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A remarkable radiation of hydrobiids in the caves and streams at Precipitous Bluff, south west Tasmania (Mollusca: Caenogastropoda: Hydrobiidae)

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Title:
A remarkable radiation of hydrobiids in the caves and streams at Precipitous Bluff, south west Tasmania (Mollusca: Caenogastropoda: Hydrobiidae)
Series Title:
Subterranean Ecology
Creator:
Eberhard, Stefan
Clark, S.A.
Ponder, W.F.
Studdert, J.B.
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Magnolia Press
Subterranean Ecology, Scientific Environmental Services
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English

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Biology ( local )
Cave Ecology ( local )
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Australia

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General Note:
A radiation of hydrobiid snails is described from caves at Precipitous Bluff, southern Tasmania. The radiation (10 species) comprises two closely related genera; Pseudotricula, endemic to the Precipitous Bluff caves, and Nanocochlea found in these caves and surface streams and seepages, but also known from elsewhere in southern Tasmania. Two Precipitous Bluff species of Nanocochlea inhabit streams outside the caves, while one is found within the caves; all three are new. Seven species of Pseudotricula are described, six being new. Up to five species, one Nanocochlea and four Pseudotricula, are found in single stream habitats characterised by diverse sediment and slow to moderate flows. One Pseudotricula species is found in a swifter, larger stream with cobbles and gravel and another on smooth limestone surfaces in fast flowing water and water falls. A cladistic analysis (based on morphology) confirmed the monophyly of Pseudotricula, but the monophyly of Nanocochlea is less robust. -- Authors
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K26-00131 ( USFLDC DOI )
k26.131 ( USFLDC Handle )
4262 ( karstportal - original NodeID )

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Description
A radiation of
hydrobiid snails is described from caves at Precipitous Bluff,
southern Tasmania. The radiation (10 species) comprises two
closely related genera; Pseudotricula, endemic to the
Precipitous Bluff caves, and Nanocochlea found in these caves
and surface streams and seepages, but also known from elsewhere
in southern Tasmania. Two Precipitous Bluff species of
Nanocochlea inhabit streams outside the caves, while one is
found within the caves; all three are new. Seven species of
Pseudotricula are described, six being new. Up to five species,
one Nanocochlea and four Pseudotricula, are found in single
stream habitats characterised by diverse sediment and slow to
moderate flows. One Pseudotricula species is found in a
swifter, larger stream with cobbles and gravel and another on
smooth limestone surfaces in fast flowing water and water
falls. A cladistic analysis (based on morphology) confirmed the
monophyly of Pseudotricula, but the monophyly of Nanocochlea is
less robust. --
Authors



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ZOOTAXAA radiation of hydrobiid snails in the caves and streams at Precipitous Bluff, southwest Tasmania, Australia (Mollusca: Caenogastropoda: Rissooidea: Hydrobiidae s.l .)W. F. PONDER, S. A. CLARK, S. EBERHARD & J. B. STUDDERTMagnolia Press Auckland, New Zealand 1074

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W. F. PONDER, S. A. CLARK, S. EBERHARD & J. B. STUDDERTA radiation of hydrobiid snails in the caves and streams at Precipitous Bluff, southwest Tasmania, Australia (Mollusca: Caenogastropoda: Rissooidea: Hydrobiidae s.l .) ( Zootaxa 1074) 66 pp.; 30 cm. 1 Nov. 2005 ISBN 1-877407-32-1 (paperback) ISBN 1-877407-33-X (Online edition) FIRST PUBLISHED IN 2005 BY Magnolia Press P.O. Box 41383 Auckland 1030 New Zealand e-mail: zootaxa@mapress.com http://www.mapress.com/zootaxa/ 2005 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326(Print edition) ISSN 1175-5334 (Online edition)

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1074Accepted by D. Geiger: 15 Aug. 2005; published: 1 Nov. 2005 3ZOOTAXA ISSN 1175-5326 (print edition) ISSN 1175-5334 (online edition) Copyright 2005 Magnolia Press Zootaxa 1074: 166 (2005) www.mapress.com/zootaxa/A radiation of hydrobiid snails in the caves and streams at Precipitous Bluff, southwest Tasmania, Australia (Mollusca: Caenogastropoda: Rissooidea: Hydrobiidae s.l .)W. F. PONDER1, S. A. CLARK2, S. EBERHARD3 & J. B. STUDDERT11 Australian Museum, 6 College Street, Sydney, NSW, 2010, Australia. Email: winstonp@austmus.gov.au2 Previously Australian Museum, now Department of Biological Sciences, Biodiversity and Systematics. University of Alabama, Box 870345, Tuscaloosa, AL 35487, United States of America3 Department of Conservation and Land Management, Science Division, PO Box 51, Wanneroo, WA 6065, AustraliaTable of contentsAbstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 The Precipitous Bluff caves and karst . . . . . . . . . . . . . . . . . . . . . . . 4 Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Genus Pseudotricula Ponder, 1992 . . . . . . . . . . . . . . . . . . . . . . . . 9 Pseudotricula eberhardi Ponder, 1992 . . . . . . . . . . . . . . . . . . . . . 11 Pseudotricula expandolabra n. sp. . . . . . . . . . . . . . . . . . . . . . . 15 Pseudotricula auriforma n. sp. . . . . . . . . . . . . . . . . . . . . . . . . 24 Pseudotricula elongata n. sp. . . . . . . . . . . . . . . . . . . . . . . . . 26 Pseudotricula arthurclarkei n. sp. . . . . . . . . . . . . . . . . . . . . . . 28 Pseudotricula conica n.sp. . . . . . . . . . . . . . . . . . . . . . . . . . 30 Pseudotricula progenitor n.sp. . . . . . . . . . . . . . . . . . . . . . . . . 34 Genus Nanocochlea Ponder and Clark, 1993 . . . . . . . . . . . . . . . . . . . . 40 Nanocochlea exigua n. sp.. . . . . . . . . . . . . . . . . . . . . . . . . . 42 Nanocochlea stylesae n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . 45 Nanocochlea damperensis n. sp. . . . . . . . . . . . . . . . . . . . . . . . 47 Discriminant function analysis . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Cladistic analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 Appendix: Station data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

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PONDER ET AL.4 2005Magnolia Press 1074ZOOTAXA AbstractA radiation of hydrobiid snails is described from caves at Precipitous Bluff, southern Tasmania. The radiation (10 species) comprises two closely related genera; Pseudotricula endemic to the Precipitous Bluff caves, and Nanocochlea found in these caves and surface streams and seepages, but also known from elsewhere in southern Tasmania. Two Precipitous Bluff species of Nanocochlea inhabit streams outside the caves, while one is found within the caves; all three are new. Seven species of Pseudotricula are described, six being new. Up to five species, one Nanocochlea and four Pseudotricula are found in single stream habitats characterised by diverse sediment and slow to moderate flows. One Pseudotricula species is found in a swifter, larger stream with cobbles and gravel and another on smooth limestone surfaces in fast flowing water and water falls. A cladistic analysis (based on morphology) confirmed the monophyly of Pseudotricula but the monophyly of Nanocochlea is less robust. Key words : karst, Nanocochlea Pseudotricula, cladistics, anatomy, stygobiteIntroduction Endemism in cave faunas is well known with some well documented examples of speciation in northern hemisphere karst areas (overviews in Culver & Holsinger 1992; Wilkens et al 2000). Radiations of hydrobiid and hydrobiid-like snails in karst and other groundwater environments have been described from other parts of the world e.g., New Zealand (Climo 1974), North America (e.g., Hershler & Longley 1986) and Europe (e.g., Giusti & Pezzoli 1977, 1981; Radoman 1983). However, only one obligate groundwaterdwelling species (stygobite) has been previously described from Australia, Pseudotricula eberhardi Ponder, 1992 from Precipitous Bluff, southwest Tasmania. One of us (SE) recollected from several caves within this karst area in 1991 and 1994 and found a number of additional species that are described herein. Samples were also collected by Arthur Clarke in 1988. Recent molecular studies (Wilke et al ., 2001) have shown that Hydrobiidae, as widely used, may not be monophyletic. In this paper the family name Hydrobiidae is used in the broad sense because the systematics of the family-group taxa involved require further studies before they are resolved. The Precipitous Bluff caves and karst Precipitous Bluff is located near the south coast of Tasmania in a remote and inaccessible part of the Tasmanian Wilderness World Heritage Area (Fig. 1). The area remains essentially undisturbed by human activities except for occasional visits by bushwalkers and cavers. The area lies within the perhumid precipitation effectiveness province of Gentilli (1972). The nearest meteorological station is at Hastings located 25 km

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2005 Magnolia Press 5HYDROBIIDAE S.L. 1074ZOOTAXA eastnortheast, where annual rainfall is 1416 mm. The cool, moist climate supports a dense vegetation cover of wet sclerophyll and cool temperate rainforest (Reid et al 1999).FIGURE 1. Schematic plan of mapped caves (black outline) at Precipitous Bluff showing hydrologic relationships (arrows indicate direction of stream flow paths), sample sites (numbered), cave entrances (black dots), and locations mentioned in text. Adapted from Hume (1989). Inserts: A Tasmania with location of Precipitous Bluff (PB in boxed area), Ida Bay (IB), Hastings (H), Vanishing Falls (VF) and Cracroft (C); B Precipitous Bluff and New River Lagoon showing surface sample sites (numbered) and location of the caves (boxed area) in relation to dolines (open circles) with sinking streams that feed Damper Cave and Quetzalcoatl Conduit, which becomes Damper Creek upon resurging at the surface. Also shown (in B) is Precipitous Bluff. The numbers shown in B are explained below. The arrows in insert B and the main figure show the direction of water flow. In the main map, the PB prefixed numbers refer to cave entrance numbers used by the Australian Speleological Federation. Sample sites shown in both the main map and in insert B are numbered as follows: 1 Bauhaus Cave (PB6) base of aven (stn PB6-1) and main streamway at downclimb (PB6-1A); 2 Bauhaus Cave (Persephone Pot PB17) lower streamway at first contact with stream stn 7; 3 Bauhaus Persephone middle streamway stn 8; 4 Bauhaus Persephone middle streamway stn 9; 5 Bauhaus Persephone upper streamway stn 10; 6 Bauhaus Screaming Stals streamway stn 11; 7 Bauhaus Screaming Stals streamway stn 12; and tributary seep near stn 12; 8 Bauhaus Screaming Stals streamway stn 13; 9 Cueva Blanca (PB4), Black Curtains streamway (=Inundation Passage); 10 Damper Cave (PB1) main streamway ca. 50 m inside entrance; 11 Damper Cave main streamway 100 m into cave; 12 Damper Cave, Honey and Cream; 13 Damper Cave seep near The Keg; 14 Damper Cave, Cane Toad Abuse; 15 Quetzalcoatl Conduit (PB3), 30 m into cave, stn PB31C; 16 Quetzalcoatl Conduit, beyond Pendulum Palace, stn PB3-2; 17 Damper Creek, 10 m downstream of Damper Cave entrance; 18 Damper Creek, first crossing; 19 Damper Creek at New River Lagoon; 20 Malani Creek; 21 Limestone Creek near New River Lagoon.

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PONDER ET AL.6 2005Magnolia Press 1074ZOOTAXA An extensive deposit of limestone occurs along the eastern flank of the New River valley and the western slope of Precipitous Bluff (Dixon & Sharples 1986). The Precipitous Bluff caves and karst are developed within Gordon Group Limestone of Ordovician age (Hughes 1957), the principal cave-bearing rock type in Tasmania. The limestone outcrops over about 10 km2 between 0300 m above sea level (asl) on the western and southwestern flanks of Precipitous Bluff (1120 m asl), which rises steeply from the shores of New River Lagoon, the estuary of the New River (Kiernan 1995). The caves are developed on the lower flanks of Precipitous Bluff and receive allogenic recharge from streams descending from the upper slopes of the mountain which sink underground on contact with the limestone at about 300 m asl. Autogenic recharge is also received from rainfall directly on the limestone outcrop, which percolates downward via vertical shafts and fissures in the limestone. The descending subsurface flow paths coalesce at lower levels to form an integrated dendritic cave drainage network which discharges from resurgence caves situated at base level on the slope-plain juncture. There are two resurgence caves associated with two drainage subsystems Quetzalcoatl Conduit and Damper Cave (Fig. 1). The Quetzalcoatl Conduit subsystem is relatively simple and essentially consists of a single, linear base-level conduit, probably mostly fed by a sinking stream located to the northeast of Elusive Bluff. As a consequence of its relatively large surface catchment area draining from the higher slopes of Precipitous Bluff, combined with rapid conduit through-flow, this subsystem experiences a variable and flashy flow regime, which periodically floods the cave completely as well as mobilising and redistributing quantities of fine sediment. In contrast to Quetzalcoatl Conduit the Damper Cave subsystem experiences flow regimes that are less variable and extreme, owing to a proportionately greater recharge contribution from diffuse sources. Inflows are smaller in size but numerous and these follow more intricate subterranean flow paths to base level. The Damper main stream, Cueva Blanca, and Bauhaus main stream are also quite responsive to rainfall events in the catchment, when surface runoff from the slopes of Precipitous Bluff rapidly sinks into the caves. However, the smaller, predominantly seepage fed streams (Persephone streamway, Hydrobiid Highway, Cane Toad Abuse) would have less variability in flow regime. The Damper Cave subsystem encompasses considerable structural complexity with numerous tributary networks developed at different vertical levels, and between different caves. Nonetheless, they are hydrologically integrated via conduit connections. From upstream to downstream the hydrologic flow path is Persephone Cave to Bauhaus to Cueva Blanca to Damper Cave (Fig. 1). The Quetzalcoatl Conduit and Damper Cave subterranean drainage subsystems lie adjacent to each other, and their resurging streams merge shortly downstream on the surface as Damper Creek before discharging into New River Lagoon. Hence, due to their close geographic proximity and downstream hydrologic connectivity, any observed faunal and hydrochemical similarities between the two subsystems may not be unexpected.

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2005 Magnolia Press 7HYDROBIIDAE S.L. 1074ZOOTAXA Certainly this appears to be the case in respect of a few measured hydrochemical parameters although very limited sampling has been undertaken. Water quality parameters in Damper Cave and Quetzalcoatl Conduit were measured on one occasion (Table 1) and both showed very similar water chemistry suggesting that any faunal differences between these two caves is likely to be due to other factors:TABLE 1. Chemistry of water from Damper Cave and Quetzalcoatl Conduit sampled on 22nd November 1990 by R. Eberhard (data cited in Kiernan 1995). The cave fauna at Precipitous Bluff was surveyed as part of a state-wide biodiversity survey of Tasmanian karsts undertaken by Eberhard et al (1991b). This study discovered that the Precipitous Bluff karst supported the richest assemblage of locally endemic and obligate cave-dwelling species (> 12 spp.) in Tasmania (Eberhard 1992a). Most of this assemblage comprised terrestrial cave obligate species (troglobites) including described species of Coleoptera (Moore 1978), Opiliones (Hunt 1990; Hunt & Hickman 1993) and Araneae (Platnick & Forster 1989), in addition to other undescribed species of Opiliones, Araneae, Pseudoscorpionida, Oniscidea, and Collembola (Eberhard et al 1991b). An interesting aquatic cave fauna was suggested by the discovery of Pseudotricula eberhardi Ponder, 1992. Besides this snail, the sampled aquatic macrofauna included mayflies (Ephemeroptera), and crustaceans (Amphipoda, Syncarida) (Eberhard et al 1991b; Eberhard 1992a). Of these other taxa at least two an undescribed species of paramelitid amphipod ( Antipodeus sp.) and an undescribed species of psammaspid syncarid ( Eucrenonaspides sp.) possessed clear stygomorphies (loss of eyes, pigment and elongation of appendages) indicating their status as obligate subterranean forms or stygobites (Eberhard et al 1991b). Additional sampling may well increase the number of stygobites, particularly as the aquatic microand meiofauna have not been sampled.Damper CaveQuetzalcoatl Conduit pH7.77.6 HCO3 mg/l 99.099.0 Cl-mg/l 23.624.7 SO4 2-mg/l 3.93.4 Ca2+mg/l 26.026.0 Mg2+mg/l 3.63.2 K+mg/l 0.620.64 Na+mg/l 11.912.0

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PONDER ET AL.8 2005Magnolia Press 1074ZOOTAXA Abbreviations Repositories AMS, Australian Museum, Sydney, New South Wales, Australia. QVM, Queen Victoria Museum, Launceston, Tasmania, Australia. TMAG, Tasmanian Museum and Art Gallery, Hobart, Tasmania, Australia Other abbreviations AL, aperture length. AW, aperture width. CV, convexity ratio. SEM, scanning electron microscope. SL, shell length. SW, shell width. TW, number of teleoconch whorls. Materials and Methods Collection of specimens took place between 18th December 1988 and 30th March 1994 during surveys of the limestone caves and adjacent surface streams in the Precipitous Bluff karst area in southern Tasmania (see Appendix for details). Specimens were collected using a combination of hand sieves to sweep the surface, washing of substrate, and picking them from the substrate by hand. A binocular microscope was used to sort material into relevant groups, whereby they were fixed in 10% formalin, buffered with sodium bicarbonate. Specimens were then preserved in 5% saltwater formalin. Adult specimens were randomly picked from samples and were measured using a digitalised graphics tablet linked to a computer as described by Ponder et al (1989). The convexity ratio (CV) was calculated using the method described by Ponder et al (1989). A subset of the measured specimens, usually three males and females, were dissected using a binocular dissecting microscope with an attached drawing apparatus. The radula, operculum and anatomy, including pigmentation, mantle cavity, digestive system, male and female reproductive systems (see appendix for further details), were examined for all but P elongata n. sp. These characters, together with the shell characters, were scored and entered into a DELTA (Dallwitz et al 1993) file. DELTA was used to generate descriptions. Shells and opercula used for SEM work were cleaned in freshwater using a small paint brush, fine jewellers, fine tipped tungsten wire, and in some cases using a sonicator. Cleaned specimens were then mounted on aluminium SEM stubs using either carbon tack, or carbon glue. Radulae (typically three per species) were extracted by dissolving the animal in a strong sodium hydroxide solution and incubating at approximately 60C for 6

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2005 Magnolia Press 9HYDROBIIDAE S.L. 1074ZOOTAXA hours. Each radula was then washed four to five times in distilled water, and mounted on a glass cover slip using fine forceps to break and spread them. The cover slip was mounted on an aluminium SEM stub with double-sided carbon tape and carbon glue. All SEM stubs were gold-coated prior to SEM examination. SYSTAT ver. 10 (SYSTAT Software Inc., Richmond, California, USA) was used for statistical analyses. PAUP* (ver. 4.0b10) (Swofford 1998) was used to perform maximum parsimony (cladistic) analyses. Genus Pseudotricula Ponder, 1992 Type species: Pseudotricula eberhardi Ponder, 1992 (original designation). Description Shell. Moderate to large in size (adults 1.7 to 4.4 mm SL), conical. Protoconch of about 1.5 whorls, typically sculptured with small, close pustules with some coalesced into weak, irregular radial ridges; separation of protoconch from teleoconch distinct, with varix-like border. Teleoconch with spire much longer to slightly shorter than length of aperture. Aperture ovate, slightly angled and typically channelled posteriorly, inner lip attached to parietal wall or partially to completely detached, although never markedly so; usually lower part raised well above base. Outer lip usually prosocline, with terminal growth, typically markedly expanded and flared, simple in some species; weak external varix present in one species ( P. elongata n. sp.). Periphery rounded, base simple, imperforate in both adults and juveniles. Usually semi-opaque to opaque, white, sometimes yellowish or brownish; with a thin pale yellowish periostracum. Operculum. Ovate, paucispiral, flat, columellar edge slightly convex, outer edge strongly convex; outer surface simple, paucispiral, nucleus markedly eccentric; inner surface with weak ridges (probably representing reduced pegs) or (more often) white smear only. Radula. Central teeth large, cutting edge broad, indented mid dorsally, with 45 cusps on either side of narrow sharp median cusp; narrow, thin lateral projections emerge at about 45 from mid laterally; base with short, rounded basal tongue not extending beyond lateral projections; two small to moderate basal cusps emerge from ventral face of tooth on either side of basal projection, innermost largest. Lateral teeth with short cutting edge, about 36 small cusps on inner side of narrow median cusp and 47 small cusps on outer side; neck prominent, near vertical; lateral flange about twice as long to slightly less than twice as long as cutting edge; prominent U-shaped projection on base below cutting edge; inner edge short, inner side of base excavated. Inner marginal teeth with 1533 tiny cusps on rather wide cutting edge (ratio of cutting edge to shaft on inner marginal teeth about ; sides approximately parallel, outer edge thickened. Outer marginal teeth narrow, thickened on inner edge, distal end with 1432 minute cusps; ratio of cutting edge to shaft about

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PONDER ET AL.10 2005Magnolia Press 1074ZOOTAXA Head-foot. Simple, unpigmented, with long cephalic tentacles, unpigmented eyes present in weak bulges at outer bases of tentacles; snout of moderate length, tapering, weakly bilobed distally. Foot short, rounded posteriorly. Non-genital anatomy. Pallial cavity elongate, osphradium large and oval, towards posterior end of ctenidium; ctenidium with small ctenidial filaments and long efferent vein (ctenidium occupies only anterior of pallial cavity). Hypobranchial gland variably developed. Kidney and pericardium usually about in pallial roof; renal gland orientated longitudinally. Stomach with anterior and posterior chambers about equal or posterior smaller; style sac moderately long; no posterior caecum. Rectum with long S-shaped coil; overlying but not indenting pallial oviduct. Male reproductive system. Prostate gland about within pallial cavity; compressed to oval in section, with very thin ventral wall; pallial vas deferens opens at about length of pallial portion. Penis located on right side of head well behind base of right tentacle; lacking glands but has an inconspicuous to moderately developed swelling in the distal portion just behind a papilla-like distal end; distal portion long and tapering; medial part simple and parallel sided; basal part wide to moderate. Female reproductive system. Ovary simple sac; coiled oviduct smooth, firm, not embedded in connective tissue, initially inverted U-shaped, usually straight distal to seminal receptacle (one species with bend), usually reaching posterior end of bursa (sometimes a little more or less) then sharply bent to run anteriorly; Seminal receptacle ovoid to pyriform, with short duct, opening to oviduct, located opposite middle to ventral part of left side of bursa copulatrix. Bursa copulatrix rather large, ovoid to pyriform, posterior to albumen gland except for small overlap on right side, either just behind posterior pallial wall or extending in front of it; bursal duct arises from anterior or ventroanterior wall, simple to undulating, joins oviduct dorsally at posterior pallial wall or in front of it as far anterior as junction of albumen and capsule glands. Common duct straight. Albumen gland or more within pallial roof, shorter to longer than capsule gland. Capsule gland compressed-oval in section, divided into three glandular zones, short anterior and posterior translucent white zones and long yellowish to orange middle zone; anterior end blunt to steeply tapering; genital opening short, terminal to slightly anterior of capsule gland. Ventral channel extended into weakly to moderately developed vestibule anteriorly. No brood pouch. Distribution and habitat Species of Pseudotricula are restricted to stream and seep habitats in the Precipitous Bluff caves. Remarks The relationships of this genus with Nanocochlea Ponder and Clark, 1993 are now well established, with molecular data (Perez et al 2005) showing they are sister taxa.

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2005 Magnolia Press 11HYDROBIIDAE S.L. 1074ZOOTAXA Anatomically the two genera are also very similar. They share the same type of ctenidium which only occupies the anterior half of the pallial cavity and with the filaments reduced in size and narrower than the normal broadly-triangular type. Other common features include the long, S-shaped rectal coil, unpigmented eyes, stomach morphology and genital anatomy. It could be argued that the fauna from Precipitous Bluff described herein shows a transition between the type species of Nanocochlea and Pseudotricula both of which lie at the extremes of what perhaps could be regarded as a single genus. The main differences between Nanocochlea and Pseudotricula are shell characters. The shell in typical Pseudotricula is broadly conical in shape, rather than elongate conic to pupiform, with a relatively large aperture that typically bears an expanded, prosocline outer lip. Other differences are detailed below under Nanocochlea The shell differences between the type species of Nanocochlea and Pseudotricula are considerable and their close relationship was not fully realised at the time of their description. Anatomically the two genera are very similar and this similarity is reflected in their sister-group relationship (Perez et al 2005). The paper describing Pseudotricula is listed as in press in Ponder et al (1993), but it was actually published shortly before in 1992. The species of Pseudotricula are rather uniform in anatomy, radular and opercular morphology, the latter differing only in the presence or absence of a peg on the inner surface. However, the species that we recognise here are readily discriminated on shell morphology (see below). The penial swelling was not noted in the original description of P. eberhardi because it is very weakly developed in that species (confirmed by re-examination). It is, however, present in the other species of Pseudotricula and an identical structure is also seen in several species of Nanocochlea Pseudotricula eberhardi Ponder, 1992 Figures 2A; 3A, B; 4A. SynonymyPseudotricula eberhardi Ponder, 1992: 26, plate 1AH, fig. 1A,B, fig. 2A,B, fig. 3AC; Kierman & Eberhard, 1990, pl. 3.1.Type material Holotype: AMS C.165052, Black Curtain Streamway, Cueva Blanca, PB4, 31 MAR 1986. Paratypes: AMS C.165053, same data (1 dry, 3 wet), TMAG E20151; AMS C.165054, same locality, 18 Dec. 1988 (9 dry, 20+ wet); TMAG E20152.

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PONDER ET AL.12 2005Magnolia Press 1074ZOOTAXA Other material examined Cueva Blanca : AMS C.165390, PB4 (1 dry topotype); AMS C.203693, Black Curtain Streamway, PB4-1C (12 dry, 14 wet).FIGURE 2. Shells of Pseudotricula species. A Pseudotricula eberhardi holotype, C.165052, Black Curtain Streamway, Cueva Blanca; BE Pseudotricula expandolabra B holotype, C.439673, Persephone Streamway, Bauhaus; C paratype, C. 439674, Persephone Streamway, Bauhaus; D C.201811, Persephone Pot, Bauhaus; E C201495, Persephone Streamway, Bauhaus; F Pseudotricula auriforma holotype, C.439672, main streamway, Damper Cave. Scales: A F 500 m m.Description Shell (Fig. 1A). Length up to 2.8 mm; broadly conical (SW/SL 0.710.80, mean 0.74, n = 10); spire low to moderate, straight to slightly convex in outline; last whorl evenly rounded; suture narrowly channelled; otherwise whorls evenly convex. Protoconch microsculpture small pits and pustules, with some weak spirally arranged wrinkles. Teleoconch up to 2.6 whorls in adult; aperture oval to pear-shaped; large, longer than spire (AL/SL 0.570.68, mean 0.63, n = 10); outer lip prosocline, straight to slightly sinuous, with strong reflection; external varix absent; notch present in posterior corner of aperture;

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2005 Magnolia Press 13HYDROBIIDAE S.L. 1074ZOOTAXA inner lip moderately thickened to thick and broad, adhering to, or narrowly separated from, parietal wall. Dimensions. See Table 2.TABLE 2. Shell dimensions and teleoconch whorl counts of Pseudotricula eberhardi Ponder. Operculum Yellowish; inner surface with white smear. Pallial cavity Ctenidium narrow; 913 very small filaments; osphradium between posterior end and middle of ctenidium; hypobranchial gland moderately developed; renal organ extends forward ca. into pallial cavity; pericardium in pallial roof. Radula (Fig. 3A, B). Central teeth (Fig. 3B): dorsal edge with shallow indentation; 5 lateral cusps, median cusp of medium width, sharply pointed, less than twice as long as adjacent cusps. Lateral teeth : dorsal edge straight to lightly convex; with 3 cusps on inner and 4 on outer side; median cusp of medium width, sharply pointed, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection bluntly pointed. Marginal teeth : Inner with 1517 cusps; outer with 1420 cusps. Stomach Stomach with posterior chamber a little smaller than anterior chamber. Male genital system. Testis of about 1 whorl; prostate gland oval to elongate pyriform; oval in section; pallial vas deferens straight to slightly undulating. Penis (Fig. 3A) with very weak swelling in mid-distal portion (hardly apparent); distal end tapering, similar to papilla in other taxa, medial section tapering; of medium length; penial duct in medial section of penis almost straight to undulating; base of penis moderately wide, with moderate folds; penial duct straight to undulating. Female genital system Ovary of 0.51.0 whorls; oviduct extends to posterior edge of bursa copulatrix; joins bursal duct at posterior pallial wall. Bursa copulatrix of medium size, not extending to posterior pallial wall; globular; with bursal duct arising from middle of anterior edge of bursa to antero-ventral edge; straight, wider at bursal end. Seminal receptacle at middle of inner wall of bursa copulatrix; pyriform. About to more than of albumen gland in front of posterior pallial wall; capsule gland about same to about length of albumen gland; circular to oval in section; anterior end blunt; ventral channel simple, approximately parallel-sided throughout; vestibular area indistinct; genital opening subterminal.SLSWALAWBWCVTW Holotype2.832.141.851.622.472.50 C.203693(9) paratypes Minimum2.331.651.341.331.940.112.30 Maximum2.661.961.691.652.350.162.62 Mean2.501.861.571.462.140.132.52 Standard Dev.0.120.120.110.110.130.020.10

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PONDER ET AL.14 2005Magnolia Press 1074ZOOTAXA FIGURE 3 : Radulae of Pseudotricula species. A, B, Pseudotricula eberhardi C.165053, Black Curtain Streamway, Cueva Blanca; CH, Pseudotricula expandolabra, C, D, C.201495, Persephone Streamway, Bauhaus, E, G, H, C.201811, Persephone Pot, Bauhaus, F, C.201822, main streamway, Bauhaus; I, Pseudotricula arthurclarkei C.203671, paratype, Quetzalcoatl Conduit; JL, Pseudotricula conica C.203676, paratype, Cane Toad Abuse Streamway, Damper Cave. A, C, G, K, central, lateral inner and outer marginal teeth; B, E, L, central teeth; D, outer marginal teeth; F, J, lateral and inner marginal teeth; H, lateral tooth; I, central and lateral teeth. Scales: A, B, E, HJ, L 10 m m; C, F, G, K, 20 m m; D, 5 m m. Distribution and habitat Found only on smooth rock surfaces in fast-flowing falls and streams in Cueva Blanca.

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2005 Magnolia Press 15HYDROBIIDAE S.L. 1074ZOOTAXA Remarks Pseudotricula eberhardi has only been found in very strongly flowing water in Cueva Blanca. A record of this species from Persephone Pot (Ponder 1992) are specimens of P. expandolabra n. sp. This species is distinguished from other congeners by its short, slightly convex spire and evenly convex whorls. Pseudotricula expandolabra n. sp. Figures 2B, E; 3CH; 4B; 5AF; 6AD; 7A,B; 8A; 9C; 10AC. Type material Holotype: AMS C.439673, Screaming Stals Streamway, Bauhaus, stn 12, 23 Dec, 1991. Paratypes: AMS C.201493, same locality stn 12 (6 dry, 12 wet); QVM, 9:20536 (5 wet). Figured (shell) paratypes: AMS C.439674, same locality stn 12.FIGURE 4. Dorsal views of penes of Pseudotricula and Nanocochlea species. A,. Pseudotricula eberhardi ; B, Pseudotricula expandolabra ; C, Nanocochlea stylesae ; D, Nanocochlea damperensis ; E, Pseudotricula conica Scales A, B, E, 0.2 mm; C, D, 0.1 mm. Abbreviations: ds, distal swelling; pd, penial duct.

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PONDER ET AL.16 2005Magnolia Press 1074ZOOTAXA FIGURE 5. Shells of Pseudotricula species. AF, Pseudotricula expandolabra AC, C.201489, Screaming Stals streamway, Bauhaus, DF, Persephone Pot, Bauhaus, D, F, C.201811, E, C.201485, Persephone Pot, Bauhaus; GI, Pseudotricula auriforma G, H, streamway near entrance, Damper Cave, I, main streamway, Damper Cave. Scales: 1 mm.

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2005 Magnolia Press 17HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 6. Protoconchs and teleoconch microsculpture of Pseudotricula and Nanocochlea species. AD, Pseudotricula expandolabra, A, B, D C.201493, paratype, main streamway, Bauhaus, C, C.201495, Persephone Streamway, Bauhaus; EG, Pseudotricula conica E, F, C.201464, Cane Toad Abuse Streamway, Damper Cave, G, Persephone Pot, Bauhaus; Honey and Cream Streamway, Damper Cave; H, Nanocochlea exigua C.201475; I, J, Nanocochlea stylesae C.201476, paratypes, Malani Creek; KM, Nanocochlea damperensis K, L, C.203685, paratypes, first crossing, Damper Creek, M, C.201282, Near New River Lagoon, E side, Limestone Ck, where tagged track crosses; N, O, Pseudotricula progenitor C.201821, main streamway, Bauhaus. A, B, E, F, H, KN, protoconch; C, D, G, I, J, O, microsculpture. Scales: A, CE, J, K, M, O, 200 m m; B, FI, L, N, 100 m m.

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PONDER ET AL.18 2005Magnolia Press 1074ZOOTAXA Other material examined Bauhaus : AMS C.201822, main streamway, stn PB6-1A (16 dry, 18 wet); AMS C.201486, Screaming Stals streamway, stn 11 (8 dry 14 wet); AMS C.201489, same locality, stn 13 (15 dry, 20+ wet); C.201485, Persephone, stn 7 (4 dry, 11 wet); C.201491, Persephone streamway, stn 8 (8 dry, 10 wet); C.201492, same locality, stn 9 (9 dry); C.201495, same locality, stn 10 (8 dry, 6 wet); C.439676, same locality, stn 10, figured specimen; C.201811, Persephone Pot, stn PB17-8R (1 dry, 4 wet); same locality, middle and upper streamway, stn PB17-2a.2 (20+ dry, 20+ wet); C.439675, same locality, stn PB17-2a.2, figured specimen; C.201815, same locality, lower streamway, stn PB17-2a.1, (20+ dry, 20+ wet); C.165055. Damper Cave : AMS C.201494, seep near The Keg, stn 3 (1 dry); AMS C.201490, Honey and Cream streamway, stn 4 (10 dry, 10 wet); AMS C.203678, Cane Toad Abuse streamway, stn PB1-2A (1 dry, 6 wet). Etymology Expando Latin, spread out, labrum Latin, lip; refers to the expanded outer lip of the aperture. Description Shell (Fig. 2B, E; 5AF; 6AD). Length up to 3.8 mm; with conical spire and large, expanded aperture (SW/SL 0.610.74, mean 0.69, n = 43); spire low to moderate, straight to slightly convex in outline, last whorl evenly rounded; usually with shallow subshoulder depression on last two whorls. Protoconch microsculpture of distinct, small pits (Fig. 6B). Teleoconch of up to 3.6 whorls in adult; aperture oval to pear-shaped; large, about equal to spire or slightly longer (AL/SL 0.520.63, mean 0.57, n = 43); outer lip prosocline; straight to or weakly sinuous, with strong reflection; external varix absent; notch present in posterior corner of aperture; inner lip thick, broad, firmly adhering to, or narrowly separated from, parietal wall. Dimensions. See Table 3.TABLE 3. Shell dimensions and teleoconch whorl counts of Pseudotricula expandolabra n.sp. SLSWALAWBWCVTW Holotype3.492.382.091.872.940.083.21 Figured Paratype3.452.41.961.772.850.072.87 Figured specimen (D)3.312.21.791.682.670.092.75 Figured Specimen (E)3.141.991.721.582.490.13.00 C.201493 paratypes(15) Minimum3.272.271.881.672.590.072.90 Maximum3.822.712.272.043.060.163.50 Mean3.622.532.111.892.900.103.11 Standard Dev.0.140.130.110.100.110.020.16......continued on the next page

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2005 Magnolia Press 19HYDROBIIDAE S.L. 1074ZOOTAXA Operculum (Fig. 7A, B). Yellowish; inner surface with or without white smear. Pallial cavity (Fig. 8A). Ctenidium narrow; 1216 very small filaments; osphradium between posterior end and middle of ctenidium; hypobranchial gland variably developed; renal organ extends forward ca. into pallial cavity; pericardium more than in pallial roof. Radula (Fig. 3CH). Central teeth (Fig. 3E): dorsal edge with shallow indentation; 45 lateral cusps, median cusp of medium to narrow, sharply pointed, less than twice as long as adjacent cusps. Lateral teeth (Fig. 3 F, H): dorsal edge with shallow indentation; with 34 (usually 4) cusps on outer and 35 on inner side; median cusp of medium width, sharply to bluntly pointed, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection bluntly pointed. Marginal teeth : Inner (Fig. 3F) with 1621 cusps; outer (Fig. 3D) with 1726 cusps. Stomach (Fig. 9C). Stomach with posterior chamber and anterior chamber about equal in size. Male genital system. Testis of 1.01.25 whorls; prostate gland oval to kidney-shaped to elongate pyriform; compressed in section. Pallial vas deferens straight to slightly undulating. Penis (Fig. 4B) weak swelling in mid-distal portion; distal end long, papillalike; medial section tapering to parallel sided, of medium length; penial duct strongly undulating; base of penis moderately wide; with moderate folds; penial duct straight to undulating. Female genital system (Fig. 10AC). Ovary of 1.0 whorls; oviduct extends to posterior edge of bursa copulatrix; with one bend distal to seminal receptacle; joins bursal duct about half way between posterior pallial wall and capsule gland. Bursa copulatrix large, extending to posterior pallial wall or slightly in front; globular to elongately oval; with bursal duct arising from middle of anterior edge of bursa; straight or with undulations. Seminal receptacle at middle of inner wall of bursa copulatrix or near mid ventral edge; ovoid. More than of albumen gland in front of posterior pallial wall; capsule gland longer than albumen gland; compressed oval in section; anterior end blunt;TABLE 3 (continued) SLSWALAWBWCVTW C.201495(4) Minimum3.022.121.711.602.450.062.90 Maximum3.702.662.001.862.970.173.55 Mean3.262.271.811.702.630.103.19 Standard Dev.0.300.260.130.110.240.050.30 C.201811(20) Minimum3.052.011.621.492.420.062.75 Maximum3.612.481.971.862.900.133.30 Mean3.242.201.791.652.580.103.01 Standard Dev.0.150.120.090.110.120.020.15

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PONDER ET AL.20 2005Magnolia Press 1074ZOOTAXA ventral channel simple, approximately parallel-sided throughout; vestibular area indistinct to distinct; genital opening terminal to overlapping anterior end of capsule gland.FIGURE 7. Opercula of Pseudotricula species. A, B, Pseudotricula expandolabra, C.201493, paratypes, main streamway, Bauhaus; C, Pseudotricula auriforma C.201496, paratype, main streamway, Damper Cave; D, E, Pseudotricula arthurclarkei C.203671, paratypes, Quetzalcoatl Conduit; FI, Pseudotricula conica F, G, C.203680, main streamway, near entrance, Damper Cave; H, I, C.201459, Screaming Stals streamway, Bauhaus; A, D, F, H, outer side; B, C, E, H, I, inner side; Scales: AI, 200 m m.

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2005 Magnolia Press 21HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 8. Pallial cavities of male specimens of Pseudotricula and Nanocochlea species. A, Pseudotricula expandolabra C.201493, paratype, Persephone Streamway, Bauhaus; B, Pseudotricula arthurclarkei C.203671, paratype, Quetzalcoatl Conduit; C, Pseudotricula auriforma C.201496, paratype, main streamway, Damper Cave; D, E, Pseudotricula conica D, C.203680, main streamway near entrance, Damper Cave, E, Cane Toad Abuse Streamway, Damper Cave; F, Nanocochlea exigua C.203677, paratype, Cane Toad Abuse Streamway, Damper Cave; G, Nanocochlea damperensis C.203685, paratype, first crossing, Damper Creek; H, Pseudotricula progenitor C.201821, main streamway, Bauhaus. Scales: AD, FH, 500 m m, E, 1 mm. Abbreviations: a, anus; ct, ctenidium; ev, efferent vein; os, osphradium; pc, pericardium; pe, pallial edge; ppw, posterior pallial wall; r, rectum; rg, renal gland. The stippled area represents the hypobranchial gland.

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PONDER ET AL.22 2005Magnolia Press 1074ZOOTAXA FIGURE 9. Head, penis and stomach of Pseudotricula species. A, D, Pseudotricula conica C.203680, main streamway near entrance, Damper Cave; C, Pseudotricula expandolabra C.201493, paratype, Persephone Streamway, Bauhaus; B, E, Pseudotricula progenitor C.201821, main streamway, Bauhaus. A, B, head and penis; CE, stomach. Scales: AE, 500 m m. Abbreviations: dgo, opening from stomach to digestive gland; ey, eye; int, intestine; oes, oesophagus, p, penis; pg, penial duck; sn, snout; ss, style sack; st, stomach; tn, tentacle.

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2005 Magnolia Press 23HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 10. Female reproductive systems of Pseudotricula species. A C, Pseudotricula expandolabra A, B, C.201493, paratype, Persephone Streamway, Bauhaus, C, C.201815, Persephone Streamway, Bauhaus; D, Pseudotricula auriforma C.201496, paratype, main streamway, Damper Cave; E, Pseudotricula arthurclarkei C.203671, paratype, Quetzalcoatl Conduit; F, G, Pseudotricula conica F, C203680, main streamway near entrance, Damper Cave, G, Cane Toad Abuse Streamway, Damper Cave. Scale: 500 m m. Abbreviations: ag, albumen gland; bc, bursa copulatrix; bd, bursal duct; cg, capsule gland; co, coiled oviduct; po, genital opening; pw, pallial wall; sr, seminal receptacle; vc, ventral channel. Distribution and habitat Damper Cave (Honey and Cream and Cane Toad Abuse streamways and seep near The Keg) and Bauhaus (Persephone and Screaming Stals streamways and Persephone Pot). This species is found in narrow (less than 1m in width) streams with gentle to moderate (<1L/s to about 5L/s estimated flows) within the caves. The substrate is mixed cobbles, gravel and silt and the microhabitats pools and riffles. Two samples were also found in seepages associated with this type of stream.

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PONDER ET AL.24 2005Magnolia Press 1074ZOOTAXA Remarks Pseudotricula expandolabra is similar to P. eberhardi but its shell differs in having a straight spire outline, a taller spire (P<0.001) and the whorls are indented slightly below the shoulder. While it is rather constant in shape, it varies in size. Specimens from Persephone Pot are typically smaller (Fig. 5DF) than those from the main Bauhaus cave. There also appears to be some notable anatomical variation; a specimen from Persephone Pot (Fig. 10C) has the bursal duct showing substantial undulations and the bursa copulatrix being larger and longer but a second specimen from this location is like some specimens from Bauhaus with only a single bend in the bursal duct (others have a straight bursal duct) and a smaller bursa. Given the continuity of these two systems and the similarity in the shells of specimens from the two localities, they are treated as a single species pending more detailed investigation. Pseudotricula auriforma n. sp. Figures 2F; 5GI; 7C; 10D. Type material Holotype: AMS C.439672, Damper Cave, main streamway, stn 6, 22, DEC, 1991. Paratypes: AMS C.201496, Damper Cave, main streamway, stn 6, 22, DEC, 1991 (3 dry, 2 wet); AMS C.203681, Damper Cave, streamway near entrance, stn PB11A, 28, MAR, 1994, 4 dry, 1 wet); QVM, 9:20537 (1 dry). Etymology Auris Latin, ear; forma Latin, shape; refers to the ear-shaped aperture. Description Shell (Figs 2F; 5GI). Length up to 4.4 mm; conical (SW/SL 0.590.65, mean 0.63, n = 6); spire moderate to high, straight in outline; last whorl evenly rounded, or angular to subangular in middle of whorl; suture indented/impressed, grooved/channelled or with very narrow shoulder above or below; subshoulder depression on last two whorls. Protoconch microsculpture unknown. Teleoconch up to 3.8 whorls in adult; aperture oval to pear-shaped; large, about equal to spire (AL/SL 0.470.55, mean 0.52, n = 6); outer lip prosocline, thin to weakly thickened in adult, straight, with strong reflection; external varix absent; notch usually present in posterior corner of aperture; upper half of inner lip thin to moderately thickened and moderately wide, thicker in lower half, in partial contact or narrowly separated from parietal wall. Translucent, semi-opaque; white (semi-opaque to opaque), or yellow. Dimensions. See Table 4.

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2005 Magnolia Press 25HYDROBIIDAE S.L. 1074ZOOTAXA TABLE 4. Shell dimensions and teleoconch whorl counts of Pseudotricula auriforma n.sp. Operculum Yellowish or brown; inner surface with or without white smear. Pallial cavity (Fig. 8C). Ctenidium narrow; 1415 very small filaments; osphradium between posterior end and middle of ctenidium to near middle of ctenidium; hypobranchial gland thick; renal organ extends forward ca. into pallial cavity; pericardium more than in pallial roof. Radula (Fig. 7C). Central teeth : dorsal edge with shallow indentation; 45 lateral cusps, median cusp of medium width, sharply pointed, less than twice as long to about equal to adjacent cusps. Lateral teeth : dorsal edge with shallow indentation; with 34 cusps on outer and 3 on inner side; median cusp of medium width, sharply pointed, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection bluntly pointed. Marginal teeth : Inner with 1517 cusps; outer with 1519 cusps. Stomach Stomach with posterior chamber and anterior chamber about equal in size, or with posterior chamber a little smaller than anterior chamber. Male genital system. Testis of 1.52.0 whorls; prostate gland oval to kidney-shaped; compressed in section. Pallial vas deferens straight. Penis with weak swelling in mid-distal portion; distal end long, papilla-like; medial section parallel sided, of medium length; penial duct in medial section of penis straight to slightly undulating; base of penis moderately wide, or narrow; with weak to moderate folds; penial duct straight to undulating. Female genital system (Fig.10D). Ovary of 0.91.2 whorls; oviduct extends to posterior edge of bursa copulatrix or slightly anterior to that edge; with or without one bend distal to seminal receptacle; joins bursal duct in front of posterior pallial wall about half way between posterior pallial wall and capsule gland to about junction of albumen and capsule glands. Bursa copulatrix large, slightly in front of posterior pallial wall to slightly behind; pyriform to elongately oval; with bursal duct arising from middle of anterior edge of bursa; straight or with bend. Seminal receptacle at middle of inner wall of bursa copulatrix or near mid ventral edge; ovoid to pyriform. More than of albumen gland in front of posterior pallial wall; capsule gland longer than albumen gland; compressed oval in section; anterior end tapering to blunt; ventral channel simple, approximately parallel-sided throughout; vestibular area distinct; genital opening overlapping anterior end of capsule gland. SLSWALAWBWCVTW Holotype4.462.762.462.223.440.083.6 C.201496 paratypes (5) Minimum3.642.311.711.672.720.073.50 Maximum4.262.662.261.953.170.123.75 Mean4.002.512.051.823.000.093.63 Standard Dev.0.230.170.210.130.200.020.12

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PONDER ET AL.26 2005Magnolia Press 1074ZOOTAXA Distribution and habitat Known only from the main streamway in Damper Cave. This streamway is wider (about 2m) and has a faster flow (estimated at 1020L/s) than the streams in which P. expandolabra occurs. The substrate consists of cobbles and gravel. Remarks Pseudotricula auriforma is similar to P. expandolabra but differs mainly (P<0.001) in its larger, more slender shell. It lives in a streamway that is wider and faster flowing than any of the habitats in which P. expandolabra occurs and the substrate lacks fine sediments. Pseudotricula elongata n. sp. Figure 11AC. Type material Holotype: AMS C.166854, Bauhaus, Persephone Pot, stn PB178R, 3, JAN, 1990. Paratypes: AMS C.201809, same location, middle and upper streamway, stn PB172a.2, 27, March 94 (3 dry); AMS C.203713, same location, stn 9, 23, DEC, 1991 (1 dry, 1 wet). Etymology Elongatus Latin, prolonged; refers to the elongate shell. Description Shell (Fig. 11AC). Length up to 2.8 mm; elongate-conic broadly conical (SW/SL 0.450.56, mean 0.49, n = 3);spire tall, straight in outline; last whorl evenly rounded, or angular to subangular in middle of whorl; suture grooved/channelled or with very narrow shoulder above or below. Protoconch microsculpture unknown. Teleoconch up to 4.4 whorls in adult; aperture oval to pear-shaped; small, much shorter than spire (AL/SL 0.370.44, mean 0.40, n = 3); outer lip orthocline, straight, with narrow reflection and thickened; notch present in posterior corner of aperture; inner lip thin to moderately thickened and narrow to medium width, in partial contact or narrowly separated from parietal wall. Dimensions. See Table 5.TABLE 5. Shell dimensions and teleoconch whorl counts of Pseudotricula elongata n.sp. SLSWALAWBWCVTW Holotype 2.831.321.050.821.770.084.4 C.201809(1)1.380.770.610.480.950.092.50 C.203713(1)2.311.040.900.681.530.103.65

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2005 Magnolia Press 27HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 11. Shells of Pseudotricula species. A C Pseudotricula elongata Persephone Pot, A, holotype, C.166854; B,C, paratypes, B, C.203713, C, C.201809; DF, Pseudotricula arthurclarkei C.203671, paratypes, Quetzalcoatl Conduit. Scales: A C, 500 m m; DF, 1 mm. Distribution and habitat Known only from Persephone Pot, Bauhaus, where it lives in a narrow (about 0.5m wide), slow flowing (estimated about 1L/s) stream. The substrate is composed of cobbles, gravel and silt. Remarks Pseudotricula elongata has a very distinctive, elongate shell shape and is known only from a few specimens. It is only known from Persephone Pot, where it is found living with several other species (Table 15). It is readily distinguished from all of the other species found in the caves by its large, elongate shell and distinctly sinuate outer lip.

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PONDER ET AL.28 2005Magnolia Press 1074ZOOTAXA Pseudotricula arthurclarkei n. sp. Figures 3I; 7E; 8B; 10E; 11DF; 12A. Type material Holotype: AMS C.439656, Quetzalcoatl Conduit, stn PB31C, 29, MAR, 1994. Paratypes: AMS C.203671, Quetzalcoatl Conduit, stn PB31C, 29, MAR, 1994 (20+ dry, 20+ wet); AMS C.203669, Quetzalcoatl Conduit, stn PB32B, 29, MAR, 1994 (12 dry, 20+ wet); QVM, 9:20538 (5 wet). Etymology Named for Arthur Clarke in recognition of his considerable contributions to the discovery and collection of cave fauna in Tasmania. Description Shell (Figs 11DF; 12A). Length up to 3.1 mm; broadly conical (SW/SL 0.770.88, mean 0.81, n = 11); spire low to moderate, straight to slightly convex in outline; last whorl evenly rounded, or angular to subangular in middle of whorl; whorls distinctly convex; suture simple. Protoconch microsculpture of closely-spaced, distinct small pits. Teleoconch up to 2.5 whorls in adult; aperture oval to pear-shaped; large, little longer than spire (AL/SL 0.530.67, mean 0.61, n = 11); outer lip prosocline; weakly to moderately thickened in adult; outer lip straight; with or without slight reflection; external varix absent; posterior notch absent; inner lip moderately thickened and of narrow to medium width, firmly adhering to narrowly separated from parietal wall. Shell colour white or yellowish-brown to pale brown. Dimensions. See Table 6.TABLE 6. Shell dimensions and teleoconch whorl counts of Pseudotricula arthurclarkei n.sp. Operculum (Fig. 7E).Yellowish; inner surface with white smear and 36 mediumsized pegs, or with pegs only. Pallial cavity (Fig. 8B). Ctenidium narrow; 1416 very small filaments; osphradium near posterior end to between posterior end and middle of ctenidium; hypobranchial gland moderately to poorly developed; renal organ extends forward ca. into pallial cavity; pericardium more than in pallial roof. SLSWALAWBWCVTW Holotype2.992.021.591.392.440.122.5 C203671 paratypes (10) Minimum2.592.121.631.422.310.102.00 Maximum3.072.661.911.742.700.152.45 Mean2.882.381.791.652.540.122.22 Standard Dev.0.160.150.100.090.120.020.12

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2005 Magnolia Press 29HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 12. Shells of Pseudotricula species. A, Pseudotricula arthurclarkei holotype, C.439656, Quetzalcoatl Conduit; B D, Pseudotricula conica B, holotype, Cane Toad Abuse Streamway, Damper Cave; C, C.203680, main streamway near entrance, Damper Cave; D, C.201813, Persephone Pot, Bauhaus. Scales: AD, 500 m m.Radula (Fig. 3I). Central teeth : dorsal edge moderately indented; 56 lateral cusps, median cusp narrow, sharply pointed, less than twice as long as adjacent cusps. Lateral teeth : dorsal edge straight to lightly convex or with shallow indentation; with 56 cusps on outer and 36 on inner side; median cusp of narrow to medium width, sharply pointed, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection rounded to bluntly pointed. Marginal teeth : Inner with 2833 cusps; outer with 2327 cusps.

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PONDER ET AL.30 2005Magnolia Press 1074ZOOTAXA Stomach Stomach with posterior chamber much smaller than anterior chamber. Male genital system. Testis of 0.752.0 whorls; prostate gland oval to kidney-shaped, oval in section. Pallial vas deferens straight. Penis with weak swelling in mid-distal portion; distal end long, papilla-like; medial section parallel sided, of medium length; penial duct straight to undulating; base of penis moderately wide, with moderate folds; penial duct straight to undulating. Female genital system (Fig. 10E). Ovary of 1.0 whorls; oviduct does not extend to posterior edge of bursa copulatrix; straight distal to seminal receptacle; joins bursal duct in front of posterior pallial wall about half way between posterior pallial wall and capsule gland to about junction of albumen and capsule glands. Bursa copulatrix of medium size, extending to posterior pallial wall; globular to pyriform; with bursal duct arising from middle of anterior edge of bursa; straight. Seminal receptacle at middle of inner wall of bursa copulatrix or near mid ventral edge; pyriform. About to more than of albumen gland in front of posterior pallial wall; capsule gland about same length as albumen gland; compressed oval in section; anterior end tapering to blunt; ventral channel simple, approximately parallel-sided throughout; vestibular area indistinct to distinct; genital opening overlapping anterior end of capsule gland. Distribution and habitat Known only from Quetzalcoatl Conduit, where it lives in a stream 34 m wide which has a strong flow (estimated at 1530L/s) with sediments composed mostly of gravel and silt. Remarks Pseudotricula arthurclarkei and the next two species differ from typical members of Pseudotricula in having more cusps on both marginal teeth, a long lateral shaft on the lateral teeth and in lacking a strongly reflected outer lip. Pseudotricula arthurclarkei and P. conica n.sp. appear to be a sister taxa and also differ from other Pseudotricula species in having several weak to moderate curved ridges (reduced pegs) on the inside of the operculum, stronger and more complex protoconch microsculpture and the shell colour is often brown. Pseudotricula conica n.sp. Figures 3JL; 4E; 6EG; 7FI; 8D, E; 9A, D; 10F, G; 12BD; 13AF. Type material Holotype: AMS C.439649, Cane Toad Abuse Streamway Damper Cave, stn PB12A, 28 Mar 1994. Paratypes: AMS C.203676, Damper Cave, same locality, stn PB12A (10 dry, 20+ wet); QVM, 9:20539 (5 wet).

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2005 Magnolia Press 31HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 13. Shells of Pseudotricula species. A F, Pseudotricula conica A, C.203694, Black Curtain streamway, Cueva Blanca, B, Screaming Stals streamway, Bauhaus; C, C.201813, Persephone Pot, Bauhaus, D, C.203464, Cane Toad Abuse Streamway, Damper Cave, E, C.201488, main streamway, Damper Cave, F, C.203692, Damper Cave. Scales: 1 mm.Other material examined (all AMS) Bauhaus : C.201823, main streamway, stn PB6-1A, 14 dry); C.201824, same locality, stn PB6-1A (14 dry); C.201460, Screaming Stals streamway, stn 12 (2 dry, 17wet); C.201459, Screaming Stals streamway, stn 11 (12 dry, 20+ wet); C.201469, same locality, stn 13 (3 dry); C.439498, same locality, stn 12 (2 dry); C.201813, Persephone Pot, stn PB17-2a.2, middle and upper streamway (9 dry); C.201817, same locality, stn PB17-2.1, lower streamway (20+ dry, 20+ wet); C.439650, same locality, stn PB17-2a.1 (1dry) figured specimen; C.439496, Persephone Streamway, stn 8 (4 dry); C.439501, same locality, stn 9 (2 dry); C.439499, same locality, stn 9 (1 dry); C.439500, same locality, stn 10 (3 dry).

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PONDER ET AL.32 2005Magnolia Press 1074ZOOTAXA Damper Cave : C.203692, stn PB1-1B (1 dry, 3 wet); C.201488, main streamway, stn 6 (9 dry, 16 wet); C.203680, main streamway near entrance, stn PB1-1A (20+ dry, 20+ wet); C.439652, same locality, stn PB1-1A, (1dry), figured specimen; C.201464, Cane Toad Abuse streamway, stn 2 (11 dry, 20+ wet); C.201487, same locality, stn 2 (4 dry, 4 wet); C.201470, stn 2 (6 dry); Cueva Blanca : C.166967, no stn no. (2 dry, 6 wet); C.203694, Black Curtain Streamway, stn PB4-1C (5 dry). Etymology Conicus Latin adjective, cone-shaped; refers to the conical shape of the shell of this species. Description Shell (Figs 6EG; 12BD; 13AF). Length up to 3.2 mm (usually less than 3 mm, mean 2.3 mm); conical to broadly conical (SW/SL 0.580.75, mean 0.67, n = 60); spire low to moderate, straight to slightly convex in outline; last whorl evenly rounded, or angular to subangular in middle of whorl; whorls very slightly convex; suture simple. Protoconch microsculpture of small, distinct, closely spaced pits (Fig. 6E, F). Teleoconch up to 3.3 whorls in adult; aperture oval to pear-shaped; moderately large, shorter to longer than spire (AL/SL 0.400.60, mean 0.50, n = 60); outer lip prosocline, straight, with or without slight reflection; external varix absent; posterior notch absent; inner lip thin to moderately thickened and narrow to medium width, firmly adhering to narrowly separated from parietal wall. Shell colour white or yellowish-brown to pale brown. Dimensions. See Table 8.TABLE 8. Shell dimensions and teleoconch whorl counts of Pseudotricula conica n.sp. SLSWALAWBWCVTW Holotype2.261.431.051.061.760.112.75 Figured Specimen2.071.281.010.891.580.142.60 Figured specimen2.31.511.231.061.850.12.75 C.201459(20) Minimum1.831.210.920.871.420.092.50 Maximum2.521.631.201.121.940.183.00 Mean2.221.461.071.011.710.132.76 Standard Dev.0.160.110.090.070.120.020.15 C.201813(17) Minimum1.651.010.670.571.180.012.12 Maximum2.441.471.120.991.750.223.30 Mean2.021.290.940.821.520.112.75 Standard Dev.0.250.150.120.140.170.050.31 C.203680(20) Minimum2.381.661.271.111.980.082.50 Maximum3.232.071.571.392.450.163.10 Mean2.611.841.411.262.150.122.66 Standard Dev.0.200.120.090.070.130.020.16

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2005 Magnolia Press 33HYDROBIIDAE S.L. 1074ZOOTAXA Operculum (Fig. 7FI). Yellowish; inner surface with white smear. Pallial cavity (Fig. 8D, E). Ctenidium narrow; 1014 very small filaments; osphradium between posterior end and middle of ctenidium; hypobranchial gland thick; renal organ extends forward ca. into pallial cavity; pericardium more than in pallial roof. Radula (Fig. 3JL). Central teeth : dorsal edge with moderate to deep indentation; 5 lateral cusps, median cusp of narrow to medium width, bluntly to sharply pointed, less than twice as long as adjacent cusps. Lateral teeth : dorsal edge lightly convex to moderately indented; with 47 cusps on outer and 46 on inner side; median cusp of medium to narrow, sharp to blunt, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection bluntly pointed. Marginal teeth : Inner with 2132 cusps; outer with 2232 cusps. Stomach (Fig. 9D). Stomach with posterior chamber and anterior chamber about equal in size. Male genital system. Testis of 1.25 whorls; prostate gland oval to elongate pyriform; compressed in section. Pallial vas deferens straight. Penis (Fig. 4E; 9A) with weak swelling in mid-distal portion; distal end long, papilla-like; medial section parallel sided; of medium length; penial duct in medial section straight to undulating; base of penis moderately wide; with weak to moderate folds; penial duct straight to undulating. Female genital system (Fig. 10F, G). Ovary of 0.61.0 whorls; oviduct extends to posterior edge of bursa copulatrix or slightly anterior or posterior to edge; straight distal to seminal receptacle; joins bursal duct in front of posterior pallial wall about half way between posterior pallial wall and capsule gland to about junction of albumen and capsule glands. Bursa copulatrix large, extending to posterior pallial wall or into pallial roof; elongately oval to pyriform; with bursal duct arising from middle of anterior edge of bursa or from antero-ventral edge; straight or with undulations. Seminal receptacle at middle of inner wall of bursa copulatrix or near mid ventral edge; ovoid to pyriform. About to more than of albumen gland in front of posterior pallial wall; capsule gland about same length as albumen gland to about length of albumen gland; compressed oval in section; anterior end tapering to blunt; ventral channel simple, approximately parallel-sided throughout; vestibular area indistinct; genital opening overlapping anterior end of capsule gland. Distribution and habitat This is one of the commonest and widely distributed species in the caves. It appears to prefer low energy streams with mixed substrates, although a few specimens have been found in high energy habitats, notably in Cueva Blanca. Remarks Pseudotricula conica exhibits considerable variation in shell size and shape in what

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PONDER ET AL.34 2005Magnolia Press 1074ZOOTAXA we have interpreted as a single species. In particular, samples from Cueva Blanca and Bauhaus caves are fairly consistently elongate (e.g., Fig. 12D; 13A, C), while those from Damper Cave are usually broader (Fig. 12B, C; 13DF). At least some material from the latter cave shows a range of shell morphology tall to broad with intermediate shell shapes (as for example in Fig. 13B from Bauhaus). In addition, the anatomical and radular details of the broad and narrow forms are very similar. Because of this, and the existence of at least some apparent intermediates, we treat both forms as a single species pending more detailed studies. Specimens near the entrance to Damper cave (C.203680) are larger with rather dark-coloured shells than those from further inside the cave. Pseudotricula conica is most similar to P. arthurclarkei from Quetzalcoatl Conduit but differs in its smaller size (P<0.001) although one lot from close to the entrance of Damper Cave (C.203680) is nearly as large as the former species. P. conica also tends to have less convex whorls and has weak to obsolete opercular ridges. The two taxa are not sympatric. Pseudotricula progenitor n.sp Figures 6N, O; 8H; 9B,E; 14H, I; 15J, K; 16G, H; 17LO; 18C, D. Type material Holotype: AMS C.439396, Bauhaus, Persephone streamway, stn 8, 23 DEC 1991. Paratypes: AMS C.201463, same data, 30+ (11 dry, 20+ wet); QVM, 9:20540 (5 wet); AMS C.439397, same data (Fig. 14H). Other material examined (all AMS) Bauhaus : C.438369, stn PB6-1 (5 dry); C.201821, main streamway, stn PB6-1A (10 dry, 20+ wet); C.166856, Persephone Pot, stn PB17-8R (5 dry, 25 wet); C.201812, same locality, middle and upper streamway, stn PB17-2a.2 (20+ dry, 20+ wet); C.438370, C.201816, same data (20+ dry, 20+ wet); same locality, PB17-2A (20+ dry); C.201454, Persephone, stn 7 (4 dry, 20+ wet); C.201468, Persephone streamway, stn 9 (15 dry, 20+ wet); C.201484, same locality, stn 10 (6 dry, 20+ wet). Damper Cave : C.203682, Main streamway near entrance, stn PB1-1A (3 dry, 5 wet); C.201458, main streamway, stn 6 (14 dry, 20+ wet). Etymology Progenitor Latin, founder of a family, ancestor. Refers to the apparently basal position (and plesiomorphic nature) of this species in relation to the other taxa included in Pseudotricula Description Shell (Fig. 6N, O; 14H, I; 15J, K). Length up to 2.8 mm; elongate-conic (SW/SL 0.500.64, mean 0.56, n = 22); spire tall, straight to slightly convex in outline; subsutural

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2005 Magnolia Press 35HYDROBIIDAE S.L. 1074ZOOTAXA indentation on last two whorls; periphery of last whorl evenly rounded; suture simple. Protoconch (Fig. 6N) as for genus, microsculpture largely obscured in available material. Teleoconch up to 4.0 whorls in adult; umbilicus absent in juveniles; aperture oval to pearshaped; small, shorter than spire (AL/SL 0.340.49, mean 0.40, n = 22); outer lip orthocline to opisthocline, weakly thickened in adult, straight, usually with slight reflection; posterior notch absent; inner lip thin to moderately thickened and of narrow to medium width, in partial contact or narrowly separated from parietal wall. Dimensions See Table 9.TABLE 9. Shell dimensions and teleoconch whorl counts of Pseudotricula progenitor n.sp. Operculum (Fig. 16G,H). Inner surface with or without white smear. Eyes Unpigmented. Pallial cavity (Fig. 8H). Ctenidium narrow; 910 very small filaments; osphradium between posterior end and middle of ctenidium; hypobranchial gland thick to moderately developed; renal organ extends forward ca. into pallial cavity; pericardium more than in pallial roof. Radula (Fig. 17LO). Central teeth : dorsal edge with moderate to deep indentation; 56 lateral cusps, median cusp narrow, sharply pointed, about twice as long as adjacent cusps. Lateral teeth : dorsal edge with shallow indentation; with 56 cusps on outer and 45 on inner side; median cusp narrow, sharply pointed, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection bluntly pointed. Marginal teeth : Inner with 2529 cusps; outer with 2228 cusps. Stomach (Fig. 9E). Stomach with posterior chamber and anterior chamber about equal in size or posterior chamber a little smaller. Male genital system. Testis of 0.91.5 whorls; prostate gland oval to kidney-shaped; compressed in section. Pallial vas deferens straight. Penis (Fig. 9B) with weak swelling in mid-distal portion; distal end long, papilla-like; medial section tapering to parallel sided; of medium length; penial duct in medial section of penis strongly undulating; base of penis moderately wide; with moderate folds; penial duct strongly undulating. SLSWALAWBWCVTW Holotype2.641.350.950.861.770.133.6 Figured paratype 2.781.390.980.831.650.114.0 C.201468 (20) Minimum1.911.220.890.801.390.062.75 Maximum2.821.561.090.951.770.163.75 Mean2.471.400.980.871.640.113.38 Standard Dev.0.250.080.050.030.100.020.30

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PONDER ET AL.36 2005Magnolia Press 1074ZOOTAXA FIGURE 14. Shells of Nanocochlea species and Pseudotricula progenitor A C, Nanocochlea exigua A, holotype, C.439393, Cane Toad Abuse Streamway, Damper Cave; B, C, C.201465, main streamway, Damper Cave; D, Nanocochlea stylesae holotype, C.439398, Malani Creek; EG, Nanocochlea damperensis E, C.203685, paratype, first crossing, Damper Creek; F, holotype, C.439399, first crossing, Damper Creek, F, C.201282, Near New River Lagoon, E side, Limestone Ck, where tagged track crosses; H, I, Pseudotricula progenitor H, paratype, C.439397, Persephone Streamway, Bauhaus; I, holotype, C.439396, Persephone Streamway, Bauhaus. Scales: 500 m m.

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2005 Magnolia Press 37HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 15. Shells of Nanocochlea species and Pseudotricula progenitor A C, Nanocochlea exigua A, C.203674, outside Damper Cave, Damper Creek B, C.166855, Persephone Pot, Bauhaus, C, C.201465, main streamway, Damper Cave; DF, Nanocochlea stylesae D, C.201476, paratype, Malani Creek, E, F, C.203686, first crossing, Damper Creek; G-I, Nanocochlea damperensis G,H, C.303685, seepages at Bauhaus entrance, I, C.203685, paratype, first crossing, Damper Creek; JL, Pseudotricula progenitor J, L, C.201463, paratype, Persephone streamway, Bauhaus, K, C.201454, Persephone, Bauhaus. Scales: 1 mm.

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PONDER ET AL.38 2005Magnolia Press 1074ZOOTAXA FIGURE 16. Opercula of Nanocochlea species and Pseudotricula progenitor A C, Nanocochlea exigua C.203677, paratypes, Cane Toad Abuse Streamway, Damper Cave; D F, Nanocochlea damperensis ; CE, C.203685, paratypes, first crossing, Damper Creek; F, C.201282, Near New River Lagoon, E side, Limestone Ck, where tagged track crosses; G, H, Pseudotricula progenitor C.201821, main streamway, Bauhaus; A, D, G, outer side; B, C, E, F, H, inner side; Scales: A, B, D, 200 m m, C, EH, 100 m m.Female genital system (Fig. 18C, D). Ovary of 0.50.7 whorls; oviduct extends to posterior edge of bursa copulatrix; joins bursal duct in front of posterior pallial wall at junction of albumen and capsule glands to half way between posterior pallial wall and capsule gland. Bursa copulatrix large, extending to posterior pallial wall; globular to pyriform; with bursal duct arising from middle of anterior edge of bursa, straight or with undulations. Seminal receptacle at middle of inner wall of bursa copulatrix; pyriform. More than half of albumen gland in front of posterior pallial wall; capsule gland about same length as albumen gland; compressed oval in section; anterior end blunt; ventral channel with distinct muscular vestibule; genital opening overlapping anterior end of capsule gland. Distribution and habitat Bauhaus and Damper Cave in low energy small streams with mixed substrate.

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2005 Magnolia Press 39HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 17. Radulae of Nanocochlea species Pseudotricula progenitor A C, Nanocochlea exigua A, C.201475, Honey and Cream Streamway, Damper Cave, B, D, C.201465, main streamway, Damper Cave; DE, Nanocochlea stylesae C.201476, paratypes, Malani Creek; GK, Nanocochlea damperensis G, H, C.203685, paratypes, first crossing, Damper Creek, IK, C.201282, Near New River Lagoon, E side, Limestone Ck, where tagged track crosses; LO, Pseudotricula progenitor C.201821, main streamway, Bauhaus. A, E, central and lateral teeth; B, F, I, M, central, lateral, inner and outer marginal teeth; C, D, J, N central teeth; G, L, lateral teeth; H, central lateral inner and outer marginal teeth; K, outer marginal teeth. Scales: A, GH, K, L, 10 m m; B, I, M, 20 m m; C, D, J, N, 5 m m.

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PONDER ET AL.40 2005Magnolia Press 1074ZOOTAXA FIGURE 18. Female reproductive system of Nanocochlea species and Pseudotricula progenitor A, Nanocochlea exigua C.203677, paratype, Cane Toad Abuse Streamway, Damper Cave; B, Nanocochlea damperensis C.203685, paratype, first crossing, Damper Creek; C, D, Pseudotricula progenitor C.201821, main streamway, Bauhaus. Scales: 500 m m. Abbreviations: ag, albumen gland; bc, bursa copulatrix; bd, bursal duct; cg, capsule gland; co, coiled oviduct; po, genital opening; pw, pallial wall; sr, seminal receptacle; vc, ventral channel. Remarks Pseudotricula progenitor shares characters of both Nanocochlea and Pseudotricula The small-sized, tall-spired shell is typical of Nanocochlea but the (narrowly) reflected outer lip, straight spire outline and subsutural indentation are characters found in typical Pseudotricula taxa. The protoconch microsculpture, while not able to be examined in detail, appears to be of the plesiomorphic type (shared by Nanocochlea and Pseudotricula s.s.), as are the radular characters, which are like those of Nanocochlea and P. arthurclarkei and P. conica It is probable that this species most closely resembles the ancestral Pseudotricula that gave rise to the Precipitous Bluff cave radiation. Genus Nanocochlea Ponder and Clark (in Ponder et al .), 1993 Type species: Nanocochlea monticola Ponder and Clark in Ponder et al ., 1993 (original designation).

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2005 Magnolia Press 41HYDROBIIDAE S.L. 1074ZOOTAXA Description Shell Minute to small in size (adults range from 1.6 to 2.5 mm in length), pupiform to elongate pupiform or elongate-conic. Protoconch of about 1.41.5 whorls, typically sculptured with uniform pitting; separation of protoconch from teleoconch distinct, with varix-like border. Teleoconch with spire much longer than length of aperture. Aperture ovate, slightly angled posteriorly, inner lip attached to parietal wall or partially to completely detached, although not markedly so, usually lower part not much separated from base; outer lip orthocline to opisthocline, weakly thickened, simple or with very slight reflection. Periphery rounded, base simple, imperforate in adults, rarely narrowly umbilicate in juveniles. All species described below are semi-opaque to opaque, white with a thin pale yellowish periostracum. Operculum. Ovate, paucispiral, flat, yellowish; columellar edge slightly convex, outer edge strongly convex; outer surface simple, paucispiral, nucleus markedly eccentric; inner surface usually smooth with white smear or (in one species) large, broad peg. Radula. Central teeth large, cutting edge broad, indented mid dorsally, with 45 cusps on either side of narrow sharp median cusp; narrow, unthickened lateral projections emerge at about 45 from mid laterally; base with short, narrow, rounded basal projection not extending beyond lateral projections; two basal cusps emerge from ventral face of tooth on either side of basal projection, innermost longest. Lateral teeth with short cutting edge, about 45 small cusps on inner side of narrow median cusp and 46 small cusps on outer side; neck prominent, near vertical; lateral flange more than twice as long as cutting edge; prominent U-shaped projection on base below cutting edge; inner edge short, inner side of base excavated. Inner marginal teeth with 2138 tiny cusps on rather wide cutting edge (ratio of cutting edge to shaft on inner marginal teeth about ); sides approximately parallel, outer edge thickened. Outer marginal teeth narrow, thickened on inner edge, distal end with 1729 minute cusps; ratio of cutting edge to shaft about Head-foot. Simple, unpigmented, with long cephalic tentacles, unpigmented to lightly pigmented eyes present in weak bulges at outer bases of tentacles; snout of moderate length, tapering, weakly bilobed distally. Foot short, rounded posteriorly. Non-genital anatomy. Pallial cavity elongate, osphradium large and oval, towards posterior end of ctenidium; ctenidium absent or with small (reduced from normal triangular type) ctenidial filaments; efferent vein very short to long (ctenidium occupying nearly entire length of pallial cavity to about ). Hypobranchial gland variably developed. Kidney and pericardium usually about in pallial roof; renal gland orientated longitudinally. Stomach usually with anterior chamber larger than posterior; long style sac moderately long; no posterior caecum. Rectum with long S-shaped coil; overlying but not indenting pallial oviduct. Male reproductive system. Prostate gland about within pallial cavity; compressed to oval in section, with very thin ventral wall; pallial vas deferens opens at about length of pallial portion. Penis located on right side of head well behind base of right

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PONDER ET AL.42 2005Magnolia Press 1074ZOOTAXA tentacle; lacking but has an inconspicuous to moderately developed swelling in the distal portion just behind a papilla-like distal end; distal portion long and tapering; medial part simple and parallel sided; basal part wide to moderate. Female reproductive system. Ovary simple sac; coiled oviduct smooth, firm, not embedded in connective tissue, initially inverted U-shape, usually straight distal to seminal receptacle, usually reaching posterior end of bursa (sometimes a little more or less) then sharply bent to run anteriorly; Seminal receptacle ovoid to pyriform, with short duct, opening to oviduct, located opposite middle to ventral part of left side of bursa copulatrix. Bursa copulatrix medium to rather large, globular to pyriform, posterior to albumen gland except for small overlap on right side, either behind posterior pallial wall or extending to it; bursal duct arrises from anterior or ventro-anterior wall, simple to undulating, joins oviduct dorsally at posterior pallial wall or just in front of it. Common duct straight. Albumen gland with or more within pallial roof, shorter to longer than capsule gland. Capsule gland compressed-oval in section, divided into three glandular zones, short anterior and posterior translucent white zones and long yellowish to orange middle zone; anterior end blunt to steeply tapering; genital opening short, terminal to slightly anterior of capsule gland. Ventral channel extended into weakly to moderately developed vestibule anteriorly. No brood pouch. Distribution and habitat Members of the genus are known from mountain lakes, seepages, streams and caves in southern Tasmania. Remarks Nanocochlea differs from Pseudotricula in its small, tall-spired pupiform to elongateconic shell with a thin, simple, orthocline to opisthocline, usually non-reflected outer lip. There are no obvious anatomical differences between the two genera and there is a rather minor radular differencethe basal tongue of the central teeth is narrower in Nanocochlea The outer shaft on the lateral teeth is longer in species of Nanocochlea than in typical species of Pseudotricula but P. conica P. arthurclarkei and P. progenitor also have a long shaft. Nanocochlea exigua n. sp. Figures 6H; 8F; 14AC; 15AC; 16AC; 17AC; 18A. Type material Holotype: AMS C.439393, Cane Toad Abuse streamway, Damper Cave, PB1-2A, 28 MAR 1994. Paratypes: AMS C. 203677, same data (20+ dry, 20+ wet); QVM, 9:20541 (5 wet).

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2005 Magnolia Press 43HYDROBIIDAE S.L. 1074ZOOTAXA Other material examined (all AMS) Bauhaus : C.201820, stn PB6-1A (8 dry, 3 wet); C.201819, stn PB6-1A (1 dry); C,166855, Persephone Pot, stn PB17-8R (4 dry, 12 wet); C.201810, same locality, middle and upper streamway, stn PB17-2a.2 (7 dry); C.201814, same data (6 dry, 13 wet); C.201818, same locality, lower streamway, stn PB17-2a (1 dry); C.201274, Persephone, stn 7 (6 dry, 20+ wet); C.201461, Persephone streamway, stn 8 (7 dry, 20 wet); C.201455, same locality, stn 9 (6 dry, 20+ wet); C.201456, same locality, stn 10 (1 dry); C.201479, same locality, stn 10 (8 dry, 20+ wet); C.201462, Screaming Stals streamway, stn 12 (11 dry, 20+ wet); C.201480, same locality, stn 12 (5 dry); C.201457, same locality, stn 13 (1 dry). Damper Cave : C.203683, main streamway near entrance, stn PB1-1A (3 dry); C.201465, main streamway, stn 6 (7 dry, 14 wet); C.439395, same data (2 dry, figd specimens); C.201482, Cane Toad Abuse streamway, stn 2 (8 dry, 20+ wet); C.201467, seep near The Keg, stn 3 (9 dry, 20+ wet); C.201475, Honey And Cream streamway, stn 4 (8 dry, 20+ wet); C.203679, Cane Toad Abuse streamway, stn PB1-2A (1 dry). Damper Creek : C.203674, 10m outside Damper Cave, stn PBs-2A (2 dry, 1 wet). Quetzalcoatl Conduit : C. 203672, stn PB3-1C (1 dry). Etymology Exiguus Latin, small, short. Description Shell (Fig. 6H; 14AC; 15AC). Length up to 2.2 mm; elongate-conic (SW/SL 0.390.61, mean 0.50, n = 37); spire tall, straight to slightly convex in outline; last whorl evenly rounded or with subshoulder depression; suture simple or indented/impressed. Protoconch microsculpture uniform (Fig. 6H). Teleoconch up to 3.7 whorls in adult; aperture oval to pear-shaped; small, shorter than spire (AL/SL 0.300.45, mean 0.36, n = 37); outer lip orthocline to opisthocline; weakly thickened in adult, straight; posterior notch absent; inner lip thin and narrow, in partial contact or narrowly separated from parietal wall. Dimensions. See Table 10.TABLE 10. Shell dimensions and teleoconch whorl counts of Nanocochlea exigua n. sp. SLSWALAWBWCVTW Holotype2.030.80.730.511.340.13.3 Figured specimens C.2014651.750.830.60.541.190.13.3 1.730.850.650.61.20.113.4 C263677(20) Minimum1.640.850.590.551.130.093.00 Maximum2.180.990.780.691.410.183.70 Mean1.890.920.670.621.240.123.37 Standard Dev.0.140.040.040.040.070.020.21......continued on the next page

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PONDER ET AL.44 2005Magnolia Press 1074ZOOTAXA Operculum (Fig. 16AC). Inner surface with white smear and 1 large peg. Eyes Unpigmented. Pallial cavity (Fig. 8F). Ctenidium rudimentary with no filaments; osphradium present, as in other taxa; hypobranchial gland thick to moderately developed; renal organ extends forward ca. into pallial cavity; pericardium more than in pallial roof. Radula (Fig. 17AC). Central teeth : dorsal edge with deep indentation; 45 lateral cusps, median cusp narrow, blunt to sharply pointed, about twice as long as adjacent cusps. Lateral teeth : dorsal edge with shallow to moderate indentation; with 46 cusps on outer and 45 on inner side; median cusp narrow to medium width, blunt to sharply pointed, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection bluntly pointed. Marginal teeth : Inner with 2126 cusps; outer with 1720 cusps. Stomach Stomach with posterior chamber a little smaller than anterior chamber. Male genital system Testis of 0.61.5 whorls; prostate gland about in pallial roof, oval to kidney-shaped; compressed in section. Pallial vas deferens straight. Penis with weak swelling in mid-distal portion; distal end long, papilla-like; medial section parallel sided, of medium length; penial duct in medial section strongly undulating; base of penis very to moderately wide; with moderate folds; penial duct straight to weakly undulating. Female genital system (Fig. 18A). Ovary of 0.60.7 whorls; oviduct extends to posterior edge of bursa copulatrix or slightly less; joins bursal duct at posterior pallial wall. Bursa copulatrix large, extending to posterior pallial wall; elongately oval to pyriform; with bursal duct arising from antero-ventral to ventral edge of bursa; straight or with undulations. Seminal receptacle at middle of inner wall of bursa copulatrix or near mid ventral edge; ovoid to pyriform. Two thirds to all of albumen gland in front of posterior pallial wall; capsule gland about same length as albumen gland; oval in section; anterior end tapering to blunt; ventral channel simple, approximately parallel-sided throughout; vestibular area indistinct to distinct; genital opening overlapping anterior end of capsule gland to terminal. Distribution and habitat Found mainly in small, low energy streams and seepages in the caves. A few specimens were also found in Damper Creek 10 m outside Damper Cave.TABLE 10 (continued) SLSWALAWBWCVTW C.201465(14) Minimum1.100.730.480.430.840.052.55 Maximum1.960.990.690.641.260.173.65 Mean1.650.850.600.561.100.113.31 Standard Dev.0.230.080.060.070.120.030.30

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2005 Magnolia Press 45HYDROBIIDAE S.L. 1074ZOOTAXA Remarks Nanocochlea exigua differs from other congeners in the operculum bearing a single large peg and in lacking a ctenidium (gill). In the former respect, it resembles some species included in Austropyrgus Cotton, 1942 (Clark et al 2003), as well as a number of other hydrobiid genera, including some found outside Australia. There are, however, several important characters that link N. exigua with Nanocochlea There is no gastric caecum (present in Austropyrgus and several other genera bearing similar opercular pegs), the rectum is strongly S-shaped whereas in other genera with opercular pegs it is usually straight to arched. The female genital anatomy and penial morphology is also very like other species of Nanocochlea and unlike that of Austropyrgus Assuming that the lack of the ctenidium in N. exigua has evolved from the presumed apomorphic condition seen in the other Precipitous Bluff congeners, and that no other species of Nanocochlea are known to possess an opercular peg, it is possible that the peg in this species is a secondary condition. Nanocochlea stylesae n. sp. Figures 4C; 6I, J; 14D; 15DF; 17D, E. Type material Holotype: AMS C.439398, Malani Creek, stn 14, 28 DEC 1991. Paratypes: AMS C.201476, same data, 1 dry, 20+ wet; QVM, 9:20542 (5 wet). Other material examined First crossing, Damper Creek: AMS C.203686, stn PBs-1A (2 dry, 1 wet). Etymology Named for Julie Styles who assisted one of us (SE) in the collection of the bulk of the material used in this paper. Description Shell (Fig. 6IJ; 14D; 15DF). Length up to 2.0 mm; narrowly conical (SW/SL 0.470.56, mean 0.52, n = 21); spire tall, straight to slightly convex in outline; last whorl evenly rounded; suture simple or indented/impressed. Protoconch microsculpture not examined in detail. Teleoconch up to 3.5 whorls in adult; umbilicus closed in juveniles; aperture oval to pear-shaped; moderate size, shorter than spire (AL/SL 0.330.42, mean 0.37, n = 21); outer lip orthocline, weakly thickened in adult, straight, without reflection; posterior notch absent; inner lip thin and narrow, in partial contact or narrowly separated from parietal wall. Dimensions. See Table 11.

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PONDER ET AL.46 2005Magnolia Press 1074ZOOTAXA TABLE 11. Shell dimensions and teleoconch whorl counts of Nanocochlea stylesae n.sp. Operculum Inner surface simple or with small slightly thickened lump near nucleus (possibly rudimentary peg). Eyes Unpigmented. Pallial cavity Ctenidium narrow; 911 very small filaments; osphradium near posterior end of ctenidium; hypobranchial gland very weakly-developed to apparently absent; renal organ extends forward ca. into pallial cavity; pericardium up to in pallial roof. Radula (Fig. 17D, E). Central teeth : dorsal edge with deep indentation; 5 lateral cusps, median cusp narrow, blunt to sharply pointed, about twice as long as adjacent cusps. Lateral teeth : dorsal edge with shallow to moderate indentation; with 56 cusps on outer and 5 on inner side; median cusp narrow, sharply pointed, less than twice as long as adjacent cusps; ratio of cutting edge to shaft about ; basal projection pointed to bluntly pointed. Marginal teeth : Inner with 2438 cusps; outer with 2028 cusps. Stomach Stomach with posterior chamber and anterior chamber about equal in size. Male genital system. Testis of about 2.0 whorls; prostate gland about in pallial roof; oval to elongate pyriform; oval in section. Pallial vas deferens strongly undulating at prostate, straight between prostate and penis and at base of penis. Penis (Fig. 4C) with weak swelling in mid-distal portion; distal end long, papilla-like; medial section tapering to parallel sided; of medium length; penial duct straight to slightly undulating; base of penis moderately wide; with moderate folds; penial duct straight to weakly undulating. Female genital system Ovary of 0.7 whorls; oviduct extends to posterior edge of bursa copulatrix; joins bursal duct at posterior pallial wall. Bursa copulatrix of medium size, not extending to posterior pallial wall; globular; with bursal duct arising from middle of anterior edge of bursa; straight. Seminal receptacle at middle of inner wall of bursa copulatrix; ovoid. One third of albumen gland in front of posterior pallial wall; capsule gland about same length as albumen gland; circular in section; anterior end tapering to blunt; ventral channel simple, approximately parallel-sided throughout; vestibular area indistinct; genital opening terminal. Distribution and habitat This species is apparently restricted to surface streams flowing from Precipitous SLSWALAWBWCVTW Holotype2.041.010.751.131.370.133.5 C.201476 paratypes (20) Minimum1.670.900.630.581.010.042.40 Maximum2.001.070.781.091.410.123.50 Mean1.900.990.710.691.250.093.19 Standard Dev.0.090.060.050.120.110.020.25

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2005 Magnolia Press 47HYDROBIIDAE S.L. 1074ZOOTAXA Bluffit is known from Malani Creek, approximately 1.3 km north of Damper Creek and in Damper Creek itself where a few specimens were found together with N. damperensis. Remarks Nanocochlea stylesae is somewhat similar in shape to N. exigua but has a short ctenidium, lacks a distinct opercular peg and tends to have more cusps on the inner marginal teeth. It differs from N. damperensis in its narrower shell (P<0.001) and less convex whorls and fewer ctenidial filaments. Nanocochlea damperensis n. sp Figures 4D; 6KM; 8G; 14EG; 15GH; 16DF; 17GK; 18B. Type material Holotype: AMS C.439399, first crossing, Damper Ck, stn PBs-1A, 29 MAR 1994. Paratypes: AMS C.203685, same data, 20+ dry, 20+ wet; QVM, 9:20543 (5 wet); AMS C.439400, same data (1 dry figured paratype). Other material examined (all AMS) Damper Creek : C.203673, 10m outside Damper Cave, stn PBs-2A (22 dry, 20+ wet); C.201282, near New River Lagoon, E side, Limestone Ck, where track crosses creek, JW 49, 15 FEB 1988; 20+ dry, 20 wet; figured specimen from same lot, C.343820. Bauhaus : C.303685, cave entrance. Etymology Named after Damper Creek. Description Shell (Figs. 6KM; 14EG; 15G, H). Length up to 2.3 mm; conical (SW/SL 0.500.69, mean 0.61, n = 63); spire moderate to tall, convex in outline; last whorl evenly rounded; suture indented/impressed. Protoconch microsculpture of small shallow pits and pustules. Teleoconch up to 3.3 whorls in adult; umbilicus present only in juveniles; aperture oval to pear-shaped; moderate size, shorter than spire (AL/SL 0.370.47, mean 0.42, n = 63); outer lip, orthocline, weakly thickened in adult, straight to slightly convex, without reflection; notch usually present in posterior corner of aperture; outer lip orthocline to weakly opisthocline; inner lip thin and narrow, in partial contact or narrowly separated from parietal wall. Dimensions. See Table 12.

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PONDER ET AL.48 2005Magnolia Press 1074ZOOTAXA TABLE 12. Shell dimensions and teleoconch whorl counts of Nanocochlea damperensis n. sp. Operculum (Fig. 16DF). Inner surface with or without white smear and, in some, 35 weak ridges (reduced pegs). Eyes Slightly pigmented. Pallial cavity (Fig. 8G). Ctenidium narrow; 1213 very small filaments; osphradium between posterior end and middle of ctenidium; hypobranchial gland thick to moderately developed; renal organ extends forward ca. into pallial cavity; pericardium more than in pallial roof. Radula (Fig. 17GK). Central teeth : dorsal edge with moderate to deep indentation; 45 lateral cusps, median cusp narrow, sharply pointed, about twice as long as adjacent cusps. Lateral teeth : dorsal edge with shallow indentation; with 56 cusps on outer and 46 on inner side; median cusp narrow, sharply pointed, less than, to about twice as long as, adjacent cusps; ratio of cutting edge to shaft about ; basal projection pointed. Marginal teeth : Inner with 2027 cusps; outer with 2026 cusps. Stomach Stomach with posterior chamber a little smaller than anterior chamber. Male genital system. Testis of 1.01.25 whorls; oval to kidney-shaped; oval in section. Pallial vas deferens straight to slightly undulating at prostate; straight between prostate and penis and at base of penis. Penis (Fig. 4D) with weak swelling in mid-distal portion; distal end long, papilla-like; medial section parallel sided; of medium length; penial duct in medial section strongly undulating; base of penis moderately wide; with moderate folds; penial duct straight to undulating. SLSWALAWBWCVTW Holotype2.051.360.920.861.530.22.9 Figured paratype2.071.210.840.831.530.193.0 Figured specimen (C.440551) 1.971.170.860.771.490.182.6 C.203685, paratypes (20) Minimum1.921.200.830.781.420.142.60 Maximum2.291.451.000.931.680.193.10 Mean2.081.310.900.851.550.162.81 Standard Dev.0.100.060.040.050.080.010.12 C.201282(20) Minimum1.871.160.800.721.410.142.60 Maximum2.191.380.970.891.650.213.30 Mean2.021.270.880.811.510.172.82 Standard Dev.0.080.080.040.040.070.020.19 C.303685(20) Minimum1.600.870.630.531.100.100.37 Maximum2.081.160.080.811.420.214.90 Mean1.841.050.730.711.270.154.69 Standard Dev.0.130.070.050.060.080.030.13

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2005 Magnolia Press 49HYDROBIIDAE S.L. 1074ZOOTAXA Female genital system (Fig. 18B). Ovary of 1.0 whorls; oviduct extends to posterior edge of bursa copulatrix or slightly less; joins bursal duct just in front of posterior pallial wall. Bursa copulatrix of medium size, not extending to posterior pallial wall; globular; with bursal duct arising from middle of anterior edge of bursa; straight. Seminal receptacle at middle of inner wall of bursa copulatrix; ovoid or pyriform. More than of albumen gland in front of posterior pallial wall; capsule gland up to ? length of albumen gland; oval to compressed oval in section; anterior end blunt; ventral channel with indistinct to distinct vestibule; genital opening overlapping anterior end of capsule gland. Distribution and habitat Found in Damper Creek near the entrance to Damper Cave but was not found in samples from inside Damper Cave (in the same creek). A sample of mainly empty shells from seepages at the entrance of Bauhaus cave (Fig. 15G, H) is also attributed to this species and a single dead shell (AMS C. 203714) from inside Bauhaus (Persephone Pot) is presumably washed in from a surface seepage. Specimens from the lower end of Limestone Creek on the eastern side of New River Lagoon are also attributed to this species (see remarks). The site in this creek is approximately 1.7 km S of the lower end of Damper Ck, although about 3.5 km separate the two locations where the samples were collected. Remarks Nanocochlea damperensis differs from the other Nanocochlea described to date in its broadly pupiform shell shape. Specimens from Limestone Creek are almost identical to the type material, although have more ctenidial filaments (1417), 56 lateral cusps on the central teeth and the pallial vas deferens undulates rather than being straight. Other anatomical differences are the seminal receptacle is ovoid, not globular, the capsule gland has a terminal opening in some specimens and most specimens lack a distinct white smear on the operculum. Given their close geographic proximity and overall similarity, these two populations are considered to be conspecific pending more detailed investigation. The very weak (rudimentary) multiple pegs on the operculum of this species resemble those seen in P. conica and P. arthurclarkei and have not been observed in other taxa included in Nanocochlea The shell-shape of this species resembles that of some taxa included in Austropyrgus (see Clark et al 2003). Members of that genus have distinct pegs on the operculum, lack an S-shaped rectum and have three or more basal cusps on the central teeth. Discriminant function analysis A discriminant function analysis was performed separately on the species in each genus using the shell measurements, the convexity ratio and whorl counts to test the efficacy of

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PONDER ET AL.50 2005Magnolia Press 1074ZOOTAXA these data in defining the taxa. Pseudotricula elongata was excluded from the Pseudotricula analysis due to insufficient measurements, as was P. progenitor which was included in the Nanocochlea analysis because of its similar shell shape to the other members of that group. Both these species are readily distinguished from their congeners using shell shape (see remarks under those taxa). In the Pseudotricula analysis, involving 130 specimens, four of the five species were 100% discriminated while one specimen of P. arthurclarkei was misclassified as P. conica (Wilks lambda = 0.021, approx. F = 29.487, df = 28, 430, p-tail < 0.0001). In the analysis involving the Nanocochlea taxa and P. progenitor a total of 143 specimens, all P. damperensis were correctly classified, while N. exigua had five (of 37) specimens incorrectly classified with N. stylesae N. stylesae had two classified as N. exigua and P. progenitor had one misclassified as N. damperensis (Wilks lambda = 0.025, approx. F = 47.548, df = 21, 382, p-tail < 0.0001). The plots of the first two discriminant scores are shown in Figures 19 and 20.FIGURE 19. Plot of discriminant scores derived from shell measurements, convexity ratio and whorl counts for Pseudotricula species ( P. elongata and P. progenitor excluded). P. eberhardi ; x P. arthurclarkei ; + P. auriforma ; P. conica ; P. expandolabra Gaussian bivariate ellipses are indicated for each sample (P=0.6827).

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2005 Magnolia Press 51HYDROBIIDAE S.L. 1074ZOOTAXA FIGURE 20. Plot of discriminant scores derived from shell measurements, convexity ratio and whorl counts for Nanocochlea and Pseudotricula progenitor N. damperensis ; x N. exigua ; N. stylesae ; + Pseudotricula progenitor Gaussian bivariate ellipses are indicated for each sample (P=0.6827).Cladistic analysis A maximum parsimony analysis was preformed on a dataset of 25 taxa (Table 13) and 60 characters (Table 14), all unordered and of equal weight (1). Two characters were parsimony-uninformative. A heuristic search of the dataset was undertaken with 100 random addition replicates (one tree held at each step) utilizing branch-swapping by treebisection-reconnection (TBR). The steepest descent option was not in effect and no constraints were enforced. The analysis resulted in six equally parsimonious trees of 215 steps (Consistency Index =0.502, Retention Index = 0.730, Rescaled Consistency Index = 0.367, Homoplasy Index = 0.498). The strict consensus tree is shown in Figure 21 with bootstrap (heuristic search, 1000 replicates) values of greater than 50% given. The analysis included 15 taxa that were not part of the Precipitous Bluff fauna. Of these, the brackish-water genus Tatea Tenison Woods, 1879 (see Ponder et al 1991) was used to root the tree. Other outgroup taxa included three species of Austropyrgus (see Clark et al 2003) and four species of Beddomeia Petterd, 1889 and Phrantela Iredale, 1943 (see Ponder et al ., 1993). The analysis also included the three previously described

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PONDER ET AL.52 2005Magnolia Press 1074ZOOTAXA species of Nanocochlea (Ponder et al 1993). The tree obtained confirms the monophyly of the outgroup genera with the exception of N. parva Ponder & Clark, 1993 being included with Phrantela This result is not unexpected, as there are several features of that taxon that are shared with Phrantela and it was only tentatively included in Nanocochlea by Ponder et al (1993). Consequently, we include this taxon in Phrantela pending a more detailed analysis. The species that comprise the Precipitous Bluff radiation are included in a single well supported clade that is comprised of two subclades, one of Nanocochlea (the two previously described taxa form a subclade within this) and another clade that is here referred to as Pseudotricula Nanocochlea is very weakly supported, with the only changes (both unambiguous) being shell size and the shape (in transverse section) of the prostate gland. Pseudotricula is a little better supported, with four unambiguous characters (although lacks bootstrap support): inner surface of the operculum with white smear (strongly convergent), the anterior location of the joining of the bursal duct and oviduct, relatively more of albumen gland in pallial roof and a small anterior shift in the position of the female genital opening. Ambiguous changes included: relative size of the stomach chambers, the position and shape of the bursa copulatrix, the shape of the capsule gland in section and the vestibule.TABLE 13. Data matrix used in cladistic analysis. The list of characters is given in Table 14.12345678910111213141516171819202122 Tatea huonensis 3111112211132133111211 Austropyrgus cooma 3113111211132133111211 Austropyrgus niger 311,33111211132134111211 Austropyrgus petterdianus 1113111211132133111211 Beddomeia launcestonensis 2252122111131,241-111212 Beddomeia palludinella 335312211113341-111212 Beddomeia hullii 221212211113111-1,22121,22 Beddomeia bellii 235212211113121-221212 Phrantela marginata 211211211113111-112211 Phrantela conica 211211211111,2141-212211 Phrantela warwicki 211211211113111-112211 Phrantela pupiformis 211211211113111-112211 Nanocochlea monticola 111311221111221-132241 Nanocochlea pupoidea 111311221112221-132241 "Nanocochlea" parva 111211211113141-112241 Nanocochlea damperensis 111213121111,2222,33133241 Nanocochlea exigua 11121,21221111,2223114-1,241 Nanocochlea stylesae 111211221111221-133141 Pseudotricula progenitor 21131,21221111,2222-133241 Pseudotricula eberhardi 234211322123322-133241 Pseudotricula arthurclarkei 234311221113233,431331,241 Pseudotricula auriforma 3333213222233?2-1332,341 Pseudotricula conica 23431,21221113232,32133241 Pseudotricula expandolabra 234321322223322-133241 Pseudotricula elongata 211321321,22213????????? ......continued on the next page

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2005 Magnolia Press 53HYDROBIIDAE S.L. 1074ZOOTAXA TABLE 13 (continied)232425262728293031323334353637383940414243 44 Tatea huonensis 311,231221,2??1,22231221,21111 Austropyrgus cooma 3212,311222?221333221111 Austropyrgus niger 311211,2222?221333221111 Austropyrgus petterdianus 31,21212222?221333211111 Beddomeia launcestonensis 121313122?222213111122 Beddomeia palludinella 121312,3132?222213111112 Beddomeia hullii 12131213??222213111122 Beddomeia bellii 12131213??1,222113111122 Phrantela marginata 122313311?222232121132 Phrantela conica 121312322?222232111132 Phrantela warwicki 122312312?2222,31,23111112 Phrantela pupiformis 121313311?21,22211111112 Nanocochlea monticola 22231,22312?222122,4222331 Nanocochlea pupoidea 221,2322312?1,21,22122212321 "Nanocochlea" parva 132312311?1222221111?2 Nanocochlea damperensis 22,3222231311,21,22122212321 Nanocochlea exigua 232122,331,2211,212132212321 Nanocochlea stylesae 232222,3312,3121,21122222321 Pseudotricula progenitor 22,322,3223121221,2132212321 Pseudotricula eberhardi 211321222211211,22211,2321 Pseudotricula arthurclarkei 222321,231,21,21223122212321 Pseudotricula auriforma 2113,422222,32111,2132212321 Pseudotricula conica 22,31,2321,2,331,221221132212321 Pseudotricula expandolabra 211,23222222111132212321 Pseudotricula elongata ?????????????????????? 45464748495051525354555657585960 Tatea huonensis 112312,714121121,215 Austropyrgus cooma 11121712111321-3 Austropyrgus niger 21121711122321-2 Austropyrgus petterdianus 111217111323,42222 Beddomeia launcestonensis 222116141231,221-1 Beddomeia palludinella 222213?31,221111-1 Beddomeia hullii 222213131,221,2,3211,213 Beddomeia bellii 1,2122171212321223 Phrantela marginata 11233425242311-2 Phrantela conica 23232425221211-3 Phrantela warwicki 12232415242,3211-4 Phrantela pupiformis 13232425243111-4 Nanocochlea monticola 12,3221614123111-3,4 Nanocochlea pupoidea 1222121414311,21-3 "Nanocochlea" parva 13232425241211-3 Nanocochlea damperensis 11,23217141332,3211,22 Nanocochlea exigua 11,223271414221,21,2-2,3 Nanocochlea stylesae 122211141221,21,21-3 Pseudotricula progenitor 1242,321,61413232222 Pseudotricula eberhardi 1222111412,32,31,221-4 Pseudotricula arthurclarkei 11,24211,61412,3231,21,21,22 Pseudotricula auriforma 11,2431,21,5,61413131,2122 Pseudotricula conica 11,2,3432,35,612,412,32,331,21-2 Pseudotricula expandolabra 12432,31,512,4131321,21,22,3 Pseudotricula elongata ????????????????

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PONDER ET AL.54 2005Magnolia Press 1074ZOOTAXA TABLE 14 The list of characters used in the cladistic analysis. The data matrix is given in Table 13.1. Maximum dimension of shell: 1, very small (1.5-2.5 mm); 2, small (2.6-4.0 mm); 3, large 4.16 mm. 2. Aperture size: 1, less than to twice as short as spire; 2, about equal to spire; 3, 1 to less than 2x longer than spire. 3. Shell shape: 1, elongate; 2, medium conic; 3, broadly conic; 4, ovate. 4. Spire outline: 1, concave; 2, straight; 3, convex. 5. Sub-shoulder depression: 1, absent; 2, present. 6. Umbilicus: 1, absent in adults and juveniles; 2, present in adults (open); 3, present only in juveniles. 7. Aperture shape: 1, rounded; 2, oval to pear-shaped; 3, D-shaped. 8. Outer lip: 1, thin in adult; 2, weakly to moderately thickened. 9. Outer lip reflection: 1, absent or weak; 2, strong. 10. Aperture flaring: 1, not flared; 2, flared. 11. Posterior (anal) notch: 1, absent; 2, present. 12. Slope of outer lip: 1, orthocline; 2, opisthocline; 3, prosocline. 13. Inner lip: 1, thin; 2, moderately thickened and narrow; 3, thick and wide. 14. Protoconch microsculpture: 1, coarse pits; 2, small pustules; 3, pits and pustules; 4, very fine pitting. 15. Operculum inner surface: 1, simple; 2, with white smear only; 3, with white smear and pegs; 4, with pegs only. 16. Number of opercula pegs: 1, single large peg; 2, up to 2 small to medium pegs; 3, 3-5 pegs; 4, more than 5 pegs. 17. Posterior pallial tentacle: 1, absent; 2, present. 18. Ctenidium: 1, well developed, wide; 2, narrow; 3, rudimentary; 4, absent. 19. Ctenidium length relative to pallial cavity: 1, extends all or most of pallial cavity; 2, only in anterior ?; 3, only in anterior 20. Osphradium position relative to ctenidium: 1, near posterior end; 2, between posterior end and middle, 3, near middle 21. Rectum: 1, straight or with slight arch; 2, with prominent arch; 3, short S-shape; 4, long Sshape. 22. Renal organ orientation: 1, longitudinal; 2, circular to transverse. 23. Basal cusps on central teeth: 1, one pair; 2, usually two pairs; 3, usually three or more pairs. 24. Dorsal edge of central teeth: 1, with shallow indentation; 2, moderately indented; 3, deeply V shaped. 25. Median cusp of central teeth: 1, medium width; 2, narrow. 26. Length of median cusp of central teeth (relative to adjacent cusps): 1, about 3 times longer; 1: about 2 times longer; 3, less than 2 times longer; 4, about equal. 27. Lateral edges of central teeth: 1, straight; 2, bent. 28. Dorsal edge of lateral teeth: 1, straight to lightly convex; 2, with shallow indentation; 3, moderately indented; 3, deeply V-shaped. 29. Shape of U-shaped basal tongue of central teeth: 1, wide; 2, medium; 3, narrow. 30. Median cusp of lateral teeth: 1, narrow; 2, medium; 3, broad. 31. Basal projection of lateral teeth: 1, U-shaped; 2, bluntly pointed; 3, pointed. 32. Cutting edge and shaft of lateral teeth: 1, shaft more than twice as long; 2, shaft twice or less than twice as long. 33. Cusps on inner marginal teeth: 1, up to 21 cusps; 22 cusps or more. 34. Cusps on outer marginal teeth: 1, up to 20 cusps; 21 cusps or more.......continued on the next page

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2005 Magnolia Press 55HYDROBIIDAE S.L. 1074ZOOTAXA TABLE 14 (continued) 35. Stomach chambers: 1, posterior and anterior chambers about equal; 2, posterior chamber a little smaller; 3, posterior chamber much smaller; 4, with posterior chamber larger. 36. Stomach caecum: 1, absent; 2, small; 3, large. 37. Prostate shape in transverse section: 1, broadly oval to circular; 2, oval; 3, compressed; highly compressed. 38. Ventral wall of prostate: 1, ventral silt; 2, thin ventral wall; 3, thick ventral wall. 39. Vas deferens from pallial part of prostate: 1, emerges anteriorly; 2, emerges about half way ventrally. 40. Pallial vas deferens: 1, straight to slightly undulating; 2, moderately to strongly undulating. 41. Subterminal swelling on penis: 1, absent; 2, present. 42. Penial papilla (distal end): 1, no papilla; 2, short papilla; 3, long papilla. 43. Medial section of penis: 1, short and broad; 2, medium; 3, elongate and narrow. 44. Coiled oviduct: 1, firm, not embedded in connective tissue; 2, soft, in connective tissue. 45. Coiling of coiled oviduct: 1, simple U-shape; 2, coiled or with two or more folds or loops. 46. Posterior extent of coiled oviduct: 1, does not reach posterior edge of bursa; 2, extends to posterior edge of bursa; 3, extends behind bursa. 47. Point of joining of oviduct and bursal duct: 1, just behind pallial wall; 2, at posterior pallial wall; 3, just in front of posterior pallial wall; 4, well in front of posterior pallial wall. 48. Bursa copulatrix size: 1, small; 2, medium; 3, large. 49. Bursa copulatrix extent: 1, lies behind posterior pallial wall; 2, extends to posterior pallial wall; 3, extends in front of posterior pallial wall. 50. Bursa copulatrix shape: 1, globular; 2, obliquely ovoid; 3, vertically ovoid; 4, horizontally ovoid; 5, elongately oval; 6, horizontally pyriform; 7, vertically pyriform. 51. Position of bursal duct emerging from bursa: 1, anterior edge; 2, ventral edge. 52. Seminal receptacle-position: 1, at anterior edge of bursa on left; 2, on middle of inner wall of bursa on left; 3, behind posterior edge of bursa on left; 4, at mid ventral edge of bursa on left; 5, on right side of bursa. 53. Seminal receptacle shape: 1, ovoid to pyriform; 2, narrow sac. 54. Albumen gland: 1, or less in pallial roof; 2, about to in pallial roof; 3, to less than sin pallial roof; 3, more than s in pallial roof. 55. Length of capsule gland relative to albumen gland: 1, longer than albumen gland; 2, about equal to albumen gland; 3, up to about ? albumen gland; 4, less than s albumen gland. 56. Capsule gland shape in transverse section: 1, circular in section; 2, oval in section; 3, compressed-oval in section; 3, elongate oval in section. 57. Anterior end of capsule gland: 1, tapering; 2, blunt; 3, rounded. 58. Ventral channel: 1, simple; 2, expanded anteriorly. 59. Vestibule: 1, indistinct or small; 2, moderate-sized, muscular; 3, large, muscular. 60. Genital opening position relative to anterior end of capsule gland: 1, anterior; 2, overlapping; 3, terminal; 4, subterminal; 5, well behind anterior end.Pseudotricula is comprised of three subclades: P. progenitor is the sister to the remainder of the clade; P. conica and P. arthurclarkei form a separate clade from the remaining four Pseudotricula taxa, defined by a single unambiguous character (protoconch microsculpture). The third clade, which includes the typical species of Pseudotricula has bootstrap support and is defined by 10 unambiguous character changes (see Fig. 21), notably, a D-shaped aperture, an expanded outer lip, posterior apertural notch and a reduction in the number of cusps on the marginal teeth.

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PONDER ET AL.56 2005Magnolia Press 1074ZOOTAXA FIGURE 21. Strict consensus tree obtained from the 6 most parsimonious trees obtained from a cladistic analysis of the data in Table 13. The unambiguous character changes are listed on each branch (see Table 14 for list of characters). Bootstrap values of more than 50% are included next to the relevant branches on the left. Discussion This paper presents the first description of a significant radiation of closely-related freshwater molluscs occurring within a single karst area in Australia. The eight species of hydrobiids, in two closely related genera, in the Precipitous Bluff caves is a level of diversity that appears to be unique among Tasmanian (and Australian) karsts. This radiation mirrors the major radiation of hydrobiids in surface freshwaters in Tasmania and the southeastern mainland of Australia (Ponder et al 1993, 1994; Colgan & Ponder 1994; Clark et al 2003), albeit the Precipitous Bluff radiation involves fewer genera in a much smaller geographic area. Sampling in dozens of other Tasmania karst areas has produced about 30 (mostly undescribed) hydrobiid taxa, many of which are also likely to be stygobites. These include species of Austropyrgus Beddomeia Phrantela and Nanocochlea with no known additional species of Pseudotricula (Ponder et al 1993, WFP and SAC pers. observ.). In all other Tasmanian caves, one to five species are known from each cave system and in most cases where multiple species are present they are not congeners. Two genera and five species have been found in both Gunns Plains/Mastyn Hardy and Junee Florentine/

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2005 Magnolia Press 57HYDROBIIDAE S.L. 1074ZOOTAXA Caryodes Cavern (W.F.P. S.A.C. and J.S. pers. observ.). The richness of the Precipitous Bluff radiation was revealed through repeated sampling at multiple sites and habitats within this karst area, thus demonstrating the need for intensive sampling to properly reveal subterranean biodiversity. While many other Tasmanian karsts have been sampled for hydrobiids no other major cave radiations have been revealed to date, including in other karst areas such as Vanishing Falls, Cracroft, Hastings and Ida Bay in the same geographic region of southern Tasmania (Clarke 1990, 1997; Eberhard et al 1991a; Eberhard et al 1991b; WFP unpublished data). Ida Bay in particular has been intensively sampled (see Eberhard 2001) and like Precipitous Bluff, the Ida Bay karst area is highly cavernous and contains a diverse array of aquatic and terrestrial habitats which harbour a rich assemblage of troglobites including many of the same genera (such as Nanocochlea ) found at Precipitous Bluff (Eberhard et al 1991b, 2001). Ida Bay therefore, might be considered a likely candidate for a major radiation of cave dwelling hydrobiids. However, intensive and repeated sampling in this karst has revealed only two hydrobiids, one a Nanocochlea and the other a Phrantela (Eberhard 2001; WFP, SAC and JS unpublished data). Although not properly quantified, the diversity and heterogeneity in subsurface aquatic micro-habitats may be greater in the Precipitous Bluff caves than in other karst areas in Tasmania. The Precipitous Bluff karst is notable for the high density and development of caves, with more than 30 caves recorded and > 5 km of underground passages mapped within an area < 4 km2 (Eberhard & Hume 1987; Hume 1989; Hume & Clarke 1989). Most of the underground passages contain perennially active streams of various physical dimensions, flow rates, and substrate characteristics. Hence there is a diverse range of aquatic subsurface habitats present. Within most caves there is considerable gradation and small-scale (< 110 m) heterogeneity in substrate and flow characteristics, which change in response to changes in the physical dimensions and gradient of the cave conduits which are highly non-uniform. Thus the Precipitous Bluff caves provide considerable diversity and heterogeneity of aquatic habitats at a range of spatial scales, from in-stream micro-habitat, to in-cave stream(s), to in-karst cave(s). The aquatic subsurface habitats in the Precipitous Bluff caves range from tiny pools, seepages and trickles fed by diffuse vertical percolation, to small streams (< 0.5 m wide, < 0.1 m deep, flow < 1 L/s) formed from coalescing seepage flows, to larger streams (0.51 m wide, 0.11 m deep, flow 25 L/s), and finally major streams (24 m wide, 0.12 m deep, flow > 5 L/s). The benthic substrates of streams where the snails are found are also diverse. They include smooth limestone bedrock in sections of streams characterised by high flow velocities including chutes and waterfalls (e.g., Black Curtains Streamway in Cueva Blanca where P. eberhardi is found). Most other cave streams have a lower gradient than this particular site however, and their beds are cloaked in clastic sediments derived from higher up the slope of Precipitous Bluff. In streams with moderate-gradient and moderate to high-flow velocities (e.g., Bauhaus main streamway), the sediments typically

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PONDER ET AL.58 2005Magnolia Press 1074ZOOTAXA consist of cobbles and pebbles of dolerite, sometimes partially cemented within a matrix of gravel and silt. In low-gradient streams with lower flow velocities (e.g., Persephone Pot stream in Bauhaus, main streams in Damper Cave) the amount of gravel and silt increases proportionately, and these sediments may dominate the benthic substrate within some streams or caves (e.g., Quetzalcoatl Conduit). The clastic sediments on the beds of streams, which may be several metres deep in places, provide potential interstitial habitat which was not examined in detail. However, the single site sampled (Damper Cave) did yield hydrobiids. In general, the broader-shelled species appear to prefer faster-flowing water and their larger apertures may well be an adaptation to these environments. A large aperture is capable of housing a larger foot for clamping to the substrate than narrowshelled taxa, Based on the results of the cladistic analysis, the cave radiation represents two components an in situ radiation ( Pseudotricula ) derived from a common ancestor with Nanocochlea and a separate incursion ( N. exigua ). It is likely the radiation of Pseudotricula within the caves is the result of allopatric speciation with subsequent sympatry although there remains a possibility of sympatric speciation as a consequence of high habitat diversity and niche partitioning. Despite the high degree of hydrologic connectivity between the different caves and habitats within the caves, some species appear to have strict habitat preferences (Table 15). Most of the hydrobiid taxa prefer the small, gently flowing streams with a variety of substrate types, including silt. The most diverse assemblages (up to five sympatric species) occur in such streams as Persephone and the main streamway in Damper Cave. Two species appear to be specialised and restricted to the particular substrate and flow conditions in Cueva Blanca (Black Curtains Streamway: P. eberhardi ) and Quetzalcoatl Conduit ( P. arthurclarkei ) respectively, whilst other species are excluded. The hydrobiid diversity in the caves is higher than usually seen in surface habitats in Tasmania, the highest diversity seen being six species (three genera) in one location, with single species being found in most locations (Ponder et al 1993; Ponder & Colgan 2002; Clark et al 2003). While this probably contributes to the maintenance of biotic diversity it may not be the sole factor. Other factors that may contribute to hotspots of subterranean biodiversity in karstic regions include inter alia cave density, altitude, lateral extent and planar length of cave passage, and the degree of continuity or fragmentation between karst outcrops (see Holsinger & Culver 1988; Culver & Sket 2000; Culver et al 2004). These factors have not been rigorously tested for Tasmanian karsts although it appears to be generally the case that larger karst areas, which are highly cavernous tend to harbour greater diversity (Eberhard et al 1991b, Eberhard 1996). As elsewhere in the world, historical biogeographic and geomorphic factors are also likely to have strongly influenced the evolution of Tasmanian cave faunas (Eberhard 1996), particularly Quaternary climate oscillations, which included peri-glacial and glacial

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2005 Magnolia Press 59HYDROBIIDAE S.L. 1074ZOOTAXA conditions that profoundly influenced the thermal and hydrologic regimes within virtually all Tasmanian karsts (Kiernan 1982). These climatic changes have been implicated in the isolation and speciation of populations in Tasmanian caves (Moore 1964; Kiernan & Eberhard 1990; Eberhard 1992a, 1996), consistent with the Pleistocene effect theory, which is the most widely accepted model for the evolution of terrestrial troglobites in the temperate latitudes of the Northern Hemisphere (see reviews in Barr & Holsinger 1985; Holsinger 2000). This model was invoked to explain why there is a general paucity of troglobites in Northern Hemisphere karst areas strongly affected by glaciation during the Pleistocene. This contrasts with the relatively rich troglobite faunas found in karst areas that experienced peri-glacial conditions beyond glacial margins. Beside the extirpation of faunas in karsts directly overridden by ice, this model assumed that cool climates during glacial periods favoured wide distribution of terrestrial invertebrates inhabiting moist, forest-floor litter and cave habitats. During warm interglacial climates, the forests retreated leaving populations isolated in cave habitats and the reduced gene flow between hypogean and epigean populations promoted the evolution of troglobites. Presumably aquatic faunas would also be affected by climatic changes and certainly in Tasmanian surface waters hydrobiids were apparently eliminated from highland areas, which were glaciated during the Pleistocene, whilst their low dispersal powers appear to have restricted their recolonisation after deglaciation (Ponder et al 1993).TABLE 15. Summary of habitats and hydrobiid species in the caves at, and in the general vicinity of, Precipitous Bluff.Locationdamperensis exigua stylesae progenitorr arthurclarkei conica eberhardi expandolabra auriforma8 elongata PotamopyrgusOutside caves Malani CkX Damper Ck, first crossingXX Damper Ck, 10 m outside caveXX Damper Ck, at New R Lagoon X Limestone CkX Inside caves Bauhaus XXXXX main streamwayX Persephone streamwayXXXX Screaming Stals streamwayXXX Persephone Pot(X)X XXXX Cueva Blanca XX Damper Cave XXXXX main streamwayXXXXX main streamway near entrance XXXX cane toad abuseXXX seep near the kegXX honey and cream streamwayXX quetzalcoatl conduitXX

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PONDER ET AL.60 2005Magnolia Press 1074ZOOTAXA A few specimens of one of the minute cave species ( N. exigua ) has been found in a stream flowing from Damper Cave a few metres outside the cave. Most species of Nanocochlea are typically associated with seepages and small streams and live in crevices, under stones and rocks or in moss and roots. Their bodies are unpigmented and they lack pigmented eyes (see also below). In contrast, none of the larger snails ( Pseudotricula ) have been found outside the caves. Sampling suggests that streams outside the Precipitous Bluff caves, even close to a cave entrance (Damper Cave), have a different fauna with the one exception noted above. Thus, based on their absence from collections made in surface waters at Precipitous Bluff, we consider the cave-dwelling species stygobites, despite the retention of unpigmented eyes. The loss of body and eye pigment appears to have occurred in at least some of the apparently ancestral Nanocochlea although N. damperensis has lightly-pigmented eyes. Unpigmented eyes are also found in some other stygobitic hydrobiids and are lost in others (e.g., Climo 1974). Human-induced changes to water quality, flow regimes and benthic habitats can adversely impact cave dwelling hydrobiids such as resulted from quarrying operations at Ida Bay (e.g., Eberhard 1992b, 2001). Fortunately, the isolation of Precipitous Bluff and its containment within a National Park suggests that there are few obvious conservation concerns for this fauna. However, the presence of the introduced invasive Potamopyrgus antipodarum (Gray, 1843) at the lower end of Damper Creek (stn PBs-3) is of potential concern to the surface stream-living species. This parthenogenic, highly fecund New Zealand species appears to compete with native taxa to their detriment (e.g., Ponder 1988; Schreiber et al 2002). Extensive tracts of limestone remain partly explored in the Precipitous BluffVanishing Falls-Cracroft areas and it is probable that other cave systems exist which may contain additional hydrobiid species. Acknowledgements This work was supported in part by ARC and ABRS grants. A small grant was also obtained from the Tasmanian Parks and Wildlife Service. Arthur Clarke, Janet Waterhouse, Vera Wong and Julie Styles assisted with collecting material. The map used in Fig. 1 is based on one drawn by N. Hume in 1989 for the TCC and is used with permission. Peter Middelfart assisted with some of the SEM work. Arthur Clarke assisted with the locality data and he and Martin Haase made useful comments on the manuscript. References Barr, T. & Holsinger J.R. (1985) Speciation in cave faunas. Annual Review of Ecology and Systematics 16, 3137.

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2005 Magnolia Press 61HYDROBIIDAE S.L. 1074ZOOTAXA Clark, S.A., Miller, A.C. & Ponder, W.F. (2003) Revision of the snail genus Austropyrgus (Gastropoda: Hydrobiidae). A morphostatic radiation of freshwater gastropods in southeastern Australia. Records of the Australian Museum Supplement 28, 1109. Clarke, A. (1990) A summary of recent karst research in Tasmania. Tasmanian Cave and Karst Research Group Journal, 4, 1. Clarke, A. (1997) Management prescriptions for Tasmania's cave fauna. Report to the Tasmanian Regional Forest Agreement Environment and Heritage Technical Committee. March 1997. 166 pp. Climo, F. M. (1974) Description and affinities of the subterranean molluscan fauna of New Zealand. New Zealand Journal of Zoology, 1, 24784. Colgan, D.J. & Ponder, W.F. (1994) The evolutionary consequences of restrictions of gene flow: examples from hydrobiid snails. Nautilus Supplement 2, 253. Culver, D.C. & Holsinger, J.R. (1992) How many species of troglobite are there? National Speleological Society Bulletin 54, 7980. Culver, D.C. & Sket, B. (2000) Hotspots of subterranean biodiversity in caves and wells. Journal of Cave and Karst Studies 6, 11. Culver, D.C., Christman, M.C., Sereg, I., Trontelj, P. & Sket, B. (2004) The location of terrestrial species-rich caves in a cave-rich area. Subterranean Biology 2, 2732. Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1993) Users guide to the DELTA System: a general system for processing taxonomic descriptions. 4th edition. http://biodiversity.uno.edu/delta/ Dixon, G., & Sharples, C. (1986) Reconnaissance geological observations on Precambrian and Palaeozoic rocks of the New and Salisbury Rivers, Southern Tasmania. Papers and Proceedings of the Royal Society of Tasmania 120, 874. Eberhard, S. (1992a) The invertebrate cave fauna of Tasmania: ecology and conservation biology MSc thesis, Zoology Department. University of Tasmania. 184 pp. Eberhard, S. (1992b) The effect of stream sedimentation on population densities of hydrobiid molluscs in caves. Report to Tasmanian Dept. Parks, Wildlife and Heritage, Hobart. 8 pp. Eberhard, S. (1996) Tasmanian cave fauna In : Juberthie, C. & Decu, V. (eds) Encyclopedia Biospeologica Socit Internationale de Biospeleologie, Moulis Bucarest, pp. 2093103. Eberhard, S. (2001) Cave fauna monitoring and management at Ida Bay, Tasmania. Records of the Western Australian Museum Supplement 64, 97104. Eberhard, R., Eberhard, S. & Wong, V. (1991a) Karst geomorphology and biospeleology at Vanishing Falls, south-west Tasmania. Helictite 30, 2532. Eberhard, S. & Hume, N. (1987) Report of the Tasmanian Caverneering Club 1986 speleological reconnaissance expedition to Precipitous Bluff. Speleo Spiel (Newsletter of the Tasmanian Caverneering Club) 246, 15. Eberhard, S., Richardson, A.M. & Swain, R. (1991b) The invertebrate cave fauna of Tasmania Report to the National Estate Office, Canberra. Zoology Department, University of Tasmania. 174 pp. Gentilli, J. (1972) Australian climate patterns Thomas Nelson and Sons, Melbourne. 285 pp. Giusti, F. & Pezzoli, E. (1977) Primo contributo alla revisione del genere Bythinella in Italia. "Natura Bresciana" Annuario del Museo Civico di Storia Naturale di Brescia, 14, 366. Giusti, F. & Pezzoli, E. (1981) Notulae malacologicae XXV. Hydrobioidea nuove o poco conosciute dell'Italia appenninica. Archiv fr Molluskenkunde 111, 2072. Hershler, R. & Longley, G. (1986) Phreatic hydrobiids (Gastropoda: Prosobranchia) from the Edwards (Balcones Fault Zone) aquifer region, south-central Texas. Malacologia 27, 127 172. Holsinger, J.R. (2000) Ecological derivation, colonization, and speciation. In : Wilkens, H., Culver, D.C. & Humphreys, W.F. (eds) Ecosystems of the World Vol. 30 Subterranean Ecosystems Elsevier, Amsterdam, pp. 39915.

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PONDER ET AL.62 2005Magnolia Press 1074ZOOTAXA Holsinger, J.R. & Culver, D.C. (1988) The invertebrate cave fauna of Virginia and a part of Eastern Tennessee: zoogeography and ecology. Brimleyana 14, 1164. Hughes, T.D. (1957) Limestones in Tasmania. Geological Survey and Mineral Resources 10. Tasmanian Department of Mines. Hobart. 291 pp. Hume, N. (1989) 1988/89 Precipitous Bluff Expedition part 2. Speleo Spiel (Newsletter of the Tasmanian Caverneering Club) 253, 114. Hume, N. & Clarke, A. (1989) 1988/89 Precipitous Bluff Expedition part 1. Speleo Spiel (Newsletter of the Tasmanian Caverneering Club) 252: 1. Hunt, G.S. (1990) Hickmanoxyomma, a new genus of cavernicolous harvestmen from Tasmania (Opiliones: Triaenonychidae). Records of the Australian Museum 42, 458. Hunt, G.S. & Hickman, J.L. (1993) Revision of the genus Lomanella Pocock with implications for family level classification in the Travunoidea (Arachnida: Opiliones: Triaeononychidae). Records of the Australian Museum 45, 8119. Kiernan, K.W. (1982) Glaciation and karst in Tasmania: review and speculations. Helictite 20, 11 16. Kiernan, K.W. (1995) An atlas of Tasmanian karst. Tasmanian Forest Research Council Inc., Research Report No. 10. 351 pp. Kiernan, K.W. & Eberhard, S. (1990) Karst resources and cave biology. In : Smith, S.J. & Banks, M.R. (eds) Tasmanian wilderness world heritage values Royal Society of Tasmania, Hobart, pp 287. Moore, B.P. (1964) Present day cave beetle fauna in perspective: a pointer to past climatic change. Helictite 3, 3. Moore, B.P. (1978) A new species of the Tasmanian cave carabid genus Idacarabus (Coleoptera). Australian Entomology Magazine 5, 23. Perez, K.E., Ponder, W.F., Colgan, D.J., Clark, S.A. & Lydeard, C. (2005) Molecular phylogeny and biogeography of spring-associated hydrobiid snails of the Great Artesian Basin, Australia. Molecular Phylogenetics and Evolution 34, 545556. Platnick, N.I. & Forster, R.R. (1989) A revision of the temperate South American and Australian spiders of the family Anapidae. Bulletin American Museum Natural History 190, 139. Ponder, W.F. (1988) Potamopyrgus antipodarum a molluscan coloniser of Europe and Australia. Journal of Molluscan Studies 54, 27185. Ponder, W.F. (1992) A new genus and species of aquatic cave-living snail from Tasmania (Mollusca: Gastropoda: Hydrobiidae). Proceedings of the Royal Society of Tasmania 126, 23. Ponder, W.F., Clark, G.A., Miller, A. & Toluzzi, A. (1993) On a major radiation of freshwater snails in Tasmania and eastern Victoria a preliminary overview of the Beddomeia group (Mollusca: Gastropoda: Hydrobiidae). Invertebrate Taxonomy 7, 501750. Ponder, W.F. & Colgan, D.J. (2002) What makes a narrow range taxon? Insights from Australian freshwater snails. Invertebrate Systematics 16, 57182. Ponder, W.F., Colgan, D.J. & Clark, G.A. (1991) The morphology, taxonomy and genetic structure of Tatea (Mollusca: Gastropoda: Hydrobiidae), estuarine snails from temperate Australia. Australian Journal of Zoology 39, 44797. Ponder, W.F., Colgan, D.J., Clark, G.A., Miller A.C. & Terzis, T. (1994) Microgeographic genetic and morphological differentiation of freshwater snailsthe Hydrobiidae of Wilsons Promontory, Victoria, south eastern Australia. Australian Journal of Zoology 42, 557678. Ponder, W. F., Hershler, R. & Jenkins, B. (1989) An endemic radiation of Hydrobiidae from artesian springs in northern South Australia: their taxonomy, physiology, distribution and anatomy. Malacologia 31, 10. Radoman, P. (1983) Hydrobioidea, a superfamily of Prosobranchia (Gastropoda). I. Serbian Academy of Sciences and Arts, Monograph DXLVII 16. Reid, J.B., Hill, R.S., Brown, M.J. & Hovenden, M.J. (eds) (1999) Vegetation of Tasmania. Flora of

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2005 Magnolia Press 63HYDROBIIDAE S.L. 1074ZOOTAXA Australia Supplementary Series 8. Australian Biological Resources Study, Canberra. ixx, 1 453 p. Schreiber, E.S.G., Lake, P.S. & Quinn, G. (2002) Facilitation of native stream fauna by an invading species? Experimental investigations of the interaction of the snail, Potamopyrgus antipodarum (Hydrobiidae) with native benthic fauna. Biological Invasions 4, 317325. Swofford, D.L. (1998) PAUP*. Phylogenetic analysis using parsimony (* and other methods). Version 4. Sinauer Associates: Sunderland, Massachusetts, USA. Wilke, T., Davis, G.M, Falniowski, A., Giusti, F., Bodon, M. & Szarowska, M. (2001) Molecular systematics of Hydrobiidae (Mollusca: Gastropoda: Rissooidea): testing monophyly and phylogenetic relationships. Proceedings of the Academy of Natural Sciences of Philadelphia 151, 121. Wilkens, H., Culver, D.C. & Humphreys, W.F. (eds.) (2000) Subterranean Ecosystems Ecosystems of the World, 30. Elsevier, Amsterdam. xiv + 791 pp.

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PONDER ET AL.64 2005Magnolia Press 1074ZOOTAXA Appendix: Station dataA. Outside caves Malani Ck, Precipitous Bluff, in rocky seep/stream in mixed forest, stn 14, 43 28S, 146 35E S. Eberhard & V. Wong 28 DEC 1991 (Note: Malani Ck drains the slopes of PB & is the next major creek to the north of the known caves). Damper Ck tributary, first crossing, stn PBs-1A, 43 29.25S, 146 34.5E, S. Eberhard & J. Styles, 29 MAR 1994 (Stream width c. 2 m, depth < 0.2 m, flow est. 1020 L/s, substrate cobbles-gravel, microhabitat gentle riffle). Damper Ck, 10m outside Damper Cave, green filamentous algae, stn PBs-2a, 43 29.06S, 146 34.96E, S. Eberhard & J. Styles, 29 MAR 1994 (Stream width c. 2 m, depth < 0.5 m, flow est. 10 20 L/s, substrate green filamentous algae, stream bed of silt-gravel, microhabitat gentle riffle/pool (= run)) New River Lagoon, at Damper Ck, stn PBs-3, 43 29.20S, 136 34.8E, S. Eberhard, 30 MAR 1994 (Stream width c. 4 m, depth < 1 m, flow est. 100 L/s, stream bed of silt-gravel with logs, litter; Potamopyrgus antipodarum (AMS C.202667). Limestone Ck, near New River Lagoon, where tagged track crosses creek, JW 49, 43 29.2S, 136 34.8E, J. Waterhouse, 15 FEB 1988. Shallow, flowing stream with gravel, sand and weeds. Bauhaus, cave entrance, 43 29.00S, 146 37.00E, S. Eberhard, 26 MAR 1994, litter associated with small seepages. B. Inside caves Bauhaus (PB6) cave entrance 43 29.01S, 146 35.28E. Bauhaus, in gravel, stn PB6-1, S.E. Eberhard & J. Styles, 27 MAR 1994 (Stream width < 1 m, depth < 0.3 m, flow est. < 5 L/s.) Bauhaus, base of aven near shaft entrance surface wash, stn PB6-1, S.E. Eberhard & J. Styles, 26 MAR 1994 (seepage close to surface, rocky substrate). Bauhaus, Persephone streamway, stn 8, S. Eberhard & V. Wong, 23 DEC 1991 Bauhaus, Persephone streamway, further upstream, near dead possum, stn 9, S. Eberhard & V. Wong, 23 DEC 1991 Bauhaus, Persephone streamway, stn 9, S. Eberhard & V. Wong, 23 DEC 1991 Bauhaus, Persephone streamway, stn 10, S. Eberhard & V. Wong, 23 DEC 1991 Bauhaus, Persephone streamway, near end of passage, stn 10, S. Eberhard & V. Wong, 23 DEC 1991. Collections for all these made progressively upstream from point of first entry into passage. Habitat details generally as: Stream width, 0.5 m, depth < 0.2 m, flow est. < 1 l/s. Substrate clastic sediments cobbles, gravel, silt, microhabitats gentle riffles & pools

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2005 Magnolia Press 65HYDROBIIDAE S.L. 1074ZOOTAXA Bauhaus, Screaming Stals streamway, stn 11, S. Eberhard & V. Wong, stn 23 DEC 1991 Bauhaus, Screaming Stals streamway, stn 12, S. Eberhard & V. Wong, 23 DEC 1991 Bauhaus, Screaming Stals streamway, tributary seep, stn 12, S. Eberhard & V. Wong, 21 DEC 1991 Bauhaus, Screaming Stals streamway, stn 13, S. Eberhard & V. Wong, 23 DEC 1991 Collections made progressively upstream from point of first entry into passage. Habitat details generally as: Stream width < 1 m, depth < 0.3 m, flow est. < 5 L/s. Substrate clastic sediments cobbles, gravel, silt, microhabitats gentle riffles & pools Bauhaus, Persephone Pot, at first contact with stream, stn 7, S. Eberhard & V. Wong, 23 DEC 1991 Bauhaus, Persephone Pot, in stream, stn 7, S. Eberhard & V. Wong, 23 DEC 1991 Bauhaus, Persephone Pot, in stream, stn 9, S. Eberhard & V. Wong, 23 DEC 1991 Collections made progressively upstream from point of first entry into passage. Habitat details generally as: Stream width, 0.5 m, depth < 0.2 m, flow est. < 1 l/s. Substrate clastic sediments cobbles, gravel, silt, gentle riffles and pools. Bauhaus, Persephone Pot, stn PB17-8R, slow flowing stream with silty substrate in deep zone, S. Eberhard, 3 JAN 1990. Bauhaus, Persephone Pot, middle & upper streamway, in gravel, stn PB17-2, S. Eberhard & J. Styles, 27 MAR 1994 Bauhaus, Persephone Pot, lower streamway, stn PB17-2a.1, S. Eberhard, 26 MAR 1994 Bauhaus, Persephone Pot, middle and upper streamway, stn PB17-2a.2, S. Eberhard, 27 MAR 1994 Bauhaus, Persephone Pot, lower streamway, in gravel, stn PB17-1, S.E. Eberhard & J. Styles, 26 MAR 1994 (For all these: stream width 0.5 m, depth < 0.2 m, flow est. < 1 l/s, substrate clastic sediments cobbles, gravel, silt, microhabitats gentle riffles & pools) Bauhaus, main streamway at downclimb, stn PB6-1A, S. Eberhard, 27 MAR 1994 (Stream width < 1 m, depth < 0.3 m, flow est. < 5 L/s, substrate cobbles, gravel, microhabitat riffle-pool). Cueva Blanca cave (PB4) entrance 43 29.00S, 146 34.95E. Cueva Blanca, clinging to smooth rock in turbulent shallow stream in dark zone, stn PB4, S. Eberhard, 31 MAR 1986 Cueva Blanca, A. Clarke & S. Eberhard, 18 DEC 1988. Cueva Blanca, Black Curtain Streamway (= Inundation Passage), in torrents & waterfalls, 100200m from entrance, stn PB4, A. Clarke & S. Eberhard, 18 DEC 1988. Cueva Blanca, Black Curtain Streamway, stn PB4-1, J. Styles, 30 MAR 1994. Cueva Blanca, Black Curtain Streamway, stn PB4-1C, S. Eberhard, 30 MAR 1994. (For all these: stream width ca. 1 m, depth highly variable chutes < 0.1 m to pools 1.5 m, Substrate smooth limestone bedrock. Flow high gradient streamway consisting of chutes and small waterfalls interspersed with plunge pools; flow ca. < 5 L/s. Damper Cave (PB1) entrance 43 29.06S, 146 34.96E. Damper Cave, main streamway, 100 m into cave, in dark zone, stn 6, S. Eberhard & V. Wong,

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PONDER ET AL.66 2005Magnolia Press 1074ZOOTAXA 2223 DEC 1991 (Stream width c. 2 m, depth < 0.2 m, flow est. 10 to 20 L/s, substrate cobblesgravel, microhabitat gentle riffle). Damper Cave, Cane Toad Abuse, stn PB1-2, S. Eberhard & J. Styles, 28 MAR 1994. Damper Cave, Cane Toad Abuse, streamway, stn 2, A. Clarke, 22 DEC 1988. Damper Cave, Cane Toad Abuse, streamway, stn PB1-2A, S. Eberhard & J. Styles, 28 MAR 1994. (All above: stream width ca. 1 m, depth, 0.1 m, flow est. < 1 L/s, substrate cemented cobbles, with gravel-silt. microhabitat gentle riffle). Damper Cave, seep near The Keg, stn 3, S. Eberhard & V. Wong, 22 DEC 1991 (Small seepage fed by drip water from aven near The Keg. Located several hundred metres into cave, deep zone). Damper Cave, main streamway near entrance (ca. < 50 m inside cave), in gravel, stn PB1-1 and PB1A (also listed as PB1-1A) S.E. Eberhard & J. Styles, 28 MAR 1994 (Stream width c. 2 m, depth < 0.2 m, flow est. 10 to 20 L/s, substrate cobbles-gravel, microhabitat gentle riffle). Damper Cave, main streamway near entrance (ca. < 50 m inside cave), interstitial gravel, stn PB11B, S. Eberhard & J. Styles, 28 MAR 1994. (Stream width c. 2 m, depth < 0.2 m, flow est. 10 to 20 L/s, Substrate cobbles-gravel, microhabitat (interstitial gravel sampled using Karaman-Chapuis method)). Damper Cave, streamway, Honey and Cream, stn 4, S. Eberhard & V. Wong, 22 DEC 1991 (Small tributary stream in Damper Cave. Stream width < 1 m, depth < 0.1 m, flow est. < 1 L/s, substrate pebbles-gravel-silt. Microhabitat small stream, riffle). Quetzalcoatl Conduit (PB3) entrance 43 29.20S, 146 35.00E. Quetzalcoatl Conduit, stn PB2-1C, S. Eberhard & J. Styles, 29 MAR 1994. Quetzalcoatl Conduit, 30m into cave, stn PB3-1, S. Eberhard & J. Styles, 29 MAR 1994. (Both: stream width 34 m, depth ranging from riffle/runs < 0.1 m to pools > 2 m, substrate limestone bedrock, sediments cobbles, gravel, flow est. 1530 L/s, but cave floods severely. Collections made in shallow-riffle zones with hard (rocky) substrate). Quetzalcoatl Conduit, 30m into cave, stn PB3-1C, S. Eberhard & J. Styles, 29 MAR 1994 (Stream width 34 m, depth ranging from riffle/runs < 0.1 m to pools 1.5 m, substrate limestone bedrock, cemented cobbles, in shallow-riffle zones with hard (rocky) substrate. Flow est. 15 30 L/s, but cave floods severely). Quetzalcoatl Conduit, beyond Pendulum Palace, stn PB3-2, S. Eberhard & J. Styles, 29 MAR 1994 (Stream width 34 m, depth ranging from runs < 0.5 to pools > 2 m, substrate mostly soft sediments, gravel-silt, flow est. 1530 L/s, but cave floods severely). Quetzalcoatl Conduit, stn PB3-2B, S. Eberhard & J. Styles, 29 MAR 1994 (Stream width 34 m, depth ranging from runs < 0.5 to pools > 2 m, substrate mostly soft sediments, gravel-silt, flow est. 1530 L/s, but cave floods severely).