridge at Bingara (2951.740 S 15034.650 E) (AMS C.322678); Mor- (BMNH 1859.10.24.2); Dalrympie Creek, 10.8 km from Goomburra ee (AMS C.51670); ”Yurunga” Warialda, fossil in lacustrine deposit, (28000 S 152150 E) (AMS C.129351) (QM 10534); Moonie River eroded creek bank, 80 –100 below present level of land (AMS C.33009); Ban Ban (AMS); Caboolture, Burpengary Creek (AMS C.87453); MacIntyre River, 3 m N Inverell (AMS); Kings (QM 64491); Condamine River, weir at Chinchilla (QM 5162); Cho- Creek, off MacIntyre River (AMS C.146292); N of Inverell, 8 miles wey Creek, 15 km W of Biggenden (QM 14182); Chinchilla Weir NW of Graman (AMS); MacIntyre River, Inverell, Stannifer Rd. (QM 19443, 19457); Marlborough, S at Princhester Creek (22530 S (AMS); Moonie River 40 miles NW of Collarenebri (dry bed) 150010 E) (QM 19932); Marlborough Creek, banks Godhelp (QM (AMS C.33009, C.109647); Barwon River, 10 miles downstream from 64493); Barcaldine (QM 35222); Carnarvon Gorge, Carnarvon NP Walgett (AMS); Barwon River, Brewarrina (rocks) (AMS); Nenegara (QM 44144, 64498); Banana, 6 km S Banana – Biloela (24280 S Waterhole, Paroo River, at Wanaaring (AMS); Bulloo River, Tiboo- 150250 E) (QM 50643); Blackall, W at Noonbah Stn (24070 S burra (AMS C.139443, syntypes of thrascia); Menindee, 27 ft. below 143110 E) (QM 53110); Springsure, Minerva Creek, nr Marmadilla surface, fossil (AMS); reservoir nr Silverton, W. of Broken Hill Stn (24000 S 148080 E) (QM 60714); Charleville, Warrego River (AMS); Darling River at Bourke (AMS); Pocucaree, Darling River (QM 64455); Roma, Bungil Creek (QM 64456); Taloona Homestead, (AMS).

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 237

Victoria: Victoria (BMNH 1879.1.21.10); Echuca (AMS C.1927); Ripalania queenslandica (E. A. Smith, 1882) Lake Culluleraine (AMS); Goulburn River, Shepparton (AMS). Figures 1g, 2h, 25, 26, 27 South Austalia: Coopers Creek, at Innamincka (AMS); Cullymurra Waterhole, 11 km E of Innamincka (2742.10 S 14050.110 E& Melania queenslandica E. A. Smith, 1882: J. Linn. Soc. Lond., 2742.580 S 14053.10 E) (AMS); Adelaide (AMS C.109708); Murray Zool. 16: 261. River (BMNH 1879.5.21.461-4); Murray Bridge (AMS C.109404); Ripalania queenslandica – Iredale, 1943: Aus. Zool. 10 (2): 209. Mundalla, nr Bordertown (AMS); Tailem Band, Murray River (AMS C.43663); Morgon, Murray River, fossil from Post-Pleistocene Type locality. „Saltwater Creek next to the coast, Card- (AMS C.109780). well“ and „Paroo River“, both „Queensland, Australia“, as originally given by E. A. Smith (1882); see also B. J. Smith (1992: 77). Note that if the Paroo River, tri- Ripalania Iredale, 1943 butary of the Darling River in southern QLD and Ripalania Iredale 1943: Aus. Nat. 10 (2): 209. Northern NSW (at 31 28 S 143 32 E) was meant, this second location is certainly in error, as R. queenslandi- Type species. Melania queenslandica E. A. Smith, 1882; ca does not occur there. by monotypy. Type material. 5 syntypes (BMNH 1879.5.21.421––3 und Taxonomy. This monotypic genus of Ripalania was cre- BMNH 1879.5.21.403––4) (Figure 25). ated by Iredale (1943) in order to accommodate an Aus- tralian species with clearly distinct and characteristic Taxonomy. This distinct species is easily diagnosed and shell. It was subsequently listed in most accounts and to be distinguished from other Australian thiarids. The dichotomous keys of the Australian freshwater fauna species appears to be endemic to the Jardinian of Aus- separately (see e.g. Cotton 1943; McMichael 1967; tralia (see below); however, it should be noted that B. J. Smith 1992, 1996), with the specifications of shell some shells very similar in appearance are recorded characters only repeating the original description. Ire- from New Guinea in the Berlin Collection (ZMB dale (1943) mentioned the putative affinity of Ripalania 86812). It remains to be seen if, as in case of Thiara with Melanopsis (Melanopsidae), which is not substan- amarula and Stenomelania cf. aspirans from the same tiated by any means (see Glaubrecht 1996). McMichael area, the distribution of R. queenslandica might also be (1967) pointed out that comparative anatomical studies much more extensive. In this context other named thiar- were still missing to clarify if Ripalania is possibly a ids from the Indo-West Pacific should be compared synonym of Stenomelania. Our preliminary molecular with the present species. studies support, however, an independent lineage of that name clearly separate from the latter genus and other Description Australian thiarids (Glaubrecht at al. unpubl. data). Shell (Figure 25). The thick shell shows an even surface Diagnosis. The large, relatively thick shell is almost with a brown-orange to olive-green colour. The top is smooth, with the suture hardly sunken, and almost often corroded and the whole shell is encrusted with a straight outer shape of whorls. The earliest whorls are ferrous layer. The last whorl is bigger than the other usually decollated; the last whorl being the tallest with three to six and it is two times as long as the shell is more than half of the entire height of the shell, which wide (Table 1). The suture, that is only in younger indi- is of brown-orange to olive-green colour with a white viduals more distinct, is not sunken, thus the outer columellar margin at the aperture. shape of the whorls almost plain.

Figure 25. Shell morphology of Ripalania queenslandica (Smith, 1882); type material from the type locality Saltwater Creek, Cardwell, QLD. a. Syntypes (BMNH 1879.5.21.421-3); b. Syntypes (BMNH 1879.5.21.403-4). Scale bar = 1 cm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 238 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

The aperture is holostome, with the basal part embryos (or “veligers”) are found to be embedded very slightly flaring. loosely in the compartiments of the female marsupium. External morphology. The body was found to be orange Ecology. Ripalania queenslandica was found to live in coloured (apparently due to the sediment at the collect- creeks and streams with sandy (or other soft substrate) ing spots?), only the mouth and the tentacle are darker bottom; among smaller stones or bolders, as e.g. along pigmented. The mantle edge showed a total of 15 papil- the banks of the North Johnstone River in Queensland lae; with three of this papillae thickened on the left side (Figure 2h), outside the swift flowing currents. Like of the body, whereas on the right side there is an anal Thiara amarula and Stenomelania cf. aspirans the oc- papilla together with two smaller papillae adjacent to currences of R. queenslandica appears to range from the opening of the brood pouch. smaller tributary regions, as e.g. in the Douglas Creek of the Daintree River, to the lower courses of rivers as Radula (Figure 26). The taeniglossate radula is very in the Johnstone drainage. small and slender, with 110 to 130 rows of teeth (for n = 5 specimen). The rachidian has a main mesocon, Distribution. The geographic range of Ripalania queens- which is flanked by three smaller denticles on both landica is restricted to and within the Jardinian fluvi- sides. The lateralia show a prominent, elongated main faunal province, occurring isolated in few rivers and denticle (Figure 26b) with one to two smaller denticles streams along the tropical coast of NE Queensland attached on the inner side and mostly three denticles (Figure 27). It is found isolated in the Iron Range and outside (with the size of these denticles decreasing out- Lockhart River area, and again from the Daintree River wardly). The marginalia are in two pairs of parallel drainage south to Cardwell. rows, with 8–10 denticles of about the same size in both inner and outer teeth. Material examined Queensland: „Australia“ (ZMB, coll. Paetal 1885); Line Hill, Iron Reproductive system. The brood pouch resembles very Range (12450 S 143250 E) (QM 64464); Lockhart River, SW of air- much in position and extension that described above field, 0,5 km South Claudie River crossing (12480 S 143170 E) for the ovoviviparous Thiara amarula; it is also filled (QM 21407); Daintree River (16180 S 145170 E) (AMS C.317982, with many small, very early ontogenetic stages. These C.100354) (NHMB 10.751); Daintree River, above Allanton Hill

Figure 26. Radula of Ripalania queenslandica (Smith, 1882). a–d. QLD, North Johnstone River (ZMB 106355). a. Radula ribbon; scale bar = 100 mm; b. Lateral and central teeth; scale bar = 50 mm; c. Rachidian; scale bar = 10 mm; d. Marginal teeth; scale bar =10mm.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 239

ticular his figure 9) might be instructive. There are ac- tually two shells depicted by Smith, which presumably represent two quite different taxa. One (there are actu- ally two specimens extant: BMNH 1879.5.21.495-6) is quite similar in appearance to S. riqueti (see Fig- ure 31b) and is given with a location (“Cape Upstart, Queensland”) within the possible range of this species in Australia (see below, Figure 30). The second and lar- ger specimen (BMNH 1857.9.30.8) is from the Victoria River in Northern Territory and, in contrast, resembles more typical S. venustula (see Figure 31c). See below under the species for more comments. Diagnosis. Small, slender shell with up to 8 whorls and an evenly rounded last whorl. Main sculptural elements are the widely spaced axial, sometimes pronouncedly opistho- cyrt ribs on the upper part of all whorls, while on the basal part of the last whorl there are prominent spiral grooves. Distribution. The genus is distributed widely in the in- Figure 27. Geographic range of Ripalania queenslandica sular regions of Southeast Asia, the Indo-Malay Archi- (Smith, 1882) in Queensland, Australia. Note the restriction pelago ranging far into the Pacific region, including within the Jardinian fluvifaunal province. Australia (see Figure 29 for the type species).

(16140 S 145190 E) (AMS C.317986; C.125802) (QM 16272); Dain- tree River, nr Daintree (QM 13483); Douglas Creek, at crossing, nr Dain- Sermyla riqueti (Grateloup, 1840) tree (1616.194 S 14558.60 E) (ZMB 107213); Stewart Creek, at junc- Figures 1h, 28, 29, 30, 35c–d tion with Daintree River (16180 S 145190 E) (AMS C.317984); Low Isles, nr Port Douglas (AMS); Barron River (AMS C.109650); Cairns Melania riqueti Grateloup, 1840: Actes Soc. Linn. Bord. 6 (36): 433. (AMS C.1334) (ZMB 61751); near Innisfail (AMS C.51805); Innisfail, Melania tornatella Lea, I. L. & Lea, H. C. 1851: Proc. Zool. Soc. Blackfellow Creek, Flying Fish point (AMS); North Johnstone River Lond. 18: 185. (1730.34 S 14559.55 E) (ZMB 106355) (1730.34 S 14559.55 E) Melania sculpta Souleyet, 1852: Voy. de la Bonite: 546. (ZMB 107214); Tully River (AMS C.9282); Cardwell (AMS). Type locality. Grateloup (1840: 433) mentioned “Bom- bay” as locus typicus; however, this is inconsistent with the information on the label of the type material stud- Sermyla H. & A. Adams, 1854 ied, viz. “Batavia [1. line], Samarang [2. line]”, i.e. cor- Sermyla H. & A. Adams, 1854: in Gen. Rec. Moll. 1: 53 (non Sermy- responding to Jakarta, Java, today; a possible link to la Walker, 1854 (Lepidoptera), non Chapuis, 1875 (Coleoptera)). the Samarang Expedition is not yet clear. Sermylasma Iredale, 1943: Aust. Zool. 10 (2): 208. Type material. 2 syntypes (BMNH 1907.11.2240––1) Type species. Melania tornatella Lea, 1850, which is a (Figure 28a). junior synonym of Melania riqueti Grateloup, 1840; by original designation. Taxonomy. According to Fulton (1908: 43) the collection of J. P. S. Grateloup in Bordeaux “has recently been bro- Taxonomy. The genus Sermyla was considered a subge- ken up, and the types of the species enumerated are now nus of Melanoides e.g. by Thiele (1929), Rensch (1934), in the British Museum”. Although the pictures in Grate- and Benthem Jutting (1956), while Pace (1973) affiliated loup’s work are often not very similar to the real shells, it with Thiara. However, according to our preliminary his two figures given for S. riqueti are unambigous. molecular analyses (Glaubrecht et al. unpubl. data), Ser- Nevertheless, later authors have assigned many different myla clearly represents a distinct evolutionary lineage names for geographically isolated populations of Sermyla with at least the following two species discussed here. riqueti, which are here considered synonyms. Knowledge of this taxon is scarce to date. Nevertheless, from the Pliocene of Boemiajoe on Java (Indonesia) a fossil is described and depicted as oppenoorthi Oostingh, Description 1935, resembling very much specimens of Sermyla. Confusion exists not only with respect to the locus Shell (Figures 1h, 28). The shell with up to eight whorls typicus of all three Sermyla taxa discussed here below is compact, oval and relatively small, with a maximum (see under the species), but as to the identity of these of about 20 mm in specimens outside Australia (Glau- species, in particular the Australian taxa venustula and brecht unpubl. data), but consistently smaller in those carbonata. As an example for the existing irritations known from this continent (Table 1). The flexed axial the diagnosis of venustula by E. A. Smith (1882, in par- ribbs are opisthocyrt and very prominent, as are the

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 240 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 28. Shell morphology of Sermyla riqueti (Grateloup, 1840). a. Syntypes of riqueti (Grateloup, 1840); „Batavia“ (BMNH 1907.11.22.40-41); b. Indonesia: Bali, Gumbrih River (ZMB 106474); c. Indonesia: Sulawesi, Kupa (ZMB 191388); d. Aus- tralia: QLD, Caloundra (AMS C.3215); e. Australia: NT, Towns River (AMS C.317336). a–d, scale bar = 1 cm; e, scale bar = 0.5 cm. elevated spiral ridges and grooves basally on the last Radula (Figures 35c–d). The rachidian has denticles whorl. The colour varies from dark to lighter tones, but that taper conspicuously, a character also found in the mostly from black to dark brown. congeneric venustula. In addition, the main central den- ticles of the lateralia are broader than usually seen in External morphology. No pronounced differences were most other thiarids, flanked by two relatively large den- noted to those characters known on the general anato- ticles forming a very pronounced wing-shaped plate on my of other Thiaridae. the inner side projecting towards the central teeth.

Figure 29. Distribution of Sermyla riqueti (Grateloup, 1840), with occurrences outside Australia; identifications and locality infor- mation from Glaubrecht (unpubl. data).

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 241

Figure 30. Occurrences of Sermyla riqueti (Grateloup, 1840), in Australia. Note the occurrences in some rivers of the Leichhard- tian, Jardinian and in the Krefftian fluvifaunal provinces; for details and discussion see text.

Ecology. Originally, Grateloup (1840: 433) thought Ser- locations can not be distinguished from the type materi- myla riqueti to be marine: “Je la soupconne marine”. al of S. riqueti (or reference material from Indonesia). Apparently, the species is indeed restricted to the estu- Accordingly, S. riqueti appears to have, at least at aries of tropical rivers, under influence of the tides, times, entered the Victoria River (Timor Sea drainage) where it lives on muddy and sandy substrate and feeds as well as rivers in the Gulf of Carpentaria; nothing on deritus. more is known about the locations along the eastern coast. Note that this species is here recorded for the Distribution. Sermyla riqueti is distributed from India first time to occur in Australia. to Southeast Asia and into the Indo-West Pacific, in- cluding Thailand, the Philippines, and the Sunda Is- Material examined lands in Indonesia, with records also known from the Indonesia: Bali: Gumbrih River (ZMB 106474). Sulawesi: Kupa Bismarck Archipelago and the Solomon Islands (Glau- (ZMB 191388). brecht unpubl. data); see Figure 29. Australia: Northern Territory: ? (BMNH 1857.9.30.16); Victoria For Australia mostly only few dry shells are available River (BMNH 1857.4.18.131); Towns River, E of Mataranka, below from museum collections, thus evidence for the regular road crossing (1442.940 S 13430.440 E) (AMS C.317336; stn NT occurrence of this species remains to be verified. To 15, 5781; note that these coordinates as given refer to Port Roper date, Sermyla riqueti is reported (often with only “his- instead). Queensland: Norman River, Billabong 1 km N of Normanton torical” records) from very few locations at rivers in (1739.712 S 14106.154 E) (ZMB 107209); Lizard Island (BMNH the Leichhardtian and Jardinian as well as isolated even 1854.10.28.32); Kapoh Cove Beach, North Point (AMS); Buchans in the Krefftian fluvifaunal provinces, as listed below Point, N of Cairns (AMS); Port Curtis (AMS); Caloundra, N Bris- and depicted in Figure 30. The shells from these few bane, brackish water from Two Way Creek (AMS C.3215).

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 242 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Sermyla venustula (Brot, 1877) Melania carbonata Reeve, 1859: Con. Icon. 12: pl. 13, fig. 88. (2 syn- Figures 1j, 2b–e, 31, 32, 33, 34, 35, 36 types BMNH 2001.0762; from “Port Essington” (?); see Fig- ure 32c and text below). New synonymy. Sermylasma carbonata – Iredale, 1943: Aust. Zool. 10: 209. Melania venustula Brot, 1877: in Martini & Chemnitz, Conch. Sermylasma prognata Iredale, 1943: Aust. Zool. 10: 208. (unnecessary Cab. 1 (24): 331. nom. nov. for Melania venustula) Melania venustula – Smith, E. A. 1882: J. Linn. Soc. Lond., Zool. 16: 261. Type material. 6 syntypes (MHNG, no number) (Fig- Sermylasma venustula – Iredale, 1943: Aust. Zool. 10: 208. ure 31a).

Figure 31. Shell morphology of Sermyla venustula (Brot, 1877). a. Syntypen von Sermyla venustula (MHNG); originally given as “Port Denison”, see text for discussion. b–c. Specimens on which E. A. Smith (1882) based his description and differentiation of two morphs in Sermyla venustula; b. “Cape Upstart” (leg. Brazier; BMNH 1879.5.21.495–496); c. NT, “Victoria River” (leg. J. R. Elsey; BMNH 1857.9.30.8); d. QLD, Norman River (ZMB 106713); e. QLD, Bynoe River (ZMB 106712); f. NT, Little Roper River (ZMB 106629); g. NT, Salt Creek (ZMB 106632). Scale bar = 1 cm.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 243

Type locality. “Port Denison, Nov. Holl. [Novae Hollan- Figure 32) indicate a close affinity to those here as- dia]”. This location as given by Brot (1877) and also signed to Sermyla venustula (Figure 31). B. J. Smith (1992: 77), which is today Bowen, S Towns- Diagnosis. In comparison to S. riqueti (Figure 28) the ville in Queensland, is most probably erroneous, as this shells of S. venustula (Figure 31) are much taller, highly species is not known to occur at all in the Jardinian turreted and slim; they are often, however, with less province (see below). distinct axial and spiral elements (as is in particular the Taxonomy. In the first diagnosis Brot (1877: 331) men- case in carbonata). The opisthocyrte axial ribs vary in tioned only four specimens; however, we found that the their flexing and the intervals between them. These ax- type series of the MHNG actually consists of six shells. ial ribs are, nevertheless, the dominant element already We also synonymize here carbonata Reeve, 1859, origin- in the juveniles. The whorls are not or only weakly ally described as separate species, unfortunately, without stepped, as compared e.g. to “Thiara” australis and information on the type locality. The types were assumed their juveniles. to be lost by B. J. Smith (1992: 78), but are actually in the BMNH (see Figure 32c). Only subsequently, Brot Description (1877) and E. A. Smith (1882) mentioned “Port Essing- ton” in NT as locality for this taxon, followed later by Shell (Figures 31 & 32). The polymorphic shell is slen- B. J. Smith (1992: 78). It is important to note that this der, highly turreted, with up to ten whorls, albeit the locality on Cobourg Peninsula is neither known for oc- apical tip of the adults is often slightly corroded. The currences of carbonata nor of venustula (see below); height of the shell reaches well over 30 mm (Table 1); however, it was certainly the only port and settlement shells have a black to brown colour, sometimes with used at the early times of exploration of northern Austra- spots and other lighter elements. The axial ribs are lia and possibly the only known locality of any sort for opisthocyrt, although we found that samples from the A. Brot and E. A. Smith at their time. It is also note- Little Roper River consistently have orthocline ribs (see worthy, that the nearest known occurrence of any Sermy- Figure 31f) as an exception. The basal part of the last la species today is at Howard Springs, as revealed below. whorl shows the typical spiral ridges and grooves, re- We also synonymize Iredale’s (1943) new genus Ser- spectively. The aperture is oval. mylasma, designated with venustula as type species, and to which he assigned five other species, among Juvenile shell (Figures 33 & 34). The shells of the ju- them carbonata Reeve, 1859 (see Figure 32c). As these veniles are characterised by very clearly developed ax- names were not found to represent distinct taxa and ial ribs, widely spaced and crossed by weak spiral ele- lack sufficient characters to justify separation, they are ments. There are up to five convex, well rounded synonymised here under venustula, further supported whorls which are stepped due to subsutural angulation by the results of our molecular analyses (Glaubrecht only occasionally. The sculpture of the first whorl is on et al. unpubl. data). In particular, samples from Howard the initial cap wrinkled (Figure 33d) to more or less Springs resembling carbonata in their shell (compare smooth (Figure 33h), with more or less clearly visible

Figure 32. Shell morphology of exceptionally large specimens of Sermyla venustula from Howard Springs, near Darwin, which have been named “Melania” carbonata (Reeve, 1859). a. NT, Howard Springs Creek (ZMB 106595); b. NT, Howard Springs (ZMB 106700); c. Syntypes of Melania carbonata (BMNH 2001.0762), from “Port Essington” (?); see text for details. Scale bar = 1 cm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 244 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 33. Juvenile shells from two po- pulations of Sermyla venustula (Brot, 1877), showing lateral view, apical whorls from lateral, apical view, and de- tails of the protoconch. a–d. NT, Little Roper River (ZMB 106629). e–h. NT, Salt Creek (ZMB 106632). Scale bar = 200 mm. commencement of growth lines that become increas- la from Little Roper River and Salt Creek, respectively, ingly prominent. as well as Figure 34 for those of “carbonata”-like spe- cimens from Howard Springs. External morphology. There are no clear differences found to other thiarids studied. Ecology. Like riqueti also venustula is apparently oc- curing in brackish water habitats, as it is known not Radula (Figures 35a–b). The radula corresponds to only from lower courses of rivers and streams, but also those typically known from other Thiaridae; however, further inland in the Northern Territory at localities the rachidians consistently appear to be more cuspid with partially saline conditions. Some areas of the Ro- with their sharply pointed denticles. per River drainage system, for example, at Salt Creek (see Figures 2d–e), are almost marine-like in their salt Reproductive system. The brood pouch is filled with content dissolved from sedimentary rocks (see Discus- about a dozen juveniles with shells, similar to the mode sion). It is not yet studied what kind of ecological cir- reported above e.g. for “Thiara” australis; compare also cumstances are relevant for the occurrence of Sermyla Figure 33 for those shelled juveniles in typical venustu- at Howard Springs.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 245

Figure 34. Juvenile shells of “carbona- ta”-like specimens of Sermyla venustula (Brot, 1877), showing lateral view, api- cal whorls from lateral, apical view, and details of the protoconch. a–d. NT, Ho- ward Springs (ZMB 106593). e–h. NT, Howard Springs (ZMB 106700). Scale bar = 200 mm.

Distribution. The geographic range of venustula ap- Note that the distribution given by B. J. Smith (1992: pears to be restricted in northern Australia to the cen- 77) as “NE coastal, QLD”, is not correct, as compared tral Leichhardtian fluvifaunal province, essentially with to our survey and reconstruction presented here. occurrences in the Gulf of Carpentaria drainage (Fig- ure 36). Remarkable, however, are the location of ex- Material examined ceptionally large and less sculptured specimens, re- ferred here to as “carbonata” (Reeve, 1859), at Howard Northern Territory: Howard Springs (1227.345 S 13103.146 E) Springs and its run-off creek near Darwin, isolated in (ZMB 106593, 106700); Howard Springs Creek (1227.268 S northwestern Northern Territory. Thus, while S. venus- 13103.108 E) (ZMB 106595, 107228); Little Roper River, at crossing tula is not found in the Daly River (i.e. Timor Sea drai- (1455.581 S 13307.176 E) (ZMB 106629, 106678, 107236, nage), all records presented here comprise, next to sev- 107237); Roper River, at Jalmurark Camp Ground (14 57.158 S eral rivers along the southern Gulf coast, in particular 133 13.29 E) (ZMB 107229); Elsey Creek on Roper Highway (1500.627 S 13314.417 E) (ZMB 107231); Salt Creek, nr Elsey those from the Roper River drainage, including those Creek, at crossing of Roper Highway (150.703 S 13314.417 E) from its tributaries Little Roper and Waterhouse River (ZMB 106632, 107230); Warloch Ponds on Elsey Creek, near Old as well as Salt Creek. Stuart Hwy, Elsey Station, Mataranka Area (1605.042 S

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 246 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 35. Comparison of the radula of species of Sermyla. a–b. Sermyla venustula from Australia: NT, Howard Springs (ZMB 106700). a. Lateral and central teeth; scale bar = 50 mm; b. Marginal teeth; scale bar = 10 mm; c–d. Sermyla riqueti from Indonesia: Bali, Kupa (ZMB 191388); c. Lateral and central teeth; scale bar = 25 mm; d. Marginal teeth; scale bar = 10 mm.

Figure 36. Geographic range of Sermyla venustula (Brot, 1877) in northern Australia with occurrences in the central Leichhardtian fluvifaunal province, essentially restricted to the Gulf of Carpentaria drainage; however, note the location of the exceptionally large specimens, referred here to as “Melania” carbonata (Reeve, 1859), at Howard Springs near Darwin, isolated in northwestern Northern Territory.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 247

13307.258 E) (ZMB 192019); Roper Bar, Roper River (1442.816 S is often the case). This species is the tallest of all Aus- 13430.501 E) (ZMB 192017); Mumpumapu waterhole, Phelp River tralian thiarids. The shell is solid, with only slightly drainage, Numbulwar-Roper rd., Mumpumumpu Outstation, Arnhem rounded whorls, and of black colour. Land (1422.59 S 13519.34 E) (AMS C.461353); 8 km NE of Towns River Crossing (1459.82 S 13516.28 E) (ZMB 192016, 192018); Foelsche River (1612.628 S 13653.034 E) (ZMB 107232); Kangaroo Creek (1647.553 S 13706.107 E) (ZMB 107233). Description Queensland: Gregory River at Riversleigh (1901.116 S 13843.529 E) (ZMB 107234); Bynoe River (ZMB 106712); 4.5 km NW of Normanton, Shell (Figures 1k, 37). The shell is turriform, very solid, road from Normanton to Karumba (saline ponds) (1739.43 S, 141 06.03 smooth; mostly corroded, with adults often possessing E) (VK 26.356) (ZMB 106713); Norman River, billabong 1 km N of Nor- only the last three to four whorls, while younger indivi- manton (1739.712 S 14106.154 E) (ZMB 107235). duals show up to seven whorls. The shell is mainly black with a tendency to appear greyish or blueish. The Stenomelania Fischer, 1885 last whorl is more than twice as large as the penulti- Stenomelania Fischer, 1885: Man. Conch.: 701 mate one. The apical part of the adult shell is often slightly angulated or concavely invaginated, which is Type species. Melania aspirans Hinds, 1844; by mono- also reflected in a bay-like impression of the upper typy. margin of the otherwise holostome and drop-shaped Taxonomy. To date, Stenomelania was essentially con- aperture. sidered as subgenus of Melanoides (see e.g. Morrison External morphology. The anatomy will be given in 1954; Smith 1992). In contrast, in our present generic more detail elsewhere, but was not found to be signifi- concept of Stenomelania we tentatively combine thiar- cantly different from the outer appearance of other ids with comparatively large, slender, highly turreted Australian Thiaridae. and smooth shell. A revision of this taxon is wanted and timely, as this group is certainly polymorphic and, Radula (Figure 38). The radula is extraordinary large as we assume from our ongoing studies on the constitu- and slender, as can be expected from the considerable ent species, also polyphyletic. Given these phylogenetic size of the body of this Stenomelania. The rachidian is and systematic uncertainties we here refrain from a for- characterised by the almost perfectly parallel upper und mal characterisation of the genus for the time being, basal margins, which was not found in other thiarids so but restrict our description to the species known now as far. This tooth is about three times broader than high, new to the Australian continent. not curved, with three denticles of about same size on both sides of the mesocone; in addition, there is some- times a relatively small lateral denticle at the outside, Stenomelania cf. aspirans (Hinds, 1844) flexed upwards (see Figure 38c). The lateralia are also Figures 1k, 2g, 37, 38, 39 quite straight at their upper margin. The inner and outer marginalia were found to have a varying number of Melania aspirans Hinds, 1844: Ann. Mag. Nat. Hist. XIV: 8. Melanoides (Stenomelania) aspirans – Wenz, 1938: Handb. Palo- denticles, mostly between 6–8. zool., Gastropoda, 1: 714. Reproductive system. The brood pouch of Stenomelania Type locality. “in rivers of Feejee Islands”; Fiji. cf. aspirans was found to be filled with veliger stages, all without shell, but with operculum and partially with Type material. 2 syntypes (BMNH 1844.9.23.32–31); cilia. The number of these early embryonic stages are three more possible syntypes from the Hinds Collection about several hundreds at least, in different phases of with the same locality (BMNH 2001.0760) (Fig- their development; they are embedded loosely within ures 37a–b). the tissue of the marsupium. Taxonomy. As the concept of Stenomelania is vage yet, Ecology. Stenomelania cf. aspirans is, like Thiara ama- we only tentatively assign the material described here to rula, living in various areas of the courses of tropical riv- the type species aspirans. This latter species is found to ers, but clearly above the area of influence of brackish be somewhat variable and should therefore also be re- water. The specimens known now from northern Austra- vised systematically. Moreover, we note that the name lia were found by the authors in the headwaters of the aspirans Hinds, 1844, is not the oldest valid and avail- Mowbray River (Figure 2g) in northern Queensland or able name for taxa resembling aspirans, as e.g. from the in small tributaries of the Bloomfield River, where they Astrolabe Expedition to the Indo-West Pacific there are live on substrate with mud, but also small pebbles. many similar forms described and depicted (Glaubrecht Some specimens buried themselves into the sand. The unpubl. data). Here we focus on the description on those flow velocity of the water at those places was low, shells from Australia that we found conchologically most nearly not existent (Mowbray), but in another place high similar to aspirans, and which we, therefore, here tenta- with swift currents of the running creek (Bloomfield). tively assign to the type species of Stenomelania. The species, albeit very large, is camouflaged very Diagnosis. The shell is high-spired and in fully-grown good, not easy to detect or to find at all, and apparently adults very tall, with up to 80 mm (if not decollated, as not present at any place in high abundance.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 248 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 37. Shell morphology of Stenomelania cf. aspirans (Hinds, 1844). a. Syntypes (BMNH 1844.9.23.32-31); “Feejee Islands”; b. Possible syntypes from Fiji in the Hinds Collection; (BMNH 2001.0760); c. Fiji, Sovi River (ZMB 106397); d. Australia: QLD, Mowbray River (ZMB 106344); e. Australia: QLD, Mowbray River (ZMB 106171). Scale bar = 1 cm.

Distribution. Stenomelania aspirans, as reported, is ap- The occurrence of Stenomelania cf. aspirans in Aus- parently widespread in the Australasian region, with tralia is restricted to the Jardinian fluvifaunal province, known occurrences, for example, in the Bismark Archi- where it only occurs in few streams along the tropical pelago, Solomon Islands, Vanuatu, New Caledonia, Fiji coast of Queensland (Figure 39). It is reported so far und Samoa. The distribution of the genus is even wider, only from highly isolated locations in the Iron Range albeit its limits not yet determined due to the lack of and Lockhart River area, the Bloomfield to Barron Riv- systematic knowledge about this taxon. As a systematic er region, and at Clump Point in the south. revision of its constituent species is lacking to date, Note that this species was hitherto not listed in any biogeographic information remains vage. faunal survey of Australian freshwater molluscs (e.g.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 249

Figure 38. Radula of Stenomelania cf. aspirans (Hinds, 1844); QLD, Mowbray River (ZMB 106344). a. Radula ribbon; scale bar = 200 mm; b. Lateral and central teeth; scale bar = 100 mm; c. Rachidian; scale bar = 50 mm; d. Marginal teeth; scale bar = 50 mm.

E. A. Smith 1882; Iredale 1943; B. J. Smith 1992, Material examined 1996), and is therefore here recorded for the first time Queensland. “Cape York Peninsula” (AMS C.109655); Rocky River to occur in Australia. (13490 S 145280 E; note that the coordinated given are out in the Coral Sea) (AMS C.317983); creek into West Claudie River, Iron Range (12470 S 143190 E) (AMS C.317327); Granite Creek, W of Bloomfield (1555.99 S 14519.54 E) (ZMB 107211); Mowbray River (1633.87 S 14527.83 E) (ZMB 106171, 106344, ZMB 107212) (AMS C.115338); Barron River, N of Cairns (AMS C.105172); Hart- ley’s Creek, N of Cairns (AMS C.109654); Froma Creek, N of Cairns (AMS C.158276); Clump Point (AMS C.30757).

“Stenomelania” denisoniensis (Brot, 1877) Figures 1m, 40, 41, 42, 43

Melania denisoniensis Brot, 1877: in Martini & Chemnitz, Conch. Cab. 1 (24): 234. Melanoides (Stenomelania) denisoniensis – B. J. Smith, 1992: Zool. Cat. Aus. 8.: 76. Stenomelania denisoniensis tacita Iredale, 1943: Aust. Zool. 10: 208 (4 syntypes AMS C.100606; from Cardwell, QLD; see Smith 1992: 77). [not seen] Melanoides (Stenomelania) tacita – B. J. Smith, 1992: Zool. Cat. Aus. 8.: 76. Stenomelania denisoniensis ultra Iredale, 1943: Aust. Zool. 10: 209 (2 syntypes AMS C.100608; from Clarence River, NSW; see Smith Figure 39. Geographic range of Stenomelania cf. aspirans 1992: 77). [not seen] (Hinds, 1844) in Queensland, Australia. Note the restriction Melanoides (Stenomelania) ultra – B. J. Smith, 1992: Zool. Cat. within the Jardinian fluvifaunal province. Aus. 8.: 77.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 250 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 40. Shell morphology of “Stenomelania” denisoniensis (Brot, 1877) in comparison with type material of Melanoides tuber- culata (O. F. Mller, 1774). a. Three syntypes of Melanoides tuberculata (ZMUC); b–g. “Stenomelania” denisoniensis; b. Syntypes (MHNG); “Queensland, Port Denison”; c. QLD, Alice River (ZMB 104145); d. QLD, Meelele River (ZMB 106341); e. QLD, Woo- badda River (ZMB 106342); f. NT, Rum Jungle (ZMB 106662); g. NT, Black Jungle (ZMB 106644). Scale bar = 1 cm.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 251

Type locality. “Port Denison”, Queensland, as given ori- lia”, that originate from the collection of Brot. How- ginally by Brot (1877); this corresponds today to Bo- ever, they are here not considered to represent syntypes, wen, S Townsville. but topotypical material only instead. Very early on the enormeous conchological variability Type material. 5 syntypes (MHNG, no number), all de- in “Stenomelania” denisoniensis, in terms of size of shell, picted in Figure 40b; see more below under taxonomy. sculpture and colour, was recognized, according to com- Taxonomy. Brot (1877) explicitly noted the number of ments in Brot (1877) and E. A. Smith (1882). However, specimens to be six (“Ich besitze sechs Exemplare die- this hasn’t stopped Iredale (1943) from naming two geo- ser Art”). Nevertheless, we only found one lot of this graphical variants – notably both from the east coast of taxon containing five syntypes in the MHNG, all glued Australia (see above) – as subspecies, viz. (i) ultra with on cardboard (as was the usual procedure with Brot’s broader and larger shells, but less convex whorls than ty- material). On the cardboard there is also space for a pical denisonensis, and (ii) tacita resembling those speci- sixth specimen, with remnants of glue left, but this last mens depicted in E. A. Smith (1882: fig. 5). Both were syntype missing. The MCZ owns two more shells raised even to species status later by B. J. Smith (1992), (MCZ 110243), also from “Port Dension, Qld, Austra- and the names also used by Stoddart (1985) for specimens

Figure 41. Juvenile shells of “Stenomela- nia” denisoniensis (Brot, 1877), showing lateral view, apical whorls from lateral, apical view, and details of the protoconch. a–d. NT, Towns River (ZMB 106641). e– h. QLD, Gregory Falls (ZMB 106352). Scale bar = 200 mm.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 252 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae from Queensland (tacita) and from Western Australia (ul- are visible. The colour is from light brown to yellowish tra), albeit without depicting or describing these two green up to dark brown tones; especially younger speci- forms and their differences more explicitly. mens show lighter colours, many exhibit flammulated Note that conchologically this species can not be dis- spots. The aperture is wide oval and rarely flaring on tinctly separated in many instances from Melanoides tu- the basis. Some specimens are pronouncedly angulated berculata; however, we found it in our molecular ana- by a subsutural step, which lends these shells a mark- lyses to be clearly separated from the latter taxon as edly different appearance (Figure 40d). However, these well as other Australian thiarids, so we consider deniso- are usually found in all possible intermediate forms niensis here as an independent lineage of own evolu- with normal phenotypes as described above, and appar- tionary history. In case of verification and thus being ently occur without clear geographical patterns. necessary later, we suggest a new generic affiliation for this species. Embyonic shell (Figure 41). The juveniles have four to five whorls and show a highly characteristic reticulate Diagnosis. The shell is slender and turriform, with up sculpture (“Gitterstruktur”) that it build by densely to nine convex whorls and simple suturs. The sculpture spaced intersecting axial and spiral ribs, whereas the is often smooth, without a clearly pronounced reticulate spiral ridges dominate the axial ribs. Only the first one pattern, but mostly with at least some pronounced spir- and a half whorls are without this sculpture, but show al grooves. The colour is more often brownish than the usual wrinkled pattern of euviviparous thiarids (see black. The aperture is wide oval and holostom. Glaubrecht 1996). External morphology. The mantle edge possesses usually Description 10–14 papillae. On the right side of the body there are four papillae which are more pronounced and larger, while Shell (Figure 40). The shell is elongate and slender with on the left hand side there is only one large (anal) papilla. up to nine convex whorls. The whorls are corroded only in the upper regions. The suture is simple. The surface Radula (Figure 42). Tha radula has up to 150 rows of of the shell is almost smooth, only a few spiral lines teeth with rachidian, lateralia and marginalia showing

Figure 42. Radula of “Stenomelania” denisoniensis (Brot, 1877). a–b. NT, Towns River (ZMB 106641). a. Lateral and central teeth; scale bar = 50 mm; b. Marginal teeth; scale bar = 10 mm; c–d. NT, Howard River (ZMB 106597); c. Lateral and central teeth; scale bar = 50 mm; d. Marginal teeth; scale bar ¼ 10 mm.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 253 the typical composition. The number of denticles of in- (northern) South East Coast fluvifaunal provinces. dividual teeth varies within the radula of individuals, While its main occurrences are in the coastal rivers of populations and geographical area. Usually, the margin- Queensland draining east to the Coral Sea, extending alia have about nine denticles. within the Jardinian and adjacent Krefftian province Reproductive system. The subhemocoelic brood pouch south right across the border into New South Wales, it in denisoniensis is filled with about a dozen juveniles reaches its northern most known occurrence in the Iron of different size but all with shells, plus many juveniles Range area. There is one notable isolated location re- without shells in different embryonic stages. The ported from Soda Gorge Spring, NE of Hughenden, shelled juveniles were found to possess about two to a which is the tributary region of the Flinders River maximum of three whorls. draining north into the Gulf. The species occurs in riv- ers along the southern coast of the Gulf of Carpentaria, Ecology. The species lives in lotic and lentic freshwater in many streams and rivers of the Northern Territory habitats; it feeds on detritus and algae. We found deni- including the Daly and Roper River systems, as well as soniensis exclusively in rivers and streams outside of in the Victoria River. In addition, there are a few re- the influence of brackish water. cords from isolated populations in the Kimberleys, the Distribution. “Stenomelania” denisoniensis is widely Pilbara and one single record in the SW of Western distributed along the rivers of the northern part of the Australia, where we found it in July 2004 in Ellendale Australian continent (Figure 43), with occurrences rang- Pool, south of Geraldton at the Greenough River. Re- ing from the Greyian (Pilbara) across the Leichhardtian markably (as compared e.g. to Plotiopsis balonnensis, with Timor Sea and Gulf drainages to the North and see Figure 24), this otherwise widely distributed species

Figure 43. Geographic range of “Stenomelania” denisoniensis (Brot, 1877) in Australia. Note the occurrences in the Indian Ocean or Greyian (Pilbara) province, including an isolated occurrence at its southern most limit at Greenough River, as well as the Leichhardtian, Northeast Coast and (northern) Southeast Coast fluvifaunal provinces. It lacks in the Sturtian and Mitchellian pro- vinces (as compared e.g. to Plotiopsis balonnensis, see Fig. 24).

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 254 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae lack in the Sturtian and in particular the Mitchellian 13506.20 E) (NTM P8702); Wumaitparr waterhole, N of Numbulwar, provinces, thus the entire Murray-Darling river system. Wumajbarr Outstation, Arnhem Land (1409.070 S 13542.040 E) (AMS C.461372); Mumpumapu waterhole, Phelp River drainage, Numbulwar-Roper rd., Mumpumumpu Outstation, Arnhem Land Material examined (1422.590 S 13519.340 E) (AMS C.461371); Wandoo River, Numbul- 0 0 Western Australia: SE of Geraldton, Ellendale Pool at Greenough war-Roper River rd., Arnhem Land (14 14.02 S 135 36.11 E) River (2851.630 S 11458.430 E) (ZMB 106586); Pilbara Springs, S (AMS C.461367); Wandoo River, Numbulwar-Roper River rd., Arn- 0 0 of Dampier (205101500 S 1163605000 E) (AMS); Pilbara Springs, hem Land (14 14.02 S 135 36.11 E) (AMS C.461354); Towns River, pool of spring (21370 S 1170602000 E) (AMS); Millstream, Mill- at crossing (15 02.570 S 135 12.718 E) (ZMB 106641, 107239); 0 0 stream NP, bottom end of lily pond (2135.50 S 11840 E) Wearyan River crossing (in muddy pools) (16 10.05 S 136 45.25 E) 0 0 (AMS C.324141); Pardoo Station, Banningarra Spring, NE of Port (VK 13.870); Calvert River crossing (16 56.06 S 137 21.25 E) Hedland (20020 S 199410 E) (AMS stn P27); 150 km E Nullagine (VK 25.837, 25.846); Calvert River, below junction with Bluey Creek (running waters) (214004500 S 121080 E) (AMS C.324137); Kimber- (1656.10 S 13721.520 E) (AMS C.151987); Cox River, N of cause- leys: Mt. Mathew, Mt. Hart Station, King Leopold Range (in a small way (in stagnant waterhole) (1519.394 S 13520.669 E) stream at the base of the hill) (VK 12387). (ZMB 107240). Northern Territory: “North Australia” (BMNH 1857.9.30.18); Queensland: running creek, 27.7 km Brookdale (1819.250 S Black Point Lagoon, Smith Point, Cobourg Peninsula (AMS); lagoon 13915.450 E) (VK 13.872); Lawn Hill Creek, nr the cascades, Lawn behind Ranger Station at Smith Point, Cobourg Peninsula (open water, Hill NP (1842.000 S 13829.000 E) (VK 26.335); Gregory River, Riv- weedy edge) (AMS); Holmes Creek, 12 miles NE of Darwin (AMS); ersleigh Stn (QM 64453); Gregory River crossing, Riversleigh Station Darwin River, at crossing with highway to Cox Peninsula (1244.527 (1901.150 S 13843.250 E) (VK 13.873); Leichhardt River, east S 13057.930 E) (ZMB 106704); Howard Springs, S Darwin branch 2 km below dam (20440 S 139470 E) (AMS C.300838); (1227.50 S 13103.00 E) (NTM P1013, P27464, P27466) Bynoe River crossing, Burketown to Normanton road (1751.530 S (AMS C.110450) (QM 5132); Howard River, crossing (1227.752 S 14047.580 E&1751.710 S 14048.060 E) (AMS C.338669) 13105.008 E) (ZMB 106597); Berry Springs, S of Darwin (VK 26.336); Normanton River, Glenmore, SE of Normanton River (1242.111 S 13059.854 E) (ZMB 106600, 107255) (QM 5624); (1551.199 S 14108.048 E) (ZMB 107242); Glenore crossing, Nor- Foggy Dam, Humpty Doo (AMS C.110490) (VK 2523); Middle Point manton River, 21 km SSE of Normanton (1751.210 S 14108.000 E) jungle, near Fogg Dam (VK 7724); Manton River, Weed Quad 3 (VK 26.331); Walker Creek Crossing on Normanton, Karumba road (NTM P27465); Manton River at Stuart Highway, 66 km S of Darwin (1728.220 S 14110.420 E) (VK 26.338); billabong, 1 km from (AMS); Coomalie Creek, at road crossing (1300.602 S 13106.850 Norman River, on Kurumba road (1739.640 S 1416.10 E) E) (ZMB 106645); Coomalie Creek, at rest area on Stuart highway (AMS C.338659); Norman River, billabong 1 km N of Normanton 0 0 (142 km) (13 0.88 S 131 7.4 E) (AMS C. 324138) (VK 973) (NTM (1739.712 S 14106.154 E) (ZMB 107241); Mt. Isa, Lake Moondar- P6468); Crater Lake, 7 km W of Batchelor (13 02.760 S 131 05.445 ra [Leichhardt River drainage] (QM 64474); Soda Gorge Spring, NE E) (ZMB 106660) (VK 24531); Adelaide River (BMNH of Hughenden (2037.000 S 14405.330 E) (AMS C.145015); Line 1891.11.21.151-152; 1892.1.29.193); Rum Jungle at Litchfield Road Hill, Iron Range (12450 S 143250 E) (QM 64470); Lockhart River, (13 02.604 S 130 59.862 E) (ZMB 106662); Black Jungle Spring, SW of airfield, 0,5 km S Claudie River crossing (12480 S 143170 E) Kakadu NP (spring and stream on floodplains, rainforest on black (QM 21408); 80 km N Cooktown, McIvor River (150.90 S 145060 0 0 soil, on mud) (13 02.898 S 132 09.889 E) (ZMB 106644) E) (AMS); Endeavour River Falls (1522.2700 S 14501.7700 E) (AMS C.32413) (VK 25909); Bamboo Creek at the junction with (ZMB 106338, 107244); McLeod Creek, at crossing, tributary of En- Daly River (1340.050 S 13039.290 E) (VK 24378); Daly River deavour River (1525.505 S 14506.049 E) (ZMB 107243); Ende- Crossing (0–5 m, roots along side of the river, not in max. flow but avour River, 12 miles WNW of Cooktown (AMS); Cooktown water still strongly flowing) (1346.020 S 13042.610 E) (AMS C.437129); Laura River (1534.680 S 14427.410 E) (AMS C.323809); Victoria River (BMNH 1857.9.30.17); Upper Vic- (ZMB 106339); Boggy Creek, W Normanby tributary (1549.97 S toria River, North Australia (17180 S) (BMNH 1857.11.18.26); Vic- 14452.910 E) (ZMB 106373); Three Mile Creek/Poison Creek, tribu- toria River, 5 km downstream from Victoria Highway bridge, 195 km tary of Endeavour River (1525.789 S 14507.04 E) (ZMB 106356, W of Katherine (AMS C.324142); Roper River, old bridge crossing at 107245); Bloomfield River (AMS C.487); Granite Creek, W of Mataranka Homestead (1455.50 S 1336.50 E) (AMS C.317320); Lit- Bloomfield (1555.99 S 14519.54 E) (ZMB 107246); Bloomfield tle Roper River, at crossing (1455.581 S 13307.176 E) River crossing, 2.9 km on N side (15560 S 145200 E) (QM 24004); (ZMB 107253); Roper River, just below crossing at Mataranka Woobadda Creek (1557.969 S 14522.858 E) (ZMB 107247); Woo- (14560 S 13370 E) (AMS C.151987); Stevie’s Hole at Waterhouse badda River (1558 S 14522.480 E) (ZMB 106342); Meelele River River, Elsey NP (1455.782 S 13308.732 E) (ZMB 107254); Water- house River, Mantaranka tourist resort (QM 5156); Roper River, (15 58.250 S 145 23.850 E) (ZMB 106341, 107248); Wonga Beach, 0 0 Roper Falls, 4 km E of Jamurak (1457.401 S 13315.018 E) 14 km NNE of Mossman (16 20.30 S 145 25.00 E) (VK 21.054); (ZMB 107252); Mataranka Falls, Roper River, Elsey Park (1457.300 Mossman River, creek at crossing (ZMB 104150); creek entering S 13315.000 E) (VK 24.377); Elsey Cemetery, 11 km of Mataranka Mossman River, at Mossman Gorge (AMS C.127104, C.426370); Springs (155.150 S 1337.440 E) (AMS); Elsey River, Elsey Ceme- Mossman River, outside Mossman Gorge NP (AMS); Mossman River tery (ZMB 106651); Salt Creek, nr Elsey Creek, at crossing of Roper Gorge, small creek nr parking area (AMS stn 53a); Mowbray River, Highway (1500.703 S 13314.417 E) (ZMB 106685, 107238); War- nr Port Douglas (AMS C.324143); W side of Daintree River Valley, loch Ponds on Elsey Creek, nr old Stuart Hwy, Elsey Station nr head of small creek off forestry road (AMS); Kewarra Beach, N (1605.042 S 13307.258 E) (ZMB 190197); Roper River at Roper of Cairns (AMS); Barron River (BMNH 1885.3.18.3–6) Bar camping ground (1442.71 S 13430.78 E) (AMS C.338661, (AMS C.127065); creek off Mulgrave River, N Cairns (17140 S C.338657); Goyder River, Arnhem Land (AMS); eastern Goyder 145460 E) (AMS); Mulgrave River, near Goldsboroug, Cairns (QM River crossing, Central Arnhem Land (1301.370 S 13458.370 E) 64451); Mulgrave River, nr Cairns (AMS); Barron Falls (AMS C.461369); Gove Peninsula, SE of Nhulunbuy, Arnhem Land (AMS C.9283); Stony Creek, nr Barron River (QM 4991); Barron (1214.560 S 13653.170 E) (AMS C.461356); Gove Peninsula, Nhu- River, Hemmings Rd. (QM 64503); Barron River, Picnic Crossing lunbuy Arnhem Land (lagoon) (1210.520 S 13647.060 E) (QM 64478); Barron River Falls (AMS C.51468); Barron River, (AMS C.461350); Rose River catchment, southern end of Parsons George Road, half way to hydro station at river access (1651.632 S Ranges, Numbulwar District, 150 km N of Roper Bar (1343.400 S 14539.791 E) (ZMB 107250); Barron River, below 150 m Lake Pla-

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 255 cid (1652.17 S 14540.405 E) (ZMB 107249); spring beside Barron Part II: Variation in morphological River, 12 miles N of Atherton (AMS C.324144); Crystal Cascades, Redlynch, nr Cairns (QM 64476); Yarrabah Rd., Pine Creek (17000 S characters 145500 E) (QM 48184); Pine Creek, NW of Malbon Thompson Rd., Cairns (AMS stn 44); Musgrave River, 3 km downstream from Gold- The conchology in particular of freshwater snails has sborough Bridge (1711.400 S 14544.130 E) (VK 26.346); tributary been found to be highly variable (see e.g. discussion in of Musgrave River, nr Gouldsborough Valley Camping Area Rensch 1934; Glaubrecht 1996, 2004, 2009, and litera- (1714.190 S 14546.280 E) (VK 26.333); Dowah Creek, at Freshwater Creek junction, W Cairns (AMS C.324140); Pelican Creek (BMNH ture therein). Nevertheless, in the past features of the 1884.12.27.1-7); nr Gregory Falls (1735.5700 S 14552.2900 E) shell have figured most prominently in species diag- (ZMB 106352); Tinaroo Dam & Lake Tinaroo, Atherton Tableland noses, although this often ignores or at least underesti- (AMS C.158125, C.158280); Lake Tinaroo (AMS C.158280); Cham- mates the amount of intraspecific variability due to, bri Lakes, Atherton Tableland, Lake Eacham (17170 S 145370 E) for example, effects such as ecophenotypy and/or geo- (QM 14002); Atherton Tableland, Lake Tinaroo (17100 S 145330 E) graphical variation. Apparently, this is also the case in 0 0 (QM 46349); Tinaroo (17 08 S 145 35 E) (QM 64505); Johnson Australian Thiaridae, as is evident from our comments River, Innisfail (AMS C.51806); creek 10 miles of Innisfail, nr John- under each species in the systematic account (see son River (AMS C.109653); Burdekin River (BMNH 1846.10.7.33- 35; 1879.5.21.399-401, 406-7); Bellenden-Ker Range, nr Babinda above Part I). Here we report on some comparative (AMS C.51353, C.109651); Fisher´s Creek, Palmerston Rock, at Pal- morphological aspects across these taxa, exemplified merston Hwy (1734.1670 S 14553.8760 E) (ZMB 107251); Mena by the adult and juvenile shells as well as the radula Creek, Innisfail (AMS); South Mission Beach (1756.8400 S for particular species. The data available now provides 14603.2900 E) (ZMB 106340); Cardwell, Rockingham Bay (BMNH the basis for further evaluation of the (eco)phenotypical 1879.5.21.397-8, 415-6, 433-4); Fisher’s Creek, Palmerston Highway and geographical disparity in context with our ongoing (AMS C.126522); Upp Lynd River, Mt. Garnet to Mt. Surprise (QM molecular genetic studies, to be extended to the popu- 1620); Elizabeth Creek, E of Mt. Surprise (1808.351 S 14419.414 E) (ZMB 106726); Ross River (in weir) (1921.730 S 14643.930 E) lation level. (AMS C.338675); Saltwater Creek, Atherton (rocky creek) (AMS C.109132); Burdekin River, 100 km from the coast (in sand, shallow water) (VK 7780); Rosetta Plains, Burdekin River Adult shell (AMS C.8892); Rosella Plains, nr Cardwell (BMNH); Missionary Bay, Hichinbrook Island, South Creek (BMNH); Townsville, Aplins The most relevant shell parameters are compiled in Weir, Ross River (QM 38407); Townsville (in drain of salt pan nr Table 1, comprising a total of 1435 specimens for all town) (AMS); nr Almaden, Chilagoe Railway, Four Miles Creek these parameters measured for the 11 species-level taxa (AMS C.54093); off Ingham Road, Townsville (AMS); nr Eungella, discussed above and including the entire spectrum of Broken River (AMS); Alice River, at Eubenangee NP entrance growth (i.e. age dependent) stages from individual po- (1724.520 S 14558.850 E) (AMS C.338679); Charter Towers, Tank pulations or samples representing them. Note that for College (QM 1600); Charters Towers, NW at Toomba Stn, Great Ba- single parameters (e.g. shell height alone) more speci- salt Wall (19580 S 145340 E) (QM 53803); Allingham Creek, Bluff Downs Stn, 80 km WNW Charters Towers (19430 S 145360 E) (QM mens were measured in some species, as is evident 64450); Alligator Creek, Charters Towers, Wando Vale Rd crossing from comparing Table 1 with the total numbers in the (QM 64452); Calcifer Creek, nr Chillagoe (AMS C. 54417); Proser- following Figures 44 and 45. pine (QM 4411, 35346); Lethe Beach, 5 km S of Proserpine on Bruce We found the standard deviation to be high in parti- Highway (AMS); Conway River NP, circuit track, E of Prosperpine cular in those cases where there is a large difference (20160 S 148460 E) (QM 35346); Myrtle Creek, crossing on Bruce between the juvenile and adult shells and when older Highway, 5 miles N Proserpine (AMS); McKinley Creek, nr Mackay shells are prone to corrosion causing decollation, as is (VK 977); Isaac and Burnett Rivers (BMNH 1885.6.12-60); Fitzroy often the case e.g. in Stenomelania cf. aspirans and River (BMNH 1879.5.21.468-72); Fitzroy Island (AMS C.58394); Rockhampton (BMNH 1879.5.21.420.7); Rockhampton, Yeppen La- Thiara amarula (see Figure 44a). Alternatively we have goon (AMS C.118148); Rockhampton, western side of Frogmore La- used the heights of the last shell whorl or that of the goon (AMS C.109652); Alligator creek, 20 km N of Rockhampton last three whorls as a better indicator (see Figure 44b). (AMS); Addy Creek (BMNH); Andromache River crossing, via Gu- However, as is evident from these figures, the results nyarra (20340 S 148290 E) (QM 26376); Port Curtis (AMS); Mt. obtained for the comparison of the entire shell height Cooper, Rolleston River (AMS C.109429); Hays Inlet, nr Kallangur (H) as well as the height of the last whorl (BW) are (QM 28789); Gloucester Passage (QM 33609); Brookfield, Moggill very similar, revealing Stenomelania cf. aspirans and (27300 S 152300 E) (QM 64471); North Pine River, Dayboro (27150 S 152500 E) (QM 64472); Kedron Brook at Brook St. (QM Thiara amarula as being the tallest among the Austra- 64473); Fox Bridge, Mazlin Creek (QM 64479); Beantree Bridge, lian thiarids. The only exception among this otherwise Mazlin Creek (QM 65404); Kenilworth, above Mary River, Little highly uniform group of freshwater snails is found in Yabba Creek junction (QM 64495); Peterson Creek (QM 64506); Bur- Sermyla riqueti, being the smallest species. rum River, Howard (QM 64507, 64508); Paluma, Mt Spec NP, Crystal As a general quantitative indicator for the overall Creek (QM 59919); Walkamin, granite (QM 64465); Palms Island shape (“gestalt”) of the thiarid shell the ratio of shell 0 (QM 64469); Blackmans Creek, SW Galdstone (24 26.30 S height (H) and shell width (W) can be used, as visua- 15125.300 E) (AMS C.324136); Boyne River at Rosedale, S Gald- lized in Figure 45; this also allows to gain a measure- stone (2413.400 S 15115.300 E) (AMS); Brisbane, Sinnamon Pk. (QM 64475). ment independent of the direct effect of shell height New South Wales: Clarence River, S Grafton (29400 S alone. Accordingly, we found two groups among the 15255.9830 E) (AMS C.170629). Australian Thiaridae, which reflect different overall

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 256 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Table 1. Measurements for shell parameters of species of Australian Thiaridae, with min./max. values, mean, standard devia- tion (all in mm) and number of whorls (N).

height width length width last body N aperture aperture whorl

Thiara amarula min./max. 6.5/51.7 4.0/29.9 2.0/31.2 2.6/15.0 4.9/47.0 2–4 (n ¼ 92) mean 23.4 13.6 15.1 7.0 20.3 SD 14.0 7.9 8.6 4.0 12.6 "Thiara" australis min./max. 5.8/32.9 2.9/13.3 2.7/13.9 1.7/6.7 3.2/18.9 2–9 (n ¼ 576) mean 17.3 7.8 7.6 3.9 11.4 SD 4.7 1.8 1.9 0.9 2.8 Melanoides tuberculata min./max. 16.6/22.7 5.6/7.3 4.6/6.8 2.7/3.6 8.4/10.8 8–9 (n ¼ 24) mean 19.8 6.4 5.9 3.2 9.5 SD 1.4 0.4 0.6 0.2 0.6 Melasma onca min./max. 8.6/26.4 3.9/11.0 4.2/11.9 2.0/14;5.8 7.4/20.1 3–9 (n ¼ 176) mean 18.4 8.0 8.4 4.1 12.5 SD 3.0 1.1 1.2 0.6 1.7 Plotia scabra min./max. 9.6/15.6 4.9/7.4 4.3/7.2 2.4/3.6 6.2/10.2 6–7 (n ¼ 18) mean 12.3 5.9 5.6 3.0 7.8 SD 1.5 0.6 0.7 0.3 1.0 Plotiopsis balonnensis min./max. 7.4/29.5 3.5/12.2 3.5/10.4 1.8/6.4 4.7/18.0 4–9 (n ¼ 71) mean 16.5 7.5 6.6 3.8 10.6 SD 4.7 2.4 1.7 1.1 3.2 Ripalania queenslandica min./max. 9.0/33.5 5.5/13.3 4.8/12.8 2.4/6.2 9.4/22.5 2–6 (n ¼ 153) mean 21.6 9.2 8.7 4.2 14.9 SD 4.0 1.5 1.6 0.7 2.4 Sermyla riqueti min./max. 5.8/8.9 2.7/4.2 2.7/4.5 1.4/2.2 3.8/5.9 4–6 (n ¼ 17) mean 7.4 3.6 3.5 1.8 4.9 SD 0.9 0.4 0.4 0.2 0.6 Sermyla venustula min./max. 9.6/33.4 3.5/15.8 3.3/12.4 1.8/5.9 5.5/18.4 4–10 (n ¼ 146) mean 19.4 7.0 6.8 3.5 10.6 SD 4.6 1.7 1.7 0.8 2.6 Stenomelania cf. aspirans min./max. 8.7/78.9 4.4/26.3 4.1/30.0 1.5/15.4 5.5/50.8 2–7 (n ¼ 40) mean 50.7 16.1 17.2 8.8 30.3 SD 21.0 5.9 7.1 3.9 13.3 "Stenomelania" denisoniensis min./max. 8.3/36.4 3.0/12.8 2.6/11.9 1.5/7.0 4.5/20.1 3–9 (n ¼ 122) mean 19.8 7.9 7.8 4.0 12.0 SD 5.2 1.7 1.6 0.9 2.5

shell shape, with T. amarula on one side and S. cf. as- In addition to this more general interspecific charac- pirans on the other side representing the end points of terization, the shell parameters gained in the present this continuum. In general, together with this latter spe- study will be used for a more in-depth evaluation of cies we found also “Stenomelania” denisionensis, Ser- interpopulational variation within individual species. myla venustula and Melanoides tuberculata to be char- Preliminary results (not shown here) indicate for Plo- acterized as of more elongated and turreted shape, tiopsis balonnensis, for example, that populations from whereas the other species have a more rounded to glo- the eastern part of its range reach on average signifi- bose shell shape. cantly larger shell size than those found isolated in As is evident from these comparisons, shell dimen- water pools of the Finke River drainage in Central Aus- sion hardly help in distinguishing between pairs of “cri- tralia. Other shell features, such as e.g. sculpture or col- tical” (i.e. fairly similar) species, such as e.g. australis ouration, are also found to vary across the geographic versus balonnensis or denisoniensis versus tuberculata, range of individual species, but await future analyses in which resemble each other in dimension and overall order to account for ecophenotypic versus other possi- shape. However, it should be noted that e.g. Melanoides ble correlations. On the other hand, Melasma onca (see tuberculata is slightly set apart in its H/W ratio from Figure 13) and even more so Ripalania queenslandica “Stenomelania” denisoniensis,asisSermyla venustula (Figure 25) exhibit quite homogenous sculpture and col- not only from riqueti but also from australis, with our of their shells not varying in the same degree as which it often was found to co-occur. found in other thiarid species. It is in particular diffi-

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 257 cult to differentiate, on the other hand, the juvenile whorls than in others (see Figure 40, compare e.g. c stages of many, often syntopic taxa among the Austra- with g). lian thiarids. For example, free-ranging juveniles of “Thiara” australis and Sermyla venustula are notor- Juvenile shell iously difficult to separate immediately where they co- occur, as e.g. in several locations along the Roper River The juvenile shells we discuss here are those only that drainage (see Figure 2). were found in the brood pouches of Australian thiarids, The latter species is also found to be extremely which in their majority actually exhibit an eu-vivipar- polymorphic in its adult shell, with very variable po- ous strategy (see below Part III), thus bearing shelled pulations occurring even within the same river sys- juveniles inside their marsupium. In general these tem. For example, shells of S. venustula depicted in shells are similar to the adults with respect to their Figures 31f & 31g are from two adjacent localities of sculptural features. For example, the shape and pattern the Roper River system, only found at a distance of of ornamentation with axial and spiral elements in ju- few kilometers apart from each other. Nevertheless, venile Plotia scabra, as in Figure 18a, is very similar to as discussed under this species (see there) they have the shells of the adults as depiected in Figure 17. How- distinct shell features and colour pattern, making ever, it should be noted that the ornamentation only them prone to superficially be assigned to distinct gradually become more similar to that in the adult (morpho-)species. A quite similar case is found in shell. Thus, not only on the first two to three whorls shells assigned here to venustula but resemble very but also on subsequent ones other sculptural elements much those described originally as “carbonata” (see might still dominate. Figure 32). While the syntypes from “Port Essington” We found Melasma onca to be the only species (see above under the species) exhibit only few axial among those that brood shelled juveniles, in which elements or ribs on the most upper whorls, other these only possess three whorls when found in the mar- specimens of this species from Howard Springs, supium (Figure 14). The orthocline ribs typically seen which are attributed tentatively to “carbonata” here, in adult M. onca are consequently not yet present in have much more pronounced axial ribs on their later these early ontogenetic stages. However, we recognized whorls. in this species that the most apical part of the embryo- In a similar way the shells of “Thiara” australis (see nic shell is often covered by a thin, translucent cover- Figure 8) and of “Stenomelania” denisoniensis (see Fig- age of unknown nature (Figure 14d), not seen in other ure 40) which are both widely distributed exhibit a rich Australian thiarids. In contrast to the latter species, the spectrum of conchological variability. In the former in juveniles of all other thiarids reach a much larger shell particular the form and number of axial ribbing is height with up to seven whorls completed within the highly variable, as well as of the spiral ridges and shell female’s marsupium (see also below Part III, Fig- colour. In contrast, in “Stenomelania” denisoniensis we ure 47). found that the shell is in some populations much more For species distinction features of the juvenile shells pronouncedly angulated in the apical part of the (last) are generally neither sufficient nor better suited than

Figure 44. Box plots with mean, standard deviation and min.-max. distribution for two shell parameters of Australia Thiaridae. In grey are on the left hand side the two largest species Thiara amarula (Linnaeus, 1758) and Stenomelania cf. aspirans (Hinds, 1844); marked in black is the smallest species Sermyla riqueti (Grateloup, 1840). a. Height (H); b. Height of last body whorl (BW).

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 258 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae those of the adult shell. Polymorphic species such as in the other three taxa, with “Stenomelania” denisonien- “Thiara” australis and “Stenomelania” denisoniensis sis possessing the largest of these “protoconchs” (Ta- exhibit a similar variability in their juvenile shell as re- ble 2). The shape of the embryonic shell varies consid- ported earlier for their adults. These differences can erably among individuals of one clutch even within the clearly be seen among specimens with equal number of same brood pouch of an animal of a given species, such whorls, as e.g. in australis (compare Figures 9a, e and as e.g. P.balonnensis (compare Figure 22), thus hardly j); therefore, we conclude that shell variability is inde- providing distinct species-specific characters. pendent of the individual developmental stage reached Typical for eu-viviparous thiarids, though, is the by these juveniles. We found many intermediate stages wrinkled sculpture of the initial cap (i.e. the “An- among the various shells of different age among the ju- fangsschlchen” or “Initialkalotte”), as was also re- veniles of a female’s marsupium or within a single ported for other non-Australian Thiaridae (see Glau- samples from one locality representing the population brecht 1996). We found it to be present in all five eu- (compare Figures 9a and e). viviparously brooding thiarids, with a more or less These intermediate phenotypes can also be found marked transition to subsequent whorls as can be seen among distinct species, such as e.g. in the juveniles of in australis (Figure 9m), onca (Figure 14d), balonnensis “Thiara” australis (Figure 9j) which exhibit a net-like (Figures 22d, h, m), venustula (Figures 33d, h), and de- or reticulate ornamentation with pronounced spiral nisoniensis (Figures 41d, h). ridges, as it is otherwise also found in the juvenile shells of “Stenomelania” denisoniensis (Figure 41) and Radula Melanoides tuberculata (for the latter species see fig- ures in Glaubrecht 1996). A similar sculpture was Ever since Troschel’s (1857) epochal study of the gas- found in specimens of Sermyla venustula (Figure 34a). tropod radula this feature is of central importance in In contrast to the specimen depicted in this latter fig- molluscan systematics on a higher taxonomic level. ure, another juvenile of the same species and locality However, it remains to be evaluated whether the radula shows a quite distinct ornamentation with more typical is also useful on the level of species or between popu- concave axial ribs, though (Figure 34b; see also Fig- lations. In thiarids, unfortunately, the radula is appar- ure 33). ently of only limited use in species distinction, as in However, we found some remarkable differences in our study of the Australian taxa we found it to be rela- the standard size parameters taken for the embryonic tively constant and conservative in its essential features whorls of these shells within the marsupium, as com- between species on the one hand, while it is very vari- piled in Table 2 for a total of 28 “protoconchs” from able in some details within species on the other hand. the five native Australian thiarids that frequently brood As consistent feature we found that the taenioglossate shelled juveniles in their marsupium; for a comparison radulae in Australian thiarids exhibits between 90–130 with the presumably non-native eu-viviparous thiarids rows, with the length and width of the radula ribbon Plotia scabra and Melanoides tuberculata refer to Glau- being correlated with apparently the shell height (i.e. brecht (1996). While the diameter at first whorl appears to size) of the animals. Accordingly, the radula is about be fairly constant, with about an average of 200–250 mm 2 mm in lenght and 0.5 mm in width in smaller species, in all five species, the size of the inital cap varies. As while e.g. in larger species such as Thiara amarula and indicated by the height and width parameter, it is on Stenomelania cf. aspirans it is more than four times average smaller in P.balonnensis and S. venustula than larger. As noted under the species, only the radula of

Figure 45. Box plots with mean, stand- ard deviation and min.-max. distribution for the height/width (H/W) index of the adult shells of Australia Thiaridae. In grey on the left hand side are species with more rounded shell shape in com- parison to those indicated in black on the right hand side with more turreted, elongated shell phenotype.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 259

Table 2. Measurements for parameters of the juvenile shells of Australian Thiaridae found in the brood pouch (in m).

height width diameter

"Thiara" australis (n = 9) mean SD 142.1 21.6 89.2 16.3 237.6 39.9 Melasma onca (n = 3) mean SD 153.2 11.7 91.6 4.1 238.4 7.6 Plotiopsis balonnensis (n = 10) mean SD 46.2 4.0 69.9 6.8 211.7 11.7 Sermyla venustula (n = 3) mean SD 107.5 14.2 66.4 6.2 196.5 15.6 "Stenomelania" denisoniensis (n = 3) mean SD 227.4 77.3 95.1 26.0 248.5 58.6 the latter taxon, with its almost perfectly rectangular Of these, only 6 animals were found to not possess a rachidian and the sharply pointed mesocone of the la- brood pouch filled with shelled juveniles. In other teralia (Figure 38), was found to be significantly dis- cases, we also found a very low frequency of non-grav- tinct to allow species-specific identification of aspirans id females, usually in the range of less than 10 percent, based exclusively on the radula. as it is also known for Melanoides tuberculata (Glau- On the other hand, the radula was found to be vari- brecht unpubl. data). able in details of the denticle numbers and shape in In the course of our study on viviparous strategies in most Australian species. For example, in “Thiara” the Australian thiarids as detailed below, we gathered australis it differs in animals from across the geo- further data that indicate an only low frequency or even graphic range of this species (see Figure 10). Also, in complete lack of males in a given population. Thus, we Plotiopsis balonnensis the shape of individual denti- conclude that many Australian thiarids reproduce by cles of the rachidian and the lateralia varies in indivi- parthenogenesis or at least by maintaining only very dual populations across the distributional range (see low abundancies of males in their populations. Figures 23b, f). In addition, the number of denticles in the marginalia of most taxa is variable, without a spe- cific pattern discernable. So far we failed to find a Viviparity and life history strategy clear-cut correlation between these minor differences in the dentition patterns of their radula within a given Life histories have been defined as the probabilities of species or between individual populations or shell survival and the rates of reproduction at each age in morphologies. the life-span (Partridge & Harvey 1988). As a proxy to the second aspect of life histories strategies Glaubrecht (1996, 1999, 2006) suggested to differentiate within the viviparous Thiaridae two reproductive modes, depend- Part III: Reproductive biology ing on the duration of ontogenetic stages to remain within the marsupium of the female, which at the same Sex ratio and parthenogenesis time might also reflect a differential degree of parential Hitherto, it has been assumed that all Thiaridae are re- investment, even including nourishment of the embryos producing parthenogenetically, as was in particular de- and juveniles via nutritive tissue. While in some thiar- scribed only in Melanoides tuberculata or in Plotia sca- ids only very early ontogenetic stages, viz. eggs or em- bra (see Glaubrecht 1996). We here evaluate the bryos without shell, develop and are released as veli- available data for the Australian thiarids; however, we gers, other thiarid species brood and even transform only indirectly contribute towards this question here, as their subhemocoelic brood pouch into a “pseudoplacen- we estimated the frequency of females by them posses- ta” (Glaubrecht 1996) that, via matrotrophy apparently sing a brood pouch filled with early ontogenetic stages help to nourish the developing juveniles, as e.g. in the (either eggs, embryos or shelled juveniles), but did not SE Asian thiarid Tarebia granifera. These juveniles estimate the frequency of males by routineously check- hatch at a later stage, i.e. as crawling young with shells ing testis tissue. Even more difficult is a sex determina- of three to five or even more whorls. These two distinct tion using the differential morphology of the pallial go- strategies have been termed (i) ovo-viviparity in case of noducts. Thus, while we can be positive about a gravid brooding and releasing veligers, versus (ii) eu-viviparity female we were not able to definitely differentiate be- in those cases where the females brood shelled juve- tween non-gravid adult females (i.e. of equal size as niles which they might also nourish. For discussion of the largest specimens out of a sample studied) and terminology see e.g. Fretter (1984) and for thiarids males, albeit we assume from our observation that the Glaubrecht (1996). former case appears to be rare if not at all non-existent We have above under each species reported on in most thiarids. In this context, we e.g. dissected a to- these reproductive modes among the viviparous strate- tal of 66 specimens of about equal shell size of “Steno- gies of Australian Thiaridae which, however, does not melania” denisoniensis from Poison Creek, a tributary correlate with distinct reproductive anatomies, i.e. the of the Endeavour River in Queensland (ZMB 106356). development of the brood pouch and “placental” tissue

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 260 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Figure 46. Reproductive biology of three thiarid species Melasma onca, Sermyla venustula and “Thiara” australis from four local- ities along the Roper River (NT); data are given for N = 64 adult specimens collected at the end of the dry season in 2005. a. Height of the adult shell (in mm) in comparison with the rough amount of shelled juveniles (given for four classes) found in the brood pouch; b. Frequency (in percentage) of adults with and without any early ontogenetic stages or shelled juveniles found in the brood pouch.

(Glaubrecht unpubl. data). In summary, we found that Sermyla riqueti actually only contain eggs and embryos while all thiarids retain their eggs within the brood up to very early developmental stages before being re- pouch, four of the Australian taxa studied here, viz. leased as veligers. These resemble those found in other Thiara amarula, Ripalania queenslandica, Stenomela- ovo-viviparous Thiaridae, such as e.g. Balanocochlis nia cf. aspirans and Sermyla riqueti, release free glans and Thiara cf. rudis (Glaubrecht et al. unpubl. swimming veligers. In contrast, in all other Australian data). They exhibit a clear body division with visceral taxa the embryos were continuously withheld in the mass and cephalopodium; the embryos are surrounded incubatory pouch for further development and only by a very thin egg membrane and possess next to cilia hatch as shelled juveniles. This later eu-viviparous in later stages also an operculum, as will be described strategy was found in all five endemic species, viz. in detail elsewhere. “Thiara” australis, Plotiopsis balonnensis, “Stenomela- The average diameter of these early embryos is less nia” denisoniensis, Sermyla venustula and in Melasma than 0.1 mm, filling the female’s brood pouch as den- onca, as well as in the two (presumably) non-ende- sely packed “egg masses”. We have only very roughly mics Melanoides tuberculata and Plotia scabra. Some estimated their numbers in the marsupia of the females of the aspects associated with these different life his- dissected, arriving at figures of several hundreds to tory strategies will be briefly outlined in the follow- even thousand. The duration of brooding and the exact ing. timing when these veligers are released is unknown for Thiaridae in general and, unfortunately, also for the Australian taxa under study; the release of free-ranging Ovo-viviparity and veliger veligers in the above cited observation was artificially The first author has evidence, by direct observation of induced, but is was noted that these veligers of T. ama- captivated animals from Queensland that were held in rula and S. aspirans survived in tab water for a couple an aquarium from November to December 1996 at the of days. It is also noteworthy that within the marsupium AMS, that both Thiara amarula and Stenomelania cf. of Australian ovo-viviparous thiarids non-shelled em- aspirans release free swimming veliger from their bryos in different ontogenetic phases are found simulta- pouches. Subsequent anatomical studies of these and neously, comprising gastrula-like stages as well as rela- other conspecifics revealed that the marsupia of these tively advanced stages of veligers with clearly visible two species as well as of Ripalania queenslandica and operculum.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 261

Eu-viviparity and shelled juveniles be given. However, as general rule we conclude that the larger the juveniles in the brood pouch the lower the The majority of Australian Thiaridae withhold their lar- number of (shelled) juveniles – and vice versa, i.e. the val stages inside the marsupium until shelled juveniles smaller the juveniles the larger the total number of have developed, with their shells comprising up to sev- those found in the marsupium. The latter was in parti- en whorls before they hatch as crawling young. These cular the case in Melasma onca, where we found the juvenile shells found in the brood pouch document their juveniles to be rarely larger than one millimeter, but ontogeny to the extend that the wrinkled sculpture of with the marsupium filled with several hundreds of em- their “Anfangsschlchen” and the more or less clearly bryonic stages. In contrast, all other Australian species visible marked transition after the first whorl when only possess relatively few, but fairly advanced juve- growth lines start (see e.g. Figure 22d) and with gradu- niles stages. ally other ornamentation developing on subsequent In this context we studied in some more detail repre- whorls all give witness to their peculiar eu-viviparous sentative samples of three thiarid species from different development inside the female’s body (for details and localities along the Roper River, Northern Territory, discussion see Glaubrecht 1996). viz. for Melasma onca (n ¼ 12 specimens), Sermyla ve- We found in the marsupium of the five eu-vivipar- nustula (n ¼ 25) and “Thiara” australis (n ¼ 27). First, ous Australian thiarid species that all growth stages we correlated the height of the female shell with the are present at the same time, often with the majority frequency of its marsupial content, differentiated into being the eariest developmental stages while only few four distinct classes according to ontogenetic stages (i.e. up to a number of one or two dozen) juveniles found within the clutch. As is evident from Figure 46, already possess relatively large shells. As a general in all three species the amount of shelled juveniles in rule we found during dissections that the most ad- the brood pouch is positively correlated with adult vanced and thus oldest stages lay most anterior and shell size, i.e. larger females have more shelled juve- adjacent to the brood pouch pore or marsupial open- niles (Figure 46a). On the other hand, those females ing, as was also described and depicted e.g. for Mela- without shelled juveniles (i.e. with eggs and very early noides tuberculata in Glaubrecht (1996). In the Austra- embryos only) are on average not smaller than those lian taxa we did not study yet, down to histological animals with more advanced stages. The frequency of detail, the compartimentalization often noted in these these gravid females without shelled juveniles in their marsupia nor the exact composition of the brood brood pouch was found to be about a quarter to more pouch tissue. However, the marsupium was found to than a third in the three thiarid species under study be of more or less similar anatomy as that seen in (see Figure 46b). other eu-viviparous Thiaridae; though, with no indica- In addition we found that M. onca holds the largest tion of a clearly developed nutritive tissue seen, as it number of embryonic stages per brood pouch, with was in contrast reported and depicted e.g. for Tarebia most stages being in the size range of up to 0.5 mm granifera (see Glaubrecht 1996). and 1.0 mm, but only very few more advanced and As the number of shelled juveniles is highly variable shelled embryos with a size of up to 1.5 mm (Fig- in the thiarid taxa studied so far, no reliable figures can ure 47). In contrast, the sympatric and syntopic thiarids

Figure 47. Number of individuals found for five different ontogenetic stages (from embryos without shell to juveniles with shells larger than 1.5 mm) in the three thiarid species Melasma onca, Ser- myla venustula and “Thiara” australis from the Roper River (NT); data are giv- en for N = 64 adult specimens collected at the end of the dry season in 2005.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 262 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

S. venustula and “T.” australis both show a fairly even Thiaridae in Australia. These taxa as differentiated and distribution of all different developmental stages in documented herein, allocated to specific genera and their marsupium, but both have a significantly lower names, are only partially consistent with those species number of juveniles than M. onca, which is thus clearly and listed in earlier catalogues of the non-marine fauna exceptional among the eu-viviparous Australian Thiari- of this continent (see introduction under Thiaridae). dae. The eleven species we identified by the morphology of their shell and radula in concert with their geographical occurrences and reproductive biology; they are also Discussion corroborated as distinct clades by our molecular genetic data, to be reported elsewhere. Three of these taxa, viz. This study of Australian Thiaridae is done in the frame- Stenomelania cf. aspirans, Sermyla riqueti and Plotia work of the systematics and evolutionary ecology of scabra, are recorded here for the first time. Note that freshwater gastropods among the Cerithioidea. Based possibly two more lineages have to be named, for aus- on the taxonomy and reconstruction of phylogenetic re- tralis and denisoniensis, respectively, in case of verifi- lationships we attempt to integrate the study of specia- cation and thus necessity; see for more under the taxo- tion mechanisms and radiation events with life his- nomic remarks for these two species. tories, i.e. reproductive strategies (as proxy in the case This new systematization explicitly suggests a new of these invertebrates), and an evaluation of competing concept for the formerly widely delineated genus biogeographical hypotheses concerning vicariance ver- Thiara, which was found in our analyses to be poly- sus dispersal and/or ecological causation, as outlined in phyletic (Glaubrecht et al. unpubl. data). Therefore, we Glaubrecht (1996, 2000, 2009). here restrict its use to a monophyletic clade with the In this context, for the Australian taxa in particular a type species amarula also present in Australia, but to suite of phenomena are undoubtedly highly relevant but the exclusion of three other species that are to be allo- await detailed studies, corresponding not only to bio- cated in their own genera, viz. “Thiara” australis, Plo- geographical facts (such as the chorological relation- tiopsis balonnensis and Plotia scabra. In case of ships with extralimital occurrences or endemism in spe- australis we found the (albeit also paraphyletic) cific regions) but also to specific adaptations to cope Indonesian thiarid Tarebia granifera to be its adelpho- with special Australian environments and ecological taxon; however, this phylogenetic affinity remains to be circumstances, such as e.g. increasing aridity, unpre- verified. Therefore, we here refrain for the time being dictable precipitation and ephemerality of freshwater from using either Thiara, Tarebia or a new name, but bodies correlated with marked hydrological fluctuations indicate this unresolved situation by using “Thiara”in of many rivers and streams and high salinities in tem- quotation marks only. A similar unresolved case is evi- porally standing waters, as briefly outlined in the intro- dently “Stenomelania” denisoniensis, which was neither duction. In addition, viviparous strategies, as found in found to be genetically allied to the type species aspir- the Thiaridae, have been perceived of as key innova- ans nor to the morphologically often very similar Mel- tions accompanying not only the colonization of fresh- anoides tuberculata. Since species of both genera are water in general, but also effecting the sometimes strik- currently under study we refrain here also from apply- ingly different distributional patterns as well as ing a new generic allocation which, however, might endemic occurrences among several taxa of these lim- turn out to be necessary. nic Cerithioidea from the Australasian and adjacent re- gions. It was the aim of the present account to provide the Taxonomical diversity versus morphological disparity systematic, biogeographic and reproductive biology data as basis for these future investigations, looking As discussed in general by Glaubrecht (2004, 2009), into significant evolutionary phenomena, such as frag- the widely adopted typological practice during the 19th mentation of species’ ranges, geographical separation of and way into the 20th century of naming several allopa- populations potentially leading to speciation, and/or hy- tric populations, in isolated fashion and often based on bridization. Although the many details of these histori- single specimens only, as if representing putatively dis- cal processes and their evolutionary consequences re- tinct (morpho-)species has led to a plethora of species main to be studied for Australia’s freshwater fauna, we and subspecies names in malacology. This often re- will summarize and discuss some of the relevant as- sulted in a considerable amount of taxonomic redun- pects here for the Thiaridae. dancy and, consequently, inflation of biodiversity as discussed recently for molluscs in Glaubrecht (2009; see also literature therein). Evidently, the factor for the Systematics of Australian Thiaridae average number of species described for a given time, As a result of the present study, but in combination taxon or region and those that were later regarded as with our ongoing molecular phylogenetic investigations being valid species was found, albeit on a different ba- (Glaubrecht et al. unpubl. data), we propose the exis- sis than here, to be between 1 : 5 and 1 : 6. Interest- tence of 11 species in 8 genera (currently named) of ingly, we here found out of the maximum number of

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 263

16 named species for the Australian thiarids as listed brecht & Rintelen 2008). Supporting Benthem Jutting’s by Iredale (1943) and B. J. Smith (1992) a total of 11, (1934: 328) certainly correct statement on the thiarid or 69 %, species to exist according to our analyses, radula that in its form the function is expressed, we suggesting a synonymy ratio of 1 : 4.5 and thus to be found for the Australian taxa hardly any major denti- slightly lower than that of 38 % estimated for other tion feature in a given species to allow for specific dis- molluscs so far. tinction. Apparently, all taxa under study seem to use In general, freshwater gastropods were found to exhi- their radula in the same fashion, as all of them are det- bit a pronounced individual conchological variability, ritus and algae feeders, mostly occurring on or in soft which has been attributed to the environmental condi- substrate with high sandy to muddy components. Only tions of their habitats that widely fluctuate on a tempor- in Stenomelania cf. aspirans we here report some dis- al and spatial scale (see e.g. Rensch 1929, 1934; Glau- tinct features of the radula that, consequently, might po- brecht 1993, 1996; Dillon 2000). However, even the tentially indicate an (albeit only slightly?) different most pronounced of these conchological features, such food source and/or feeding strategy. as e.g. shell size, shape and sculpture as well as colour, Therefore, those characters of shell plus radula that hardly allow for definite species identifications, as was were classically utilized most in limnic malacology exemplified among the freshwater Cerithioidea for Me- (e.g. Martens 1883; Westerlund 1892; Thiele 1928, lanopsidae (Glaubrecht 1993, 1996) and Pachychilidae 1929; Rensch 1934; Starmhlner 1969, 1973, 1976, (Glaubrecht & Khler 2004; Khler & Glaubrecht 1983, 1986, 1993; Glaubrecht 1996, 2008) are now 2001, 2003, 2006; Khler et al. 2008). In this context, known to be of only very limited use in genus or even we found in the present study on Australian Thiaridae species level taxonomy. Also, the juvenile shell (even that the use of conchological features is of limited use when preserved in the brood pouch) only reflect the only. Shell characters allow for the overall differentia- variability of the shells in the adults. Surely, these facts tion among most of the Australian taxa, but less so hamper any quick and clear differentiation of species within what is here considered, as a consequence, to or even distinct populations in context of biosyste- represent biospecies. matics and biogeography, as was realized right at the As detailed in the introductory section, E. A. Smith time of the advent of molecular genetics. However, to (1882) compiled a total of 12 thiarid species for Aus- date only Stoddart (1985) applied other methods, when tralia, which Iredale (1943) increased to 16 later, ac- using isoenzyme electrophoresis investigating 16 popu- cepted or at least considered most recently also by lations of two Australian thiarid species from 14 local- B. J. Smith (1992) in his catalogue. Stoddart (1985) ities across the continent. He arrived at the conclusion summarized the situation when noting that “Australian of having dealt with 13 populations of Thiara [= Plo- thiarids have been referred to a large number of named tiopsis] balonnensis while the remaining three were re- forms, frequently on the basis of a few morphological presentatives of Thiara [= “Stenomelania”] denisonien- characters, and frequently to forms with very limited sis; identified on the basis of their shell, he was distribution”. We found that in particular those species unable to differentiate these thiarids on a finer scale, with wider distribution across the Australian continent, though. like “Thiara” australis, Plotiopsis balonnensis and “Ste- In case of thiarids the application of a particular spe- nomelania” denisoniensis, are often extraordinary vari- cies concept is not made easy given the prediction, able even within single populations but all the more which is supported here, that these gastropods repro- among populations widely separated across river sys- duce largely via, at least partial, parthenogenesis (see tems and the various regions of the continent. Single above Part III). This was shown for Melanoides tuber- specimens of “problematic” species pairs such as culata (Jacob 1957, 1958; Berry & Kadri 1974) and as- “Thiara” australis and Plotiopsis balonnensis (including sumed also by Stoddart (1985) for two Australian thiar- also Plotia scabra) as well as “Stenomelania” deniso- ids. Morrison (1954: 375) in discussing the enormeous niensis and Melanoides tuberculata were found difficult shell variability in thiarids in context with parthenogen- to be identified based solely on their shell. In contrast, esis noted, wisely, that “wise indeed is the scientist who species with more restricted geographic range were in can tell whether a clone is a species or not, and be general also less variable, which holds true in particular right every time, in the case of the Thiaridae”. Stoddart for Melasma onca as a species easily distinguishable (1985) applied the evolutionary species concept (ESC) due to its fairly constant conchology (in addition, of following Wiley (1978, 1981; see for discussion also course, to those with highly distinct shells anyway, like Willmann 1985), as in his opinion this concept “stres- Stenomelania cf. aspirans and Thiara amarula; see Fig- ses the relevance of the process of speciation to species ure 45). definitions and provides the most appropriate frame- Although the radula is known to provide another dis- work for the taxonomy of asexual organisms”. Although tinct character since Troschel (1857), its usefulness in this statement is debateable for several reasons, admit- differentiation of freshwater Cerithioidea on the species tedly, the biological species concept is also not without level is debateable (see e.g. Martens 1883; Sarasin & problems in application here. The BSC was introduced Sarasin 1898; Benthem Jutting 1934; Rensch 1934; by Mayr (1942: 120) and since then widely discussed; Glaubrecht 1996; Glaubrecht & Khler 2004; Glau- see literature survey with references updated and dis-

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 264 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae cussion with respect to limnic gastropods e.g. in Glau- production. We have additional (direct) evidence for brecht (2004, 2009). males in thiarids, as we know of spermatophores, for Explicitly, the BSC uses the reproductive criterion in example, in Thiara amarula from Queensland (as re- sexually reproducting organisms. It case of the thiarids, ported in Glaubrecht & Strong 2004) and in Balanoco- however, it remains to be seen in how far they are actu- chlis glans (Glaubrecht & Brinkmann unpubl. data); ally prone to parthenogenesis. For example, for popula- furthermore, male genital anatomy was described e.g. tions of Melanoides tuberculata in Israel Ben-Ami & by Starmhlner (1969, 1976) and Schtt & Glaubrecht Heller (2005) reported sexually as well as asexually re- (1999) for species such as Thiara amarula and Steno- producing individuals, thus contradicting the general as- melania aspirans. sumption that indeed all thiarids reproduce via apo- mixis. Apparently, at least in M. tuberculata there are Reproductive biology both modes realised, securing the exchange of genetic information by sexual reproduction as was shown in The majority of Australian thiarids are eu-viviparous, earlier allozyme studies (Livshits & Fishelson 1983) i.e. they incubate their embryos that subsequently de- and excluding the possibility of gynogenesis, i.e. velop multi-whorled shells inside the brood pouch until parthenogenesis with the development of eggs to be in- hatching as crawling juveniles. In contrast, four of the duced by contact with sperm, though. Given the fact thiarid species discussed here have only veliger stages that we have (albeit indirect) evidence for the presence in their brood pouches, thus being ovo-viviparous, viz. of males at least in low frequency (of up to 10 %) in Thiara amarula, Stenomelania aspirans, Ripalania some Australian thiarids, we assume that also in other queenslandica and Sermyla riqueti (for discussion of cases thiarids either maintain this low level of males or terminology see below). Accordingly, the Australian by other means switch between asexual and sexual re- thiarids exhibit distinct life histories, here described in

Figure 48. Geographical ranges of “Thiara” australis (Lea & Lea, 1851) (black symbols), restricted to the wet-dry tropical north of Australia, and Plotiopsis balonnensis (Conrad, 1850) (grey symbols) with wide distribution across the continent. For discussion see the text.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 265 terms of their different reproductive strategies, which populations sexual reproduction is maintained, irrespec- are potentially also correlated with differential parental, tive on the one hand of the obvious advantages, but i.e. maternal, investment (Glaubrecht 1996). In this also on the other hand in context with the “Red Queen” context, viviparity has not only been discussed as an hypothesis and the role of parasites, as discussed e.g. important key innovation for freshwater colonization for Potamopyrgus antipodarum (Lively 1987, 1992; (e.g. Cunnington 1920; Sunderbrink 1929; Seshaiya Jokela & Lively 1995; Lively & Jokela 2002; Jokela 1936, 1940; Morrison 1954; Hubendick 1962; Davis et al. 2003) or Melanoides tuberculata in Israel (Ben- 1982; Stoddart 1983; Fretter 1984; Starmhlner 1973, Ami & Heller 2005). 1984), but this specific mode of reproduction and re- It should also be considered for Australian thiarids cruitment strategy is also of paramount importance that parthenogenesis, in combination with viviparity with respect to the specific environmental conditions (see below), might be directly correlated and/or even for freshwater organisms in Australia, given the ex- causally linked – in context of the unpredictability of treme climatic variables of this continent, such as, for precipitation and the ephemerality of water bodies – example, its aridity and seasonality of rainfall resulting with the survival of populations in or rapid (re-)coloni- in monsoonal flooding of vast areas right after long zations of new habitats and areas right after or in the droughts. Although these factors and correlations were course of flooding, potentially being an adaptation to recently been exemplified and discussed using verte- the monsoonal regime of northern Australia. brates (in particular terrestrial reptiles) (see Shine & Brown 2008), the same has barely been touched to date for aquatic invertebrates such as thiarids. Only some Viviparous modes and matrotrophy preliminary remarks are outlined here based on the available insights yet. Interestingly, limnic Cerithioidea exhibit a wide array of reproductive modes that are otherwise only found in all Caenogastropoda taken together (Fretter & Graham Life history strategies of Thiaridae 1994; Webber 1977; Aldridge 1983; Fretter 1984; Reid from the Australian perspective 1990). Ovipary represents the plesiomorphic condition in this superfamily and viviparity has evolved several Parthenogenetic reproduction has found much interest times independently (Glaubrecht 1996, 1999, 2006; in the past in evolutionary biology, not only with re- Khler et al. 2004). In this context Fretter (1984) con- spect to the origin of sex. Clonal reproduction in natur- sidered true viviparity and brood care to be realized al populations has obviously many advantages over sex- only when there is direct flow of nutrients from the fe- ual modes, with growth rates in the former often being male to the embryo “involving transfer of organic much accelerated over the latter, as all individuals with- food”. Although Fretter & Graham (1994) therefore ex- in the population are able to contribute (Maynard Smith cluded the presence of this mode of viviparity in any 1978). In addition, these clones are considered instru- caenogastropod, a nutritive function of the epithel of mental in fast colonization of new habitats and areas, the brood pouches in some species of thiarids was as- as even a single female can give rise to a new popula- sumed earlier by Ramamoorthi (1950) and Muley tion (Baker 1955). Nevertheless, most faunas are domi- (1977). It was also postulated by Glaubrecht (1996) nated by sexually reproducing species, with asexual or- based on histological serial sections of Tarebia grani- ganisms being in the minority (Bell 1982). There is a fera that nutrients are provided via matrotrophy (via a vast literature available discussing these phenomena placenta-like epithel) in thiarids. In contrast, ultrastruc- (e.g. Williams 1975; Hamilton et al. 1990; Kondrashov tural analyses of the brood pouch tissue in Melanoides 1993). tuberculata from Israel did not reveal indications for Also in malacology there are some classical case stud- true viviparity sensu Fretter (Ben-Ami & Hodgson ies, such as the New Zealand freshwater hydrobiid 2005). Accordingly, juveniles are considered by these Potamopyrgus antipodarum (Gray, 1843) (see e.g. Jokela latter authors to be simply kept in the brood pouch of et al. 2003) or the thiarid Melanoides tuberculata (see some thiarids, but not to be nourished maternally; thus, e.g. Jacob 1957, 1958; Berry & Kadri 1974; Ben-Ami in accordance with Fretter (1984) the term ovo-vivipar- & Heller 2005). However, in both cases reproduction is ity was applied, as it is basically a lecithotrophic devel- not exclusively parthenogenetic. In populations of tu- opment, also to this kind of incubation within the mar- berculata, for example, the frequency of males was supium. found to vary between 40 % in the French West Indies In the present study we have not found clear evi- (Samadi et al. 1998) and up to 66 % in Israel (Livshits dence for matrotrophy in Australian Thiaridae. How- & Fishelson 1983; Heller & Farstey 1990). We here an- ever, we refrain from adopting the term ovo-viviparity ticipate a similar phenomenon in Australian Thiaridae, in those cases where shelled juveniles are incubated for with varying frequencies of males (even less than longer spans of time within the female marsupium. We 10 %), but propose that reproduction is not generally find it necessary to clearly differentiate as done here parthenogenetically only. It remains to be investigated between (i) an obviously ovo-viviparous strategy, when why and how in individual thiarids and their Australian only veligers are released, and (ii) an eu-viviparous

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 266 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae strategy, when more advanced juveniles hatch with lected species, albeit controversially and with now hav- shells of three to even seven whorls. These two distinct ing “something of a bad name” (Partridge & Harvey strategies were found to be realized in Australian thiar- 1988: 1453). Background to this is the observation that ids as summarized here: r-selected species with high reproductive rates (i.e. the Malthusian parameter or intrinsic rate of increase) (i) ovo-viviparity: evolve under density independent conditions compared Thiara amarula, Ripalania queenslandica, Steno- to K-selected species that have low reproductive rates melania cf. aspirans, Sermyla riqueti. evolving under density dependent, i.e. saturated condi- (ii) eu-viviparity: tions (with K for carrying capacity). “Thiara” australis, Plotiopsis balonnensis, “Steno- Glaubrecht (1996) used this theoretical concept of melania” denisoniensis, Sermyla venustula, r- versus K-selected species that was suggested by Melasma onca, Melanoides tuberculata, Plotia sca- MacArthur & Wilson (1967) for certain cases in biology bra. only, and adopted their terminology in order to describe Only when applying this crucial distinction in the re- differential reproductive strategies involving highly vary- productive mode (as proxy for different life history ing numbers of offspring that are released and/or incu- strategies) and reflected by different terminology, we bated in the marsupia of thiarid gastropods; for details will be able to also discern some of the potentially eco- and discussion see there. Accordingly, species with ovo- logical and evidently biogeographical correlations (see viviparous mode are termed as “rbp”-strategists, i.e. pos- below). sessing many eggs and veligers in the brood pouch and, In this context it is noteworthy to mention again the thus, having a high intrinsic rate of reproduction, in com- exceptional case of Melasma onca. Although we con- parison to those species with eu-viviparous mode that are sistently found shelled juveniles in the marsupium of termed “kbp”-strategists, i.e. possessing less juveniles in females in this species, these stages were less advanced their brood pouch and, thus, a lower intrinsic rate of re- and of smaller size compared to all other species listed production (albeit potentially a higher amount of mater- above in this category (see Figure 47). In addition, we nal investment). We suggest to apply these two terms also found that the number of embryonic stages in the mar- to the Australian thiarids, focussing here on the criterion supia of onca were, with ~ 200 juveniles, highest of the approximate number of embryonic stages within among all eu-viviparous thiarids (< 50) and intermedi- the marsupium as (the only available) proxy of reproduc- ate compared to those species listed above as ovo-vivi- tive rate, but not necessarily, of course, of reproductive parous (> 1000). Melasma onca occurs only in the success in terms of overall fitness, or accompanied to flood-drought dominated rivers of the “top end” of date by density dependency data. Northern Territory, where it often occurs not only sym- In general, we expect to find rbp-species under envir- patrical but syntopical with “Thiara” australis and Ser- onmental conditions where there is the need for a flex- myla venustula. It can only be speculated at this time ible and opportunistic mode of reproduction, such as that these facts might be a key towards an ecological e.g. under highly unpredictable conditions. As argued explanation (e.g. in terms of competitive interaction?) in Glaubrecht (1996), these conditions are, for example, of differences in reproductive modes and life history given for thiarids in or close to estuarine environments strategies of distinct thiarid species as further discussed as for example assumed for the ovo-viviparous Thiara below. amarula and several species of Stenomelania with an rbp-strategy. In contrast, the kbp-strategy is realized among eu-viviparous thiarids, like Melanoides or Tare- The concept of r-strategies versus K-strategies bia. In case of the Australian Thiaridae the rbp-species Within life history theory the rate of reproduction plays are relatively restricted in their geographic range, i.e. an eminent role, as it evolved in response to the impact the Jardinian province (with the exception of S. riqueti of different environments on the survival and fertility; that is discussed below as an occasional, opportunistic it is also constrained by genetic variance and evolution- marine invader to the northern coastal areas); for a gra- ary history (see e.g. Partridge & Harvey 1988). Thus, phic summary see Figure 49. In contrast, all kbp-spe- only within an evolutionary ecology theory the diversity cies have a less restricted, often widespread distribution of reproductive strategies is intelligible. In this context, on the Australian continent, again with the exception of a dichotomy in life history tactics have been described Melasma onca that is more restricted in the Northern for species that have early in their life a high number Territory. of offsprings versus those species that become fertile We are aware of the debates on MacArthur & Wil- later and give birth to less offspring. MacArthur & Wil- son’s (1967) original concept in context with the com- son (1967) have discussed several of these aspects and plex life history theory that can not be reviewed or re- coined a specific terminology (r- and K-selection), in- ferred to here in any depth desirable. Instead we itially used to compare colonizing with established po- suggest to use the two terms rbp and kbp only to de- pulations. Considering in particular density dependence, scribe certain biological phenomena found in thiarids these terms have been applied since then in various ap- which are reflected by them best. In remains to be ana- proaches to the study of r-selected species and K-se- lysed in more detail to what extend the rbp- and kbp-

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 267

Figure 49. Summary of the geographic ranges of eleven species of Thiaridae in Australia, according to their occurrences in fluvi- faunal provinces and major drainage systems, as documented in the present paper. The provinces and drainages are delineated as in Figure 3 (see introduction). Bars in black represent primary native occurrences as reconstructed here, with the status being either (e) – endemic to Australia, or (a) – authochthonous in Australia, but widely distributed elsewhere. Bars in grey mark the distribution of those taxa not primary native in Australia, i.e. being either (i) – possibly introduced, or (b) – possible occassional brackish water invasions only, or (c) – possibly cryptic species in Australia. Hatched bars for two taxa of the Leichhardtian mark only few occurrences also in the Carpentaria and Timor Sea drainages, respectively. For details and discussion see text.

strategies can be indeed correlated with the special en- discussed in the following as having quite distinct his- vironmental conditions found in the arid but also wet- torical contingencies; in addition, particularly their dis- dry or monsoonal rain influenced regions of the Austra- tinct reproductive strategies have important implications lian continent. It is open to future studies, for example, for the biogeographical patterns found in these Austra- whether these distinct reproductive strategies are corre- lian ‘marsupial’ freshwater gastropods. lated in those thiarid species being distributed in the Acknowledging the resulting regionalization as wet-dry tropics of northern Australia with the highly purely instructive for descriptive purpose we have in seasonal and variable (both spatially and annually) pre- this study integrated some of the various suggestions in cipitation, as discussed for vertebrates (Shine & Brown terms of Australian biogeographical areas through cor- 2008). It needs to be tested also if this developmental relating the major river drainage systems with fluvifau- plasticity of thiarid reproduction is either a reflection or nal provinces (see introduction and Figure 3). The maybe even the result (in terms of a flexible adaptive synthesis of our findings is summarized and visualized response) of the unpredictable but monsoonal rainfall in Figure 49, and will be discussed in the following in leading to stochastic flooding of rivers in Australia. terms of the most evident ecological and historical as- pects. Some striking biogeographical pattern in the Australian thiarids are, nevertheless, immediately ob- Biogeography vious; first of all the widespread occurrences of Plo- It was long known that several of the thiarid species tiopsis balonnensis and “Stenomelania” denisoniensis under study here not only occur on the Australian con- over vast areas of the continent. tinent but in other regions also. Nevertheless, this Ceri- At the same time most striking is the vicariant occur- thioidean freshwater family exhibit some remarkable rence of the conchologically often quite similar “Thiara endemisms at the generic and species level, since six australis” in the Leichhardtian province, restricted to out of eleven species (i.e. about half) of the Australian both the Timor and Gulf division, where balonnensis is Thiaridae only occur there, viz. “Thiara australis”, Plo- lacking (see Figure 48). We so far failed to record a tiopsis balonnensis and “Stenomelania” denisoniensis, sympatric and syntopic occurrence of these two thiarids which all exhibit a fairly wide distribution on the con- in Australia, although we were able to find both in the tinent, as well as Melasma onca, Sermyla venustula and same drainage (the Gilbert-Einasleigh system) in north- Ripalania queenslandica which have more restricted ern Queensland in 2007, albeit each species at opposite ranges (see Figure 49). In contrast, Thiara amarula and ends of this extensive river course. As obvious as this Stenomelania cf. aspirans as well as Melanoides tuber- distribution pattern presents itself now as a result of culata, Plotia scabra, and Sermyla riqueti are widely our mapping efforts and as interesting as it appears to distributed also outside Australia, in particular in the be, asking any dedicated biogeographer for an explana- Malay Archipelago and the Indo-West Pacific, respec- tion, we are currently unable to offer a convincing hy- tively. The occurrences of all these thiarid species are pothesis as to its origin and/or causation that would sur-

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 268 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae pass the many speculations possible, e.g. in terms of ern coastal region. Three areas are of particular interest mutual competitive exclusion. in the thiarid context here. Most of the ecological as- It is worth noting, though, that with Melasma onca pects relevant for Australian thiarids are not yet studied and Sermyla venustula two other Australian thiarids are in extensu, but some will be highlighted here from our also found to be restricted exclusively to the Leichhard- observations so far. tian. However, both species are more abundant either in (i) First, the northern part of the continent with its the Timor Sea drainages, as in case of onca (see under tropical climate of wet and hot summers associated the species and Figure 16), or endemic to the Carpen- with monsoonal rainfalls, and dry and warm winters, taria division in case of venusta (with only one excep- where rivers are mostly either permanent (see Fig- tion; see under the species and Figure 36). Thus, re- ures 2a–b), but often also cease to flow with a series of markably in addition to the ubiquitous deninsoniensis, isolated (partly large) pools within the dry river bed but just where balonnensis lacks, there are three other (see Figures 2d–e). Both in the Timor Sea and Gulf of thiarids endemic to the northern “top end” region of Carpentaria division the topography is dominated by the Australian continent. A comparable degree of ende- dissected tablelands and ridges, interspersed by flat al- misms among the Thiaridae is only found again in the luvial plains with permanent to intermittantly flowing Jardianian province (see Figure 49), albeit the three rivers and their associated wetlands. These have been thiarid taxa occurring there, apparently authochtho- termed “northern flood-drought rivers” by Williams & nously (see below), exhibit a much more restricted dis- Allen (1987: 188), concluding that “perhaps nowhere tribution in each case. else in Australia are seasonal hydrological conditions As discussed under the species, three thiarids known so important a determinant of the nature of aquatic en- to be widespread in the Indo-West Pacific and/or Orien- vironments”. In light of the restriction to the more tal region appear to be either introductions or invasions coastal areas found here in the distribution of the four to the Australian continent. Most evident, we hypothe- thiarid species typical to this region of Australia it sized (see details under the species) that Melanoides tu- should be noted that rainfall, which is mostly asso- berculata is not an Australian native but instead, as in ciated with monsoons and tropical cyclones, is large in so many other regions of the world, an anthropogenetic northern regions, with median annual value more than element to the freshwater fauna. Although the same can 1.500 mm, but less than 400 or only 300 mm in the be assumed for Plotia scabra it is impossible to decide south (Williams & Allen 1987: 188). with certainty yet on the status of this species; alterna- All four thiarids of the Leichhardtian province, viz. tively, it can not be excluded that scabra might be a the ubiquitous “Stenomelania” denisoniensis as well as cryptic species hitherto unknown to Australia but with the endemic “Thiara” australis, Melasma onca and Ser- a longer history on this continent. Given the scarcity of myla venustula, reproduce eu-viviparously, i.e. releasing records and our unsuccessful attempts during previous fairly advanced stages hatching as crawling juveniles. years to retrieve more records of Sermyla riqueti in Nevertheless, in particular the latter two exhibit a much Australia we hypothesize that this species only rarely more restricted distribution, even occurring essentially enters the lower courses of rivers in particular of the in only specific drainage systems like the Daly and/or northern region but also along the eastern coast (see Roper Rivers. In addition, as noted above, the later two Figure 30), facilitated by it possessing a mobile free appear largely vicariant toward each other, with the no- swimming veliger that in other parts of the Indo-West table exception of occurrences along the Roper River Pacific (see Figure 29) has allowed the wide dispersal system, though (compare Figures 16 and 36). While we with ocean currents and the colonization of various re- have no indication for differences in the life histories gions. of three of these thiarids, we already remarked above In the following we will investigate in more detail on the distinct features in some details of the reproduc- now only those eight native Australian Thiaridae, for tive biology of M. onca, which has many more but which we can reasonably assume that they are either smaller juvenile shells in its marsupium as compared to endemic to Australia (n ¼ 6) or have (regular) australis, venustula (see Figure 47) and also denisonien- authochthonous occurrences (n ¼ 2) on the continent. sis. We also found a remarkable case of correlation with individual occurrence and features of the hydrology in Ecological biogeography: correlation with climate case of Sermyla venustula, which is according to our and hydrology molecular analyses (Glaubrecht et al. unpubl. data) clo- sely related to the brackish water Sermyla riqueti. The Most obvious in Australia, with its either wet-tropical latter species in found in its SE Asian range (see Fig- or semi-arid to arid conditions, are the marked hydrolo- ure 29) to inhabit exclusively brackish water regions of gical conditions of many of its rivers in connection the estuaries and lower courses of rivers and streams with e.g. the unpredictability of precipitation, but also (Glaubrecht unpubl. data). We recorded a similar asso- the isolation, ephemerality and high salinity in particu- ciation with environments of elevated salinity also for lar of water bodies in the northern and central region, the Australian venustula, which is found mostly in lo- versus more reliable but heavy rainfall in the northeast- calities close to the coast and under marine influence,

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 269 as e.g. in the Gulf of Carpentaria region in the lower lia, it is only in this region of the continent where course of the Bynoe River and at Normanton (see Fig- we find conditions quite similar to those in other tro- ure 36), where it lives at least temporarily in brackish pical and monsoonal regions within the wide distribu- water conditions. At first sight the occurrence of S. ve- tional range of at least two of these three Jardinian nustula further inland, in particular along the Roper species (note the comment on endemism under Ripa- River, seems to contradict this finding. However, we lania queenslandica), such as e.g. the eastern Sunda found that among these localities especially Salt Creek Islands and Wallacea or northern New Guinea to the (Figures 2a–b) quite deserves its name, since there are Solomons. indications for highly increased salinity. Using meas- Developmental mode is often correlated directly with urements of the electrolytic conductivity (as a proxy geographic distribution and, for example, the relation- for salinity) taken for the Roper drainage in the course ship between planktonic larvae and extensive geo- of the “Top End Waterways Project” of the Australian graphic ranges seems well established for many animal Government we found that where Sermyla venustula groups. Nevertheless, Thiara amarula, Stenomelania as- occurs at Little Roper River (1357 mS/cm) and Salt pirans and Ripalania queenslandica confront us with a Creek (4240 mS/cm) the salinity was extremely high ap- biogeographical paradoxon and enigma, as we would parently due to salt dissolved from the sedimentary generally assume that those ovi-viviparous taxa which rocks in this region. Apparently, by virtue of this spe- release veliger from their brood pouch have larger dis- cial geological feature S. venustula is able to colonize tributional ranges with a cohesive gene pool (and are, suitable habitats even far inland. However, it should be thus, less prone to speciation in different allopatric re- noted that in particular for the Roper River we have to gions once dispersed there). However, these three rbp- assume also otherwise strong marine influences even in strategists exhibit the most restricted geographical inland courses that are fairly far from the coast, creat- ranges among the Australian Thiaridae. On the other ing special hydrological conditions for thiarids. hand, once swept down from the tributaries, lower parts (ii) A second area of particular interest is the semi- of the rivers or upper estuaries, presumably coinciding arid to arid central region of Australia, where rainfall with times of high rainfall, their marine veligers could is unreliable and considerable low with median annual cross even vast areas of the sea with ocean currents value less than 300 mm, and with its endorheic (inter- which follow the coast from the north. It is tempting, nal) drainages. Being by far the most widespread cli- therefore, to regard these thiarids as autochthonous spe- matic region of the continent, throughout there we find cies that, by virtue of their veligers being swept down often only seasonal or even episodic water-bodies as by ocean currents along the Cape York Peninsula from most characteristic and dominant limnological feature, more northern conspecific populations, managed to co- in particular the Lake Eyre and adjacent internally lonize and/or survive in what appears to be a relictual drained drainages (see Williams & Allen 1987 for de- area for many Australian faunal and floral elements tails). As the only thiarid in the Sturtian province (see (see introduction). On the other hand, it is contentious Figure 49) we record Plotiopsis balonnensis to be ex- at least to interprete the occurrence of these thiarids tant in highly isolated water-bodies along the Finke along the coastal rainforest pockets of Queensland as River system, where it survived apparently for long being relictual (see below). times in the only permanent deep pools that formed As is illustrated by these few observations we can where the rivers carve through the ridges of the Mac- expect a series of (pre-)adaptations in Australian Thia- Donnell Ranges (see under the species). Results from ridae that might have fascilitated colonization of and our ongoing molecular genetic analyses to be reported survival in the rivers and streams of northern Australia. elsewhere suggest a close affinity of all balonnensis po- As discussed e.g. by Williams & Allen (1987: 193) in pulations from Finke River and, surprisingly, with Wes- this context we have to remember the multidimensional tern Australian conspecifics instead of those from more nature of requisitory adaptations and should not consid- adjacent areas in QLD or NSW (Glaubrecht et al. un- er the many ecological but also historical factors in- publ. data.). volved only isolated, as they are more often to occur (iii) The third region of immediate interest is the contemporaneously and act in concert. northeastern coastal region, or Jardinian province, where summer rainfall is monsoonal and more reliable (in northern regions), with many, albeit relatively short Historical biogeography: “terra australis incognita” streams, that have high initial gradients, gradually de- and the origin of Australian Thiaridae creasing seawards, high annual discharges, seasonally variable or erratic flows and variable turbidity (Wil- The aim of historical biogeography is the study and ex- liams & Allen 1987: 187). The occurrence of three planation of the distribution of organisms in space and thiarids in this province (see Figure 49) appears to be time on the basis of historical events (see Glaubrecht correlated to areas of the highest rainfall and directly 1996, 2000). Two distinct explanatory approaches are connected to the climatic conditions to the east of the long under debate, viz. vicariance and dispersal, re- Great Dividing Range running in parallel to the east viewed e.g. by the latter author for limnic gastropods. coast. Therefore, in contrast to other parts of Austra- In the recent two decades, molecular phylogenetic stud-

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 270 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae ies have enormeously contributed to the renaissance of cestral thiarids like Thiara amarula, Stenomelania cf. this discipline as they often allow now to reconstruct aspirans and Ripalania queenslandica, as it was the the spatial pattern and temporal sequence of historical case for many other Miocene faunal elements of Aus- events and, thus, the causation of biogeography. For tralia (see e.g. Archer et al. 1991). On the other hand, only few freshwater organisms of Australia these meth- however, Ponder (1991) argued in case of the type ods have been consistently applied so far, in particular species of the hydrobiid genus Jardinella as the most for the crustacea (see literature cited in the introduc- derived taxon within this clade, that although asso- tion). For freshwater mollusca, unfortunately, we lack ciated with rainforests it is a relatively recent invader many of those studies; however, even in their absence from artesian springs in the east of the Great Dividing some cautious inferences can be made here for Thiari- Range. He found support for the hypothesis that J. dae. thaanumi crossed the range from the west and invaded While the origin of the freshwater mussel fauna is the coastal rivers that drained east. This apparent wes- still clouded and highly speculative, for freshwater tern derivation of the eastern species of Jardinella ap- snails four ancestral sources have been assumed accord- pears to be in contrast to the now widely accepted ing to Williams & Allen (1987: 190): Gondwana (as in idea that many of the Jardinian rainforest faunal ele- case of the Hydrobiidae), SE Asia (as in case of the ments are relictual; instead, he assumed that these lim- Lymnaeidae, Planorbidae and Viviparidae), the marine nic hydobiid snails could be a rather recent phenom- environment (obviously for Neritidae and Iravadiidae) enon in the fauna of NE Queensland. Obviously, the and anthropochorous introductions of exotics (such as existence of truely relictual species in the freshwater Physa among the Physidae). Thiaridae have found no gastropod fauna in this region remains equivocal for mentioning so far. However, based on the findings pre- the time being. sented here we propose that all four of the above given (iv) Most interestingly, nevertheless, we hypothesize sources are also found in these limnic Cerithioidea, as here that the long hold view of the thiarid fauna being summarized in Figure 49. a mere appendage to the SE Asian biota is false. With (i) We have mentioned above Melanoides tuberculata a new look at the considerable degree of endemisms in and potentially Plotia scabra to be most likely anthro- particular in the northern-central region of the Leich- pochorous introductions. hardtian, we find a peculiar and, possible highly specia- (ii) We have argued that the essentially brackish lised and adapted, thiarid fauna there that managed to water to estuarine Sermyla riqueti (as closest relative persist for long times apparently. With the thiarids of to the endemic S. venustula) certainly is in Australia Australia known now to exhibit some quite remarkable due to its connection with the marine environment. cases of distinct regionality and incidences of ende- Several currents in the Arafura Sea and the Coral Sea misms, we anticipate an Australian continental, i.e. facilitate occasional invasions by virtue of veligers into Gondwanan origin of these endemic Thiaridae. A the lower courses of rivers in northern (and eastern) southern continental origin of the Thiaridae as a clade Australia. has been proposed earlier by Glaubrecht (1996) when (iii) As not only near relatives but even conspecifics discussing the occurrences of these limnic Cerithioidea of Australian thiarids can be found over the entire SE in parts of former Gondwana. Asian and Pacific region, it is tempting to assume the Further support for this new biogeographical hypoth- ovo-viviparous Thiara amarula and Stenomelania cf. esis comes from rare fossils of Australia’s freshwater aspirans to be fairly recent colonizers from the north faunal elements. Recently, Hamilton-Bruce et al. (2004) that were able to maintain populations in restricted described a non-marine gastropod Melanoides godthelpi and relictual areas along the Jardinian. As a special from the Griman Creek Formation at Lightning Ridge case of interest remains Ripalania queenslandica, for in New South Wales of Early Cretaceous (Middle to which we to date lack positive indication of conspeci- Late Albian) age. The formation being regarded as of fics in other than the Australian region (see under the limnic origin, and this gastropod which appears quite species). However, certainly as an alternative view- similar to recent thiarids, suggest the occurrence of point, it should be considered that these three thiarids Thiaridae in Australia at a time when the continent was might represent an ancient Australian freshwater fau- far away from what is today Asia, but still closely con- nal element. As such they might have survived as re- nected to the Antarctic section of Gondwana. lictual forms in the Jardinian from times long gone (or even could have been the source for other Austra- lasian populations?). As noted in the introduction, the Conclusion rainforests in NE Queensland and their fauna, that to- day are generally considered ancient and providing re- In contrast to earlier authors, who have consistently hy- fuges for relictual species, are known to have been pothesized that the Australian Thiaridae are relatively much more abundant and widespread during the Mio- recent arrivals on this continent from the north, prefer- cene. These formerly continous rainforest habitats, as- ably from SE Asia and in particular the Malay Archipe- sociated with much higher rainfall at these times, lago (see Introduction), we suggest here that Australia could have provided suitable environments also for an- – and in particular its northern coastal wet-dry region

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 271 of the Leichhardtian province – holds a unique and en- References demic species assemblage of viviparous, i.e. “marsu- pial”, thiarid gastropods of other origin. Therefore, Abbott, R. T. 1948. Handbook of medically important molluscs of the herewith we like to promote the view that not only the orient and the western Pacific. – Bulletin of the Museum of marsupials among the mammals are to be hailed as pri- Comparative Zoology 100: 285–299. celess natural heritage among the Australian biota. This Aldridge, D. W. 1983. Physiological ecology of freshwater proso- continent “down under” also possesses some unique branchs. In Russel-Hunter, W. D. (ed.). The Mollusca, 6. Ecology. Academic Press, Orlando: pp. 329–358. freshwater lineages with peculiar elements that are en- Archer, M., Hand, S. J. & Godthelp, H. 1991. Riversleigh. The story demic on a continental scale, but also some species that of animals in ancient rainforests of inland Australia. Reed Books, are restricted to only certain regions in the tropical Kew VIC. north and even to certain river drainage systems. With Baker, H. G. 1955. Self-compatibility and establishment after ‘long- the hypothesis on Australia’s distinct nature supported distance’ dispersal. – Evolution 9(3): 347–349. where endemic ancient survivors and modern lineages Barker, F. K., Cibois, A. Schikler, P., Feinstein, J. & Cracraft, J. 2004. dovetale even Darwin’s much acclaimed ‘distinct Crea- Phylogeny and diversification of the largest avian radiation. – Proceedings of the National Academy of Sciences of the United tors at work’ finally falls into place. States of America 101(30): 11040–11045. Beck, R. M. 2009. Was the Oligo-Miocene Australian metatherian Yalkaparidon a ‘mammalian woodpecker’? – Biological Journal Acknowledgements of the Linnean Society 97: 1–17. Beheregaray, L. 2008. Twenty years of phylogeography: The state of M. G. thanks Winston F. Ponder (Sydney) for initially in 1996 having the field and the challenges for the Southern Henisphere. – Mole- suggested this “little side project” and for his various help along the cular Ecology 17: 3754–3774. way, including collecting thiarids in Arnhem Land in 2007. We are most Bell, G. 1982. The Masterpiece of Nature. The Evolution and Genet- grateful to Vince Kessner (Adelaide River), who has introduced us to ics of Sexuality. University of California Press, Berkeley, Los An- and later helped in many ways with field work in Northern Territory, geles. showing us all his most favorite collecting spots at the Victoria, Daly Ben-Ami, F. & Heller, J. 2005. Spatial and temporal patterns of and Roper Rivers, for being a wonderful host in Adelaide River over the parthenogenesis and parasitism in the freshwater snail Melanoides years, and for allowing us to use the data from his collection of Austra- tuberculata. – Journal of Evolutionary Biology 18: 138–146. lian thiarids. Special thanks are also in particular to Dr. Richard C. Ben-Ami, F. & Hodgson, A. N. 2005. Ovoviviparity and the structure Willan (Museum & Art Gallery of the Northern Territory, Darwin), as of the brood pouch in Melanoides tuberculata (Gastropoda: Pro- well as to Shirley Slack-Smith and Corey S.Whisson (Western Austra- sobranchia: Thiaridae). – Journal of Morphology 263: 322–329. lian Museum) and John Stanisic (Queensland Museum, Brisbane) for Benthem Jutting, T. van 1934. ber den Bau der Radula und ihre Be- help with field work, permits and the shipping of material. deutung fr die Nahrungsaufnahme bei einigen javanischen Sß- The initial work was aided by Visiting Research Fellowships from wassergastropoden. – Verhandlungen der Internationalen Vereini- the AMS to M. G. in 1996 and in 1998. Our field work in Australia gung fr theoretische und angewandte Limnologie 6: 325–330. was continuously supported by several grants from the Deutsche Stif- Benthem Jutting, W. S. S. van 1953. Annotated list of the Non-Marine tungszentrum (formerly Stifterverband fr die Deutsche Wissenschaft) Mollusca of the Islands Ambon, Haruku, Saparua and Nusa Laut. in Essen; we are in particular thankful to the Stemmler-Stiftung (for – Treubia 22 (2): 275–318. grants in 2002, 2004, 2005) and the Rudolf und Helene Glaser-Stif- Benthem Jutting, W. S. S. van 1956. Systematic studies on the non- tung (for a grant in 2007) as well as to Dr. Marilene Macher for ne- marine Mollusca of the Indo-Australian archipelago. 5. Critical gotiating these. We are also most grateful to the following persons for revision of the Javanese freshwater gastropods. – Treubia 23 (2): their help with collection and/or loan of material, hospitality and as- 259–477. sistance, for their encouragement, advice and for discussions: Chris- Benthem Jutting, W. S. S. van 1959. Catalogue of the non-marine tian Albrecht, Yves Finet, Vince Kessner, Ian Loch, Alison Miller, Mollusca of Sumatra and of its satellite islands. – Beaufortia 7 Timothy Page, Heidi-Ann Phl, Thomas von Rintelen, Martin Robin- (83): 41–191. son, Mark Salotti, Shirley Slack-Smith, Fred Wells, Corey S. Whisson, Berry, A. J. & Kadri, A. B. H. 1974. Reproduction of the Malay Richard Willan, as well as to Peter Carabot and Ernie Frei, who in freshwater cerithiacean gastropod Melanoides tuberculata. – Jour- December 1996 first collected Thiara amarula and Stenomelania as- nal of Zoology 172: 369–381. pirans alive in the Mowbray River, thus directing our later collecting Bishop, K. A. & Forbes, M. A. 1991. The freshwater fishes of northern efforts to this area. Australia. In Haynes, C. D., Ridpath, M. G. & Williams, M. A. J. After the initial material lists having been compiled by Andrea (eds). Monsoonal Australia. Landscape, ecology and man in the Glaubrecht in 1996 in the AMS, over the course of a decade Lee northern lowlands. A. A. Balkema, Rotterdam: pp. 79–108. Hsiang Liow, Sabine Schtt, Christine Zorn, Alban Gebler, Katharina Boles, W. E. 1995. The world’s oldest songbirds. – Nature 374: 21– Rieger, Viktor Hartung and Kathrin Podlacha helped at the ZMB with 22. the compilation and mapping of all samples’ locations; Silke Tenner Braby, M. F., Pierce, N. E. & Vila, R. 2007. Phylogeny and historical and Sophia Seidel assisted with some analyses of the reproductive biogeography of the subtribe Aporiina (Lepidoptera: Pieridae): strategies in Roper River thiarids, and Gabriele Drescher with SEM implications for the origin of Australian butterflies. – Biological works. Claudia Dames and Eva Patzschke helped in various ways dur- Journal of the Linnean Society 90: 413–440. ing the final preparation of the illustrations for this paper, as well as Brandt, R. A. M. 1974. The non-marine aquatic Mollusca of Thailand. two reviewers with comments on the manuscript. – Archiv fr Molluskenkunde 105(1/4): 1–423. M. G. designed the project and together with N. B. conducted the Brot, A. 1874–1877. Die Melaniaceen (Melanidae) in Abbildungen field work, taxonomic identifications and geographical analyses; he nach der Natur mit Beschreibungen. In Martini, F. H. W. & Chem- finally also wrote the paper. N. B. undertook the morphological (and nitz, J. H. (eds). Systematisches Conchylien-Cabinet. vol. 1 (24). molecular) analyses and wrote a draft version of the paper. J. P. was Bauer & Raspe, Nrnberg: 488 pp. responsible for the final compilation of data and maps and assisted Brown, D. S. 1994. Freshwater snails of Africa and their medical im- with the figures and in writing the manuscript. portance. Taylor & Francis, London.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 272 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

Carini, G. & Hughes, J. M. 2006. Subdivided population structure and Glaubrecht, M. 2003. Arten, Artkonzepte und Evolution. Was sind phylogeography of an endangered freshwater snail, Notopala sub- und wie entstehen „biologische Arten“? In Reichholf, J. (ed.). lineata (Conrad, 1850) (Gastropoda: Viviparidae), in Western Biologische Vielfalt – Sammeln, Sammlungen, Systematik. Rund- Queensland, Australia. – Biological Journal of the Linnean So- gesprche der Kommission fr kologie, 26. Verlag Dr. Friedrich ciety 88: 1–16. Pfeil, Mnchen: pp. 15–42. Carini, G., Hughes, J. M. & Bunn, S. E. 2006. The role of waterholes Glaubrecht, M. 2004. Leopold von Buch’s legacy: treating species as as “refugia” in sustaining genetic diversity and variation of two dynamic natural entities, or why geography matters. – American freshwater species in dryland river systems (Western Queensland, Malacological Bulletin 19 (1/2): 111–134. Australia). – Freshwater Biology 51: 1434–1446. Glaubrecht, M. 2006. Independent evolution of reproductive modes in Colgan, D. J. & Ponder, W. F. 2000. Incipient speciation in aquatic viviparous freshwater Cerithioidea (Gastropoda, Sorbeoconcha) – snails in an arid-zone spring complex. – Biological Journal of the a brief review. – Basteria 69 (suppl. 3): 28–32. Linnean Society 71: 625–641. Glaubrecht, M. 2008. Adaptive radiation of thalassoid gastropods in Cunnington, W. A. 1920. The fauna of the African Lakes: a study in Lake Tanganyika, East Africa: morphology and systematization of comparative limnology with special reference to Tanganyika. – a paludomid species flock in an ancient lake. – Zoosystematics Proceedings of the Zoological Society of London 90: 507–622. and Evolution 84 (1): 71–122. Darlington, P. J. 1965. Biogeography of the Southern end of the Glaubrecht, M. 2009. On “Drawinian Mysteries” or molluscs as mod- world. Harvard University Press, Cambridge, Mass. els in evolutionary biology: from local speciation to global radia- Davis, G. M. 1982. Historical and ecological factors in the evolution, tion. – American Malacological Bulletin 27: 3–23. adaptive radiation, and biogeography of freshwater mollusks. – Glaubrecht, M. & Khler, F. 2004. Radiating in a river: systematics, American Zoologist 22: 375–395. molecular genetics and morphological differentiation of vivipar- de Bruyn, M., Wilson, J. C. & Mather, P. B. 2004. Reconciling geo- ous freshwater gastropods endemic to the Kaek River, Central graphy and genealogy: phylogeography of giant freshwater prawns Thailand (Cerithioidea, Pachychilidae). – Biological Journal of from the Lake Carpentaria region. – Molecular Ecology 13: the Linnean Society 82: 275–311. 3515–3526. Glaubrecht, M. & Rintelen, T. von 2003. Systematics, molecular ge- Dillon, R. T. 2000. The ecology of freshwater molluscs. Cambridge netics, and historical zoogeography of the viviparous freshwater University Press, Cambridge. gastropod Pseudopotamis (Cerithioidea, Pachychilidae): a relic on Edwards, S. V. & Boles, W. E. 2002. Out of Gondwana: the origin of the Torres Strait Islands, Australia. – Zoologica Scripta 32 (2): passerine birds. – Trends in Ecology and Evolution 17: 347–349. 415–435. Facon, B., Pointier, J. P., Glaubrecht, M. Poux, C. Jarne, P. & David, Glaubrecht, M. & Rintelen, T. von 2008. The species flocks of la- P. 2003. A molecular phylogeography approach to biological inva- custrine gastropods: Tylomelania on Sulawesi as models in spe- sions of the New World by parthenogenetic thiarid snails. – Mo- ciation and adaptive radiation. (Proceedings of the “Speciation lecular Ecology 12: 3027–3039. in Ancient Lake IV” Symposium, Berlin). – Hydrobiologia 615: Filardi, C. E. & Moyle, R. G. 2005. Single origin of a pan-Pacific 181–199. brid group and upstream colonization of Australasia. – Nature Glaubrecht, M. & Strong, E. E. 2004. Spermatophores of thalassoid 438: 216–219. gastropods (Paludomidae) in Lake Tanganyika, East Africa, with Fischer, P. H. & Crosse, J. C. H. 1891–92. Pour servir l’Histoire de a survey of their occurrence in Cerithioidea: functional and phylo- la faune de l’Amerique centrale et du mexique. Vol. 2. Imprimerie genetic implications. – Invertebrate Biology 123 (3): 218–236. Nationale, Paris. Grateloup, M. D. 1840. Memoire sur plusieurs espces de coquilles Flannery, T. F. 1994. The future eaters. An ecological history of the nouvelles ou peu connues. Mollusques. Lafargue, Libraire, Bor- Australasian lands and people. Reed Books, Chatswood NSW. deaux. Fretter, V. 1984. Prosobranchs. In Tompa, A. S., Verdonk, N. H. & Hamilton, W., Axelrod, R & Tanese, R. 1990. Sexual selection as an van den Biggelaar, J. A. M. (eds). The Mollusca, 7. Reproduction. adaptation to resist parasites. – Proceedings of the National Acad- Acadamic Press, Orlando: pp. 1–45. emy of Sciences of the United States of America 87: 3566–3573. Fretter, V. & Graham, A. 1994. British Prosobranch Molluscs. Their Hamilton-Bruce, R. J., Kear, B. P. & Smith, B. J. 2004. A new non- functional anatomy and ecology. The Ray Society, Andover, London. marine Early Cretaceous gastropod species from the Lightning Fulton, H. C. 1908. A list of species of shells described by Dr. Grate- Ridge, New South Wales. – Alcheringa 28: 485–492. loup, with critical notes. – Proceedings of the Malacological So- Haynes, C. D., Ridpath, M. G. & Williams, M. A. J. 1991. Monsoonal ciety of London 8: 43–44. Australia, landscape, ecology and man in the northern lowlands. Germain, L. 1921. Mollusques terrestres et fluviatiles de Syrie. Voy- A. A. Balkema, Rotterdam. age zoologique d’ Henri Gadeau de Kerville en Syrie. 1. Introduc- Heinsohn, R. 2008. Life in the Cape York Rainforest. CSIRO Publish- tion et gastropodes. Baillire, Paris. ing, Collingwood VIC. Glaubrecht, M. 1993. Mapping the diversity: geographical distribution Heller, J. & Farstey, V. 1990. Sexual and parthenogenetic populations of the freshwater snail Melanopsis (Gastropoda: Cerithioidea: of the freshwater snail Melanoides tuberculata in Israel. – Israel Melanopsidae) with focus on its systematics in the Mediterranean Journal of Zoology 37: 75–87. Basin. – Mitteilungen aus dem Hamburger Zoologischen Museum Hesse, P. P., Magee, J. W. & van der Kaars, S. 2004. Late Quaternary und Institut 90: 41–97. climates of the Australian arid zone: a review. – Quaternary Inter- Glaubrecht, M. 1996. Evolutionskologie und Systematik am Beispiel national 118–19: 87–102. von Sß- und Brackwasserschnecken (Mollusca: Caenogastropoda: Holznagel, W. E. 1998. A nondestructive method for cleaning gastro- Cerithioidea): Ontogenese-Strategien, palontologische Befunde pod radulae from frozen, alcohol-fixed, or dried material. – und Historische Zoogeographie. Backhuys Publishers, Leiden. American Malalogical Bulletin 14 (2): 181–183. Glaubrecht, M. 1999. Systematics and the evolution of viviparity in Houbrick, R. S. 1988. Cerithiodean phylogeny. – Malacological Re- tropical freshwater gastropods (Cerithioidea: Thiaridae sensu lato) view, Suppl. 4: 88–128. – an overview. – Courier Forschungsinstitut Senckenberg 215: Hubendick, B. 1955. On a small material of fresh water molluscs col- 91–96. lected by Prof. T. Gislen in Australia. – Lunds Universitets Aars- Glaubrecht, M. 2000. A look back in time: Toward an historical bio- skrift 51 (6): 1–11. geography as synthesis of systematic and geologic patterns out- Hubendick, B. 1962. Aspects on the diversity of the fresh-water fau- lined with limnic gastropods. – Zoology 102: 127–147. na. – Oikos 13 (2): 249–261.

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 273

Hugall, A. F. & Lee, M. S. Y. 2004. Molecular claims of Gondwana Lively, C. M. & Jokela, J. 2002. Temporal and spatial distributions of age for Australian agamid lizards are untenable. – Molecular parasites and sex in a freshwater snail. – Evolutionary Ecology Biology and Evolution 21: 2102–2110. Research 4: 219–226. Hugall, A. F., Foster, R., Hutchinson, M. & Lee, M. S. Y. 2008. Phy- Livshits, G. & Fishelson, L. 1983. Biology and reproduction of the logeny of Australasian agamid lizards based on nuclear and mito- freshwater snail Melanoides tuberculata (Gastropoda: Prosobran- chondrial genes: implications for morphological evolution and chia) in Israel. – Israel Journal of Zoology 32: 21–35. biogeography. – Biological Journal of the Linnean Society 93: Livshits, G., Fishelson, L. & Wise, G. S. 1984. Genetic similarity and 343–358. diversity of parthenogenetic and bisexual populations of the fresh- Iredale, T. 1943. A basic list of the fresh water mollusca of Australia. water snail Melanoides tuberculata (Gastropoda: Prosobranchia). – – The Australian Zoologist 10 (2): 118–209. Biological Journal of the Linnean Society 23: 41–54. Iredale, T. & Whitley, G. P. 1938. The fluvifaunulae of Australia. – Lydeard, C., Holznagel, W. E., Glaubrecht, M. & Ponder, W. F. 2002. South Australian Naturalist 18: 64–68. Molecular phylogeny of a circum-global, diverse gastropod super- Jacob, J. 1957. Cytological studies of Melaniidae (Mollusca) with spe- family (Cerithioidea: Mollusca: Caenogastropoda): pushing the cial reference to parthenogenesis and polyploidy. I. Oogenesis of deepest phylogenetic limits of mitochondrial LSU rDNA se- the parthenogentic species of Melanoides (Prosobranchia-Gastro- quences. – Molecular Phylogenetics and Evolution 22: 399–406. poda). – Transactions of the Royal Society of Edinburgh 63: MacArthur, R. H. & Wilson, E. O. 1967. The theory of Island Bio- 341–352. geography. Monographs in Population Biology. Princeton Univer- Jacob, J. 1958. Cytological studies of Melaniidae (Mollusca) with sity Press, Princeton. special reference to parthenogenesis and polyploidy. II. A study Mackey, B. G., Berry, S. L. & Brown, T. 2008. Reconciling ap- of meiosis in the rare males of the polyploid race of M. tubercu- proaches to biogeographical regionalization: a systematic and gen- lata and M. lineatus. – Transactions of the Royal Society of Edin- eric framework examined with a case study of the Australian con- burgh 63: 433–444. tinent. – Journal of Biogeography 35: 213–229. Jokela, J. & Lively, C. M. 1995. Parasites, sex, and early reproduction Markgraf, V., McGlone, M. & Hope, G. 1995. Neogene paleoenviron- in a mixed population of freshwater snails. – Evolution 49: mental and paleoclimatic change in southern temperate ecosys- 1268–1271. tems – a southern perspective. – Trends in Ecology and Evolu- Jokela, J., Lively, C. M., Dybdahl, M. F. & Fox, J. A. 2003. Genetic tion 10: 143–147. variation in sexual and clonal lineages of a freshwater snail. – Martens, E. von 1883. Die Weich- und Schaltiere gemeinfaßlich dar- Biological Journal of the Linnean Society 79: 165–181. gestellt. G. Freytag, Leipzig. Jonsson, K. A., Irestedt, M., Fuchs, J., Ericson, P. G. P., Christidis, L., Martin, H. A. 1978. Evolution of the Australian flora and vegetation Bowie, R. C. K., Norman, J. A., Pasquet, E. & Fjeldsa, J. 2008. through the Tertiary: evidence from pollen. – Alcheringa 2: 181– Explosive avian radiation and multi-directional dispersal across 202. Wallacea: evidence from the Campephagidae and other crown Martin, H. A. 1998. Tertiary climatic evolution and the development Corvida (Aves). – Molecular Phylogenetics and Evolution 47: of aridity in Australia. – Proceedings of the Linnean Society of 221–236. New South Wales 119: 115–136. Kearney, M. 2003. Why is sex so unpopular in the Australian desert? Maynard Smith, J. 1978. The evolution of sex. Cambridge University – Trends in Ecology and Evolution 18: 605–607. Press, Cambridge. Kearney, M. 2005. Hybridization, glaciation and geographical parthe- Mayr, E. 1942. Systematics and the origin of species. Columbia Uni- nogenesis. – Trends in Ecology and Evolution 20: 495–502. versity Press, New York. Keast, A. (ed.) 1981. Ecological Biogeography of Australia. Dr. Mayr, E. 1944. Timor and the colonization of Australia by birds. – W. Junk, The Hague, Boston. The Emu 44: 113–130. Keynes, R. D. (ed.) 1988. Charles Darwin’s “Beagle Diary”. Cam- McMichael, D. F. 1967. Australian freshwater mollusca and their bridge University Press, Cambridge. probable evolutionary relationships: a summary of present knowl- Khler, F. & Glaubrecht, M. 2001. Toward a systematic revision of the edge. In Weatherly, A. H. (ed.). Australian Inland Waters and their Southeast Asian freshwater gastropod Brotia H. Adams, 1866 (Ceri- fauna. National University Press, Canberra: pp. 123–149. thioidea: Pachychilidae): an account of species from around the McMichael, D. F. & Hiscock, I. D. 1958. A monograph of the fresh- South China Sea. – Journal of Molluscan Studies 67: 281–318. water mussels (Mollusca: Pelecypoda) of the Australian region. – Khler, F. & Glaubrecht, M. 2003. Morphology, reproductive biology Australian Journal of Marine and Freshwater Research 9: 327– and molecular genetics of ovoviviparous freshwater gastropods 507. (Cerithioidea, Pachychilidae) from the Philippines, with descrip- Merrick, J. R. & Schmida, G. E. 1984. Australian freshwater fishes: tion of a new genus Jagora. – Zoologica Scripta 32 (1): 35–59. biology and managment. Griffith Press, Netley. Khler, F. & Glaubrecht, M. 2006. A systematic revision of the Miller, A. C., Ponder, W. F. & Clark, S. 1999. Freshwater snails of the Southeast Asian freshwater gastropod Brotia (Cerithioidea: Pachy- genus Fluvidona and Austropyrgus (Gastropoda: Hydrobiidae) chilidae). – Malacologia 48 (1): 159–251. from northern New South Wales and southern Queensland, Aus- Khler, F., Brinkmann, N. & Glaubrecht, M. 2008. Convergence tralia. – Invertebrate Taxonomy 13: 461–493. caused confusion: on the systematics of the freshwater gastropod Morrison, J. P. E. 1954. The relationship of old and new world mela- Sulcospira pisum (Brot, 1868) (Cerithioidea, Pachychilidae). – nians. – Proceedings of the United States National Museum 103: Malacologia 50 (1–2): 331–339. 357–394. Khler, F., Rintelen, T. von, Meyer, A. & Glaubrecht, M. 2004. Multi- Muley, E. V. 1977. Studies on the breeding habits and development of ple origin of viviparity in Southeast Asian gastropods (Cerithioi- the brood-pouch of a viviparous prosobranch, Melania scabra. – dea: Pachychilidae) and its evolutionary implications. – Evolution Hydrobiologia 54: 181–185. 58 (10): 2215–2226. Murphy, N. P. & Austin, C. M. 2004a. Phylogenetic relationships of Kondrashov, A. S. 1993. Classification of hypotheses on the advan- the globally distributed freshwater prawn genus Macrobrachium tage of amphimixis. – The Journal of heredity 84 (5): 372–87. (Crustacea: Decapoda: Palaemonidae): biogeography, taxonomy Lake, J. S. 1971. Freshwater fishes and rivers of Australia. Thomas and the convergent evolution of abbreviated larval development. – Nelson Australia, Melbourne. Zoologica Scripta 34: 187–197. Lively, C. M. 1987. Evidence from a New Zealand snail for the main- Murphy, N. P. & Austin, C. M. 2004b. Phylogeography of the wide- tenance of sex by parasitism. – Nature 328: 519–521. spread Australian freshwater prawn, Macrobrachium australiense

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 274 Glaubrecht,M.etal.:SystematicsofAustralianThiaridae

(Decapoda, Palaemonidae). – Journal of Biogeography 31: 1065– Ponder, W. F., Hershler, R. & Jenkins, B. 1989. An endemic radiation 1072. of hydrobiid snails from artesian springs in northern South Aus- Myers, M. J., Meyer, C. & Resh, V. H. 2000. Neritid and thiarid gas- tralia: their taxonomy, physiology, distribution and anatomy. – tropods from french polynesian streams: how reproduction (sex- Malacologia 31: 1–140. ual, parthenogenetic) and dispersal (active, passive) affect popula- Ramamoorthi, K. 1950. The brood-pouch in the viviparous Melanii- tion structure. – Freshwater Biology 44: 535–545. dae (Mollusca, Gastropoda). – Journal of the Zoological Society Nation, J. L. 1983. A new method using hexamethyldisilazane for of India 2 (1): 27–33. preparation of soft insect tissues for scanning electron micro- Reid, G. D. 1990a. A cladistic phylogeny of the genus Littorina (Gas- scopy. – Stain Technology 53 (6): 347 – 351. tropoda): implications for evolution of the reproductive strategies Nguyen, T. T. T., Austin, C. M., Meewan, M. M., Schultz, M. B. & and for classification. – Hydrobiologia 193: 1–19. Jerry, D. R. 2004. Phylogeography of the freshwater crayfish Rensch, B. 1929. Das Prinzip geographischer Rassenkreise und das Cherax destructor Clark (Parastacidae) in inland Australia: histor- Problem der Artbildung. Borntraeger, Berlin. ical fragmentation and recent range expansion. – Biological Jour- Rensch, B. 1934. Sßwasser-Mollusken der Deutschen Limnolo- nal of the Linnean Society 83: 539–550. gischen Sunda-Expedition. – Archiv fr Hydrobiologie, Suppl. Nicholas, F. W. & Nicholas, J. M. 1989. Charles Darwin in Australia. 13: 3–53. Cambridge University Press, Cambridge. Riech, E. 1937. Systematische, anatomische, kologische und tiergeo- Ospovat, D. 1981. The development of Darwin’s theory. Cambridge graphische Untersuchungen ber die Sßwassermollusken Papua- University Press, Cambridge. siens und Melanesiens. – Archiv fr Naturgeschichte 6: 37–153. Pace, G. L. 1973. The freshwater snails of Taiwan (Formosa). – Ma- Rintelen, T. von & Glaubrecht, M. 2005. The anatomy of an adaptive lacological Review Suppl. 1: 3–118. radiation: a unique reproductive strategy in the endemic fresh- Page, T., Rintelen, K. von & Hughes, J. M. 2007. An island in the water gastropod Tylomelania (Cerithioidea: Pachychilidae) on Su- stream: Australia’s place in the cosmopolitan world of Indo-West lawesi, Indonesia, and its biogeographic implications. – Biologi- Pacific freshwater shrimp (Decapoda: Atyidae: Caridina). – Mo- cal Journal of the Linnean Society 85: 513–542. lecular Phylogenetics and Evolution 43: 645–659. Samadi, S., Artiguebielle, E., Estoup, A., Pointier, J.-P., Silvain, J.-F., Partridge, L. & Harvey, P. 1988. The ecological context of life history Heller, J., Cariou, M.-L. & Jarne, P. 1998. Density and varibility of evolution. – Science 241: 1449–1455. dinukleotide microsatellites in the parthenogenetic polyploid snail Pilsbry, H. A. & Bequaert, J. 1927. The aquatic mollusks of the Bel- Melanoides tuberculata. – Molecular Ecology 7: 1233–1236. gian Congo, with a geographical and ecological account of Congo Sarasin, P. & Sarasin, F. 1898. Die Sßwassermollusken von Celebes. malacology. – Bulletin of the American Museum of Natural His- Beitrge zur Naturgeschichte der Insel Celebes. Kreidel, Wiesbaden. tory 53: 69–602. Schtt, S. & Glaubrecht, M. 1999. Thiara amarula (Linn, 1758) (Cae- Pointier, J. P., Incani, R. N., Balzan, C., Chrosciechowski, P. & Pryp- nogastropoda: Thiaridae) in Australia – new evidence on the anato- chan, S. 1994. Invasion of the rivers of the littoral central region my of the reproductive system in a viviparous freshwater mollusc. – of Venezuela by Thiara granifera und Melnoides tuberculata Courier Forschungs-Institut Senckenberg 215: 181–188. (Mollusca: Prosobranchia: Thiaridae) and the absence of Biom- Schultz, M. B., Smith, S. A., Horwitz, P., Richardson, A. M. M., phalaria glabrata, snail host of Schistosoma mansoni. – Nautilus Crandall, K. A. & Austin, C. M. 2009. Evolution underground: A 107 (4): 124–128. molecular phylogenetic investigation of Australian burrowing Ponder, W. F. 1991. The eastern seaboard species of Jardinella (Mol- freshwater crayfish (Decapoda: Parastacidae) with particular focus lusca, Gastropoda, Hydrobiidae), Queensland rainforest-inhabiting on Engaeus Erichson. – Molecular Phylogenetics and Evolution freshwater snails derived from the west. – Records of the Austra- 50: 580–598. lian Museum 43: 275–289. Seshaiya, R. V. 1936. Notes on the comparative anatomy of some In- Ponder, W. F. & Clark, G. A. 1990. A radiation of hydrobiid snails in dian Melaniidae. – Journal Annamalai University 5 (2): 167–182. threatened artesian springs in western Queensland. – Records of Seshaiya, R. V. 1940. A free larval stage in the life history of a flu- the Australien Museum 42: 301–363. viatile gastropod. – Current Science 9 (7): 331–332. Ponder, W. F. & Colgan, D. J. 2002. What makes a narrow-range tax- Shine, R. & G. P. Brown 2008. Adapting to the unpredictable: repro- on? Insights from Australian freshwater snails. – Invertebrate ductive biology of vertebrates in the Australian wet-dry tropics. – Systematics 16: 571–582. Philosophical Transactions of the Royal Society 363: 363–373. Ponder, W. F. & Lindberg, D. R. 1997. Towards a phylogeny of gastro- Smith, B. J. 1992. Non-marine Mollusca. In Houston, W. W. K. (ed.). pod molluscs – an analysis using morphological characters. – Zo- Zoological Catalogue of Australia, 8. Australian Government, ological Journal of the Linnean Society 119: 83–265. Canberra: pp. 75–79. Ponder, W. F., Clark, G. A., Miller, A. C. & Toluzzi, A. 1993. On Smith, B. J. 1996. Identification keys to the families and genera of major radiation of freshwater snails in Tasmania and eastern Vic- bivalve and gastropod molluscs found in Australian inland waters. toria: a preliminary overview of the Beddomeia group (Mollusca: Co-operative Research Centre for Freshwater Ecology, Albury. Gastropoda: Hydrobiidae). – Invertebrate Taxonomy 7: 501–750. Smith, E. A. 1882. On the freshwater shells of Australia. – Journal of Ponder, W. F., Colgan, D. J., Clark, G. A., Miller, A. C. & Terzis, T. the Linnean Society (Zoology) 16: 255–317. 1994. Microgeographic, genetic and morphological differentiation Starmhlner, F. 1969. Die Gastropoden der madagassischen Binnenge- of freshwater snails – the Hydrobiidae of Wilsons Promontory, wsser. – Malacologia 8 (1/2): 1–434. Victoria, south-eastern Australia. – Australian Journal of Zoology Starmhlner, F. 1973. Die Gattung Melanopsis Ferussac 1807 auf 42: 557–678. Neukaledonien. – Malacologia 14: 242–243. Ponder, W. F., Colgan, D. J., Terzis, T., Clark, S. A., & Miller, A. C. Starmhlner, F. 1976. Ergebnisse der sterreichischen Indopazifik- 1996. Three new morphologically and genetically determined spe- Expedition 1971 des 1. Zoologischen Insitutes der Universitt cies of hydrobiid gastropods from Dalhousie Springs, northern Wien: Beitrge zur Kenntnis der Sßwassergastropoden pazi- South Australia, with the description of a new genus. – Mollus- fischer Inseln. – Annalen des Naturhistorischen Museums in can Research 17: 49–109. Wien 80 B: 473–656. Ponder, W. F., Eggler, P. & Colgan, D. J. 1995. Genetic differentiation Starmhlner, F. 1983. Results of the Hydrobiological Mission 1974 of of aquatic snails (Gastropoda: Hydrobiidae) in artesian springs in the Zoological Institute of the University of Vienna; Part VIII: arid Australia. – Biological Journal of the Linnean Society 56: Contributions to the knowledge of the freshwater-gastropods of 553–596. the Indian Ocean Islands (Seychelles, Comores-, Mascarene-Ar-

museum-zoosyst.evol.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 85 (2) 2009, 199–275 275

chipelagos). – Annalen des Naturhistorischen Museums Wien 84 Thiele, J. 1929. Handbuch der Systematischen Weichtierkunde. Vol. 1. B: 127–249. Gustav Fischer, Jena. Starmhlner, F. 1984. Results of the Austrian-Indian hydrobiological Troschel, F. H. 1856–1863. Das Gebiss der Schnecken zur Begrn- mission 1976 to the Andaman-Islands; Part IV: the freshwater dung einer natrlichen Classification. Nicolaische Verlagsbuch- gastropods of the Andaman-Islands. – Annalen des Naturhisto- handlung. Berlin. rischen Museums Wien 86 B: 145–204. Unmack, P. J. 2001. Biogeography of Australian freshwater fishes. – Starmhlner, F. 1993. Ergebnisse der sterreichischen Tonga-Samoa Journal of Biogeography 28: 1053–1089. Expedition 1985 des Instituts fr Zoologie der Universitt Wien: Webber, H. H. 1977. Gastropoda: Prosobranchia. In Giese, A. C. & Veitrge zur Kenntnis der Sß- und Brackwasser-Gastropoden der Pearse, J. S. (eds). Reproduction of marine invertebrates, 4. Aca- Tonga- und Samoa- Inseln (SW-Pazifik). – Annalen des Naturhis- demic Press, New York. torischen Museums in Wien 94–95 B: 217–306. Wenz, W. 1938–1944. Gastropoda. Teil 1: Allgemeiner Teil und Pro- Stoddart, J. A. 1983. The accumulation of genetic variation in a sobranchia. In Schindewolf, O. H. (ed.). Handbuch der Palozoo- parthenogenetic snail. – Evolution 37: 546–554. logie 6 (1). Borntraeger, Berlin: pp. 1–1639. Stoddart, J. A. 1985. Analysis of species lineages of some Australian Westerlund, C. A. 1892. Fundamenta Malacologia. Hkan Ohlsson, thiarids (Thiaridae, Prosobranchia, Gastropoda) using the evolu- Lund. tionary species concept. – Journal Malacological Society of Aus- White, M. E. 1986. The greening of Gondwana. The 400 million year tralia 7(1–2): 7–16. story of Australia’s plants. Kangaroo Press, East Roseville NSW. Sunderbrink, O. 1929. Zur Frage der Verwandtschaft zwischen Mela- White, M. E. 1994. After the greening. The browning of Australia. niiden und Cerithiiden. – Zeitschrift fr Morphologie und kolo- Kangaroo Press, Kenthurst NSW. gie der Tiere 14: 261–337. Whitley, G. P. 1947. The fluvifaunulae of Australia with particular Tenison-Woods, J. E. 1888. On the anatomy and life history of Mol- reference to freshwater fishes in Western Australia. – Western lusca peculiar to Australia. –Transactions of the Royal Society of Australian Naturalist 1: 49–53. New South Wales 1888: 106–187. Whitley, G. P. 1959. The freshwater fishes of Australia. – Monogra- Thacker, C. E., Unmarck, P. J., Matsui, L. & Rifenbark, N. 2007. phiae Biologicae 8: 136–149. Comparative phylogeography of five sympatric Hypseloetris spe- Wiley, E. O. 1978. The evolutionary species concept reconsidered. – cies (Teleostei: Eleotridae) in south-eastern Australia reveals a Systematic Zoology 27: 17–26. complex pattern of drainage basin exchange with little congru- Wiley, E. O. 1981. Phylogenetics. The theory and practice of Phyloge- ence across species. – Journal of Biogeography 34: 1518–1533. netic Systematics. Wiley, New York. Thacker, C. E., Unmarck, P. J., Matsui, L., Duong, P. & Huang, E. Williams, G. C. 1975. Sex and Evolution. Princeton University Press. 2008. Phylogeography of Philypnodon species (Teleostei: Eleotri- Williams, W. D. & Allen, G. R. 1987. Origins and adaptations of the dae) across south-eastern Australia: testing patterns of connectiv- fauna of inland waters. In Dyne, G. R. & Walton, D. W. (eds). ity across drainage divides and among coastal rivers. – Biological Fauna of Australia, vol. 1A. Australian Government Publishing Journal of the Linnean Society 95: 175–192. Service, Canberra: pp. 184–201. Thiele, J. 1928. Revision des Systems der Hydrobiiden und Melanii- Willmann, R. 1985. Die Art in Raum und Zeit. Das Artkonzept in der den. – Zoologische Jahrbcher 55: 351–402. Biologie und Palontologie. Parey, Berlin, Hamburg.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de