Memoirs of THE Queensland Museum Brisbane 10 January 2005 Volume 50 Part 2 ■5 Volume 50 Part 2 Memoirs of THE Queensland Museum Minister: Director: Managing Editor: Editorial Assistant: Hon. Anna Bligh MLA LG. Galloway, PhD P.A. Jell, PhD P. Avem, BSc PUBLISHED BY ORDER OF THE BOARD 10 JANUARY 2005 © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 61 7 3840 7555 Fax 61 7 3846 1226 National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Memoirs of the Queensland Museum may b reproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations ma be made but queries regarding the republication of any papers should be addressed to the Director. Copies ol the journal ca be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site www.qmuseum.qld.gov.au/resources/resourcewelcome.html A Queensland Government Project Typeset at the Queensland Museum Printed by Watson Ferguson & Co 35 Hamilton Road, Moorooka, Queensland 4105 A NEW SPECIES OF LERISTA (SCINCIDAE) FROM CENTRAL QUEENSLAND ANDREW P. AMEY, ALEX S. KUTT AND MARK HUTCHINSON Amey, A.P., Kutt, A.S. & Hutchinson, M. 2005 01 10: A new species of Lerista (Scincidae) from central Queensland. Memoirs of the Queensland Museum 50(2): 125-131. Brisbane. ISSN 0079-8835. Lerista chordae sp. nov. from the Desert Uplands Bioregion of central Queensland is a small skink with tetradactyl limbs and a moveable eyelid. It is readily distinguished from its congeners by the combination of fore-limb only 35-52% of hind-limb length; 2 phalanges and 3 subdigital lamellae on the 4th finger; ear aperture similar size or smaller than nostril; normally 20 midbody scale rows; four lines of dark flecks on dorsum; an ill-defined dark dorsolateral zone encompassing two half-scale widths only; an immaculate white vent and tail pale yellow in spirit. The new species occurs in open Eucalyptus woodlands characterized by very sandy soils and a sparse to dense ground cover of spinifex or other tussock grasses. The morphologically similar species Lerista quadrivincula , known from a single specimen, is redescribcd. □ Lerista, skink, new species, desert uplands , central Queensland. Andrew P. Amey, Queensland Museum, PO Box 3300, South Brisbane 4101; Alex S. Kutt, Environmental Protection Agency, PO Box 5391, Townsville 4810; Mark Hutchinson, South Australian Museum, North Terrace, Adelaide 5000, Australia; 3 September 2004. The Desert Uplands Bioregion of central Queensland is an area of Acacia and Eucalyptus woodlands, dune systems and grasslands bordering the Einasleigh Uplands, Mitchell Grass Downs and the Northern Brigalow Belt between Charters Towers, Hughenden and Blackall. It has been little surveyed in com- parison with nearby coastal eastern Queensland. Its biodiversity potential has probably been under-appreciated. Recent surveys have resulted in the discovery of a new species of Ctenotus (Couper et al., 2002) and a new species of Lerista , described herein. This suggests that the inhabitants of the diverse habitats of this bioregion especially the smaller, cryptic species, may be worth closer scrutiny, particularly as pressure for more intensive development for pastoral activity increases. Lerista is the second largest reptile genus in Australia (after Ctenotus) with 79 species recognised at the end of 2003. While many species are common and widespread, many others are known from only a few specimens from few or remote localities; L. bunglebungle Storr, 1991, L. praefrontalis Greer, 1986, L. quadrivincula Shea, 1991 and L . speciosa Storr, 1990 are recorded solely from the holotype. Knowledge of intra- and inter-specific variation is therefore very patchy within this genus. As has been pointed out elsewhere (Greer, 1987, 1989, 1990b), the remarkable range of limb reduction within the genus, from the full 5 digit condition to limbs entirely absent, even varying within a species, offers rich possibilities for study of the evolutionary processes driving this condition. The dire situation of Lerista allanae , possibly Australia’s first reptile species to be driven to extinction by human activities (Covacevich et al., 1996), exemplifies the need for sound taxonomic assessment of cryptic taxa as an aid in determining conservation priorities. METHODS Part of the type series, the first specimens to be identified, was collected during broadscale surveys conducted within the Desert Uplands to assess the patterns of vertebrate assemblage and distribution (Kutt, 2003). Sampling methods are described in Couper et al. (2002). The rest of the type series, including the holotype, was collected by hand opportunistically along the Torrens Creek-Aramac Rd. These specimens were hiding in loose bark or small partly buried branches, in sandy patches in a generally clay-soil terrain. All body measurements were taken using Mitutoyo electronic calipers. Scales were counted on the right side only of specimens examined. The total number of enlarged nuchals is given. Presacral vertebrae, caudal vertebrae and phalanges were counted on X-ray photographs taken with a Torrex 120D X-ray inspection system using 50k V, 50mA, 50s exposure. Only original tails were included in the morphometric analysis (assessed by X-ray). Abbreviations for body measurements are as follows: snout-vent 126 MEMOIRS OF THE QUEENSLAND MUSEUM length (SVL); axilla to groin (AG); original tail length (vent to tip, TL); fore-limb (axilla to tip of longest finger, LI); hind-limb (groin to tip of longest toe, L2); snout-axilla (SA); head width (widest point, HW); head length (tip of snout to posterior margin of parietals, HL); eye to ear (posterior margin of orbit to dorsal anterior margin of ear, EE). Scalation definitions follow Horner (1992). Other abbreviations used: Queensland Museum (QM), South Australian Museum (SAM), standard deviation (SD). For comparison, all species of the Lerista frosti group (L. dorsalis , L. flammicauda , L. frosti , L. quadrivincula and L. zietzi) were examined using museum specimens. SYSTEMATICS The new species is assigned to Lerista Bell, 1833, following the revised diagnosis of Greer (1986). It conforms to the frosti group (Storr et al., 1999), with moveable eyelids, a 4 + 4 digital formula, 3 supraoculars, 5 supraciliaries, 6 upper labials and midbody scales in 1 8-22 rows. These plesiomorphic characters are widespread in Lerista (Greer, 1986, 1990a) and so membership of the group does not necessarily imply close phylogenetic relationship between group members. Lerista chordae sp. nov. (Figs 1,2) ETYMOLOGY. Latin chordae , strings of a lyre, in reference to the thin black lines on the dorsum. MATERIAL. HOLOTYPE QMJ81070 Torrens Creek- Aramac Rd. 40km S Torrens Creek (21°05'30”S 145°00’16”E). PARATYPES QMJ72754-5 Bede Sin, 100km NNE of Aramac (22°22’37"S. 145 a 35*32 , 'E), QMJ74034 Ulva Stn, 100km S of Torrens Creek (21°25’00”S, 1 45°08*45 ,, E), QM J8 1 07 1 22km (road) S of Torrens Creek (20°59 , 09 1, S 1 45°0 1 , 54'*E), SAMR55681-4 Torrens Creek-Aramac Rd. 40km S Torrens Creek (21°05'30 ,, S 145°00 , 16”E). See Fig. 3 for map of collection localities. DIAGNOSIS. Readily distinguished from all other Lerista by the combination of smaller size (max. SVL 45.10mm), lower eyelid moveable; ear aperture similar size to nostril; four digits on fore- and hindlimbs, forelimb 35-52% of hindlimb length; 2 phalanges and 3 subdigital lamellae on 4th finger; normally 20 midbody scale rows; 36-38 presacral vertebrae; four lines of dark flecks on dorsum; an ill defined dark dorsolateral zone encompassing two half-scale widths only; an immaculate white vent and tail pale yellow in spirit. DESCRIPTION. Measurements. SVL (mm) 29.28-45.10 (mean=38.27, SD=5.59, n=9). Proportions (% SVL): AG=6 1 .82-69.55 (mean=65.79, SD=2.34, n=9); TL=1 12.78- 130.08 (mean=l 19.62, SD=9.20, n=3); LI =9.76- 13.63 (mean= 11.71, SD=1.24, n = 8); L2= 18.76-32.54 (mean=25.82, SD=4.14, n=9); HL=12. 17-14.46 (mean=13.15, SD=0.98, n=9), SA=25.68-31.19 (mean=28.56, SD=2.00, n=9). Proportions (% L2): L 1 =35.62-52.03 (mean=45.48, SD=5.07, n=8). Proportions (% HL): HW=52.42-64.62 (mean=57.1 1, SD=3.89, n=9); EE=38.9 1-49.69 (mean=41.80, SD=3.42, n=9). Scalation. Rostral crescent-shaped with triangular medial projection between nasals; nasals separated (n = 7) or in contact (n=2), nostril placed anteriorly and laterally; frontonasal wider than long, saddle-shaped, triangular anteriorly between nasals to meet or just fail to meet rostral, concave posteriorly; frontal longer than wide, somewhat coffin- shaped, contacts frontonasal, prefrontals, first 2 supraoculars and frontoparietals; 3 supraoculars, 2nd the largest; supraciliaries 5, rarely 4 (QMJ81071 one side only), 1st, 3rd and 4th project between prefrontal and 1st supraocular, 1 st and 2nd supraocular, 2nd and 3rd supraocular respectively; palpebrals 5 rarely 6 (QMJ72755); frontoparietals paired and distinct; interparietal distinct, angular anteriorly, rounded posteriorly; parietals in broad contact behind interparietal; 2-6 enlarged nuchals; loreals 2, 1st largest; 2 preoculars, I prcsubocular; 2 postoculars, 1 postsubocular; primary temporal in contact with pretemporal, secondary temporal, 5th and 6th supralabials; secondary temporal in broad contact with parietal, primary temporal and point contact with pretemporal; supralabials 6, 4th subocular; infralabials 6, 2 contacting postmental; 2 additional rows of enlarged chin shields; ear opening circular to vertically elliptic, similar size to nostril; midbody scale rows 20, rarely 18 (QMJ81071, SAMR55681); para- vertebrals 68-75 (mean=72.33, SD=3.08, n=9); lamellae beneath 4th finger 3 (n=8); supradigitals above 4th finger 2 (n=8); lamellae beneath 4th toe 5, rarely 4 (SAMR55682, QMJ72754 one side only) or 6 (SAMR55682, R55684 one side only); supradigitals above 4th toe 4 rarely 3 (QMJ72755, J81071, SAMR55681 one side only) or 5 (SAMR55684 both sides); lamellae beneath longest toe (3rd) 13-16 (mean 1 4.44, SD =1.01, n=9); supradigitals above longest toe A NEW SPECIES OF LERISTA 127 FIG. 1. One of the paratypes of Lerista chordae in life, Torrens Creek-Aramac Rd (21°05 , 30 ,, S 145^00’ 16”E). (3rd) 9-12 (mean=10.11, SD=0.93, n-9); subcaudals 78-8 1 (mean=79.33, SD= 1 .53, n=3). Osteology’. Presacral vertebrae 36-38 (mean= 37.33, SD=0.87, n=9) and caudal vertebrae 42 (n =3). Phalangeal formula of fore-limb 0.2.3 .4.2 (n =4), hind-limb 0.2. 3. 5. 3 (n=7). Holotype. Measurements and scale counts for the holotype are as follows: Sex: male (determined by dissection); SVL =42. 79mm; AG=28.50mm; TL=55.66mm; LI =4. 84mm; L2=9.84mm; HL=5.37mm; HW=3.00mm; EE=2.13mm; SA= 12.08mm; nasals separated, enlarged nuchals 6, supraciliaries 5, palpebrals 5, ear opening circular, midbody scale rows 20, paravertebrals 75, lamellae beneath 4th finger 3; supradigitals above 4th finger 3; lamellae beneath 4th toe 5, supradigitals above 4th toe 4, lamellae beneath longest toe (3rd) 15, supradigitals above longest toe (3rd) 12, subcaudals 78. Presacral vertebrae 37, caudal vertebrae 42. Colour Pattern (In Spirit). Dorsal ground colour grey-brown to bronze. Four longitudinal lines of fine dark chocolate brown spots running from nape to base of tail, breaking up into irregular flecking along tail, outer pair sometimes discontinuous (SAMR55681). Chocolate brown dorsolateral band from rostral to base of tail, 2 half-scale widths on flanks, ill-defined, continuing along tail where it breaks up into irregular flecking and tends to merge with dorsal flecks. Head with dark, fine flecks above. A dark vertical medial bar through rostral scale. Pale limbs mottled with dark brown above. Tail very pale yellow in spirit. Ventral surfaces immaculate white, very pale yellow on tail. Supralabials lightly to strongly edged with brown/black, sometimes infralabials also (QMJ72754-5). Lateral scales edged with dark brown/black merging with dark dorsolateral band. Regrown tail may be immaculate pale yellow (SAMR55684). In life, juveniles have bright red tail colouring (brightest on the ventral surface), the colour in adults being pale orange-yellow. Variation in Paratypes . Quantitative variation in scale characters and osteology among paratypes is given above. Paratypes varied in the degree of dark markings, from 2 faint rows of spots on the dorsum, light flecking on the top of the head and no labial edging (SAMR5568 1 ) to 4 broken lines dorsally (QMJ81071), heavy flecking on the top of the head (QMJ74034) and heavy edging on the supralabials (QMJ72754 and J75755). Sex was known for only 3 individuals (QMJ81070, SAMR55682 and R55683, examination through incision made for tissue sampling), which were all males. These exhibited wide variability in characters normally associated with sexual dimorphism in Lerista. For example, the number of presacral vertebrae is commonly higher in females than males (Greer, 1 987, 1 990b) while in the 3 known males, these ranged between 36 and 128 MEMOIRS OF THE QUEENSLAND MUSEUM 38, the total range for the species. Consequently, it was thought unnecessarily destructive to determine sex of the other individuals, and therefore any dimorphic characters. Comparison with Other Species. The combination of tetradactyl fore- and hind-limbs and a movable eyelid distinguishes this species from all currently described Lerista except L. dorsalis , L. flammicauda , L. frosti , L. quadrivincula and L. zietzi. Nasals normally widely separated, nostril similar size to ear aperture, short fore-limbs (35-52% of hind-limb) with digital formula 0.2. 3.4. 2, 3 subdigital lamellae on the 4th finger, • dorsum with 4 longitudinal lines of Hecks, and narrow, ill-defined dorsolateral band separate this species from L. flammicauda and L. zietzi (nasals usually in contact, car aperture larger than nostril, fore-limbs with digital formula 0.2. 3.4.3, 52-77% hind-limb with 5-7 lamellae on 4th finger or 63-77% hind-limb with 6-7 lamellae on 4th finger respectively, dorsum either immaculate or 2-4 narrow, discontinuous lines of flecks on nape or rump, dorsolateral band 2 scales wide, solid and sharp-edged or narrow but solid and well-defined dorsally). It is distinguished from L. dorsalis and L. frosti by an irregular dark dorsolateral stripe (vs. a wide, sharp-edged and solid dorsolateral and its immaculate ventral surface (vs. darkly edged ventral scales). It is further distinguished from L. dorsalis by the fore-limb digital formula 0.2.3.4.2 (vs 0.2.3.4.3) and 3 lamellae on the 4th finger vs. 4-6 lamellae. It is distinguished from L. quadrivincula by smaller size (maximum SVL 45.10mm vs 51.13mm), fewer presacral vertebrae (36-38 vs 42) and a pale yellow tail differently coloured to the body, immaculate below vs. tail same colour as body, reticulated below. Lerista quadrivincula is known from a single specimen from the arid northwest coast of the Pilbara, WA, - 2,500km west of the known distribution of L. chordae. DISTRIBUTION. Lerista chordae occurs in the Desert Uplands, within an area encompassing a central zone of yellow and red sandy earths (20°-23°S) and, to the immediate west, alluvial sand plains (Fig. 3). It is associated with two open woodland vegetation types (see Habitat). HABITAT. Lerista chordae is a fossorial lizard occurring in open woodlands on sandy soils with a predominantly spinifex or other tussock grass dominated ground cover. It was found in 2 associated regional ecosystems types in the Desert Uplands bioregion. The specimens from Bede Station were trapped in the low, open Eucalyptus similis (ycllowjacket), Corymbia brachycarpa , C. setosa (bloodwood) and C. dallachiana (ghost gum) woodlands with a moderate to dense spinifex ( Triodia pungens) ground cover that is mediated by fire-age and grazing intensity (regional ecosystem 10.5.1 and 10.5.2, Sattler & Williams, 1999). This A NEW SPECIES OF LERISTA 129 HG 3. Map of collection localities of Lerista chordae in the Desert Uplands bioregion. EIU = Einasleigh Uplands, MGD = Mitchell Grass Downs, DEU = Desert Uplands, BRB = Brigalow Belt. Map sources: AUSLIG (1992), Queensland Herbarium June (2001), Queensland Environmental Protection Agency (1998). vegetation is widespread and occurs on the extensive, uniform, gently undulating Cainozoic sand deposits along the Great Dividing Range. Another recently described reptile for the Desert Uplands, Ctenotus rosarium Couper et aL, 2002, also occurs in this vegetation type. The specimens from Bede Station were captured in a pitfall trap from a long unbumt site (>8 years), where the habitat was characterised by dense spinifex cover (>60%). At Ulva Station and the sites south of Torrens Creek, specimens were located on small sandy rises within old alluvial sand plains. These sandy rises are dominated by mixed bloodwood and gum open woodlands (Corymb ia terminal is, C. dallachiana , C. plena) with a variable lower tree storey and shrub layer containing species such as Grevillea parallela, Carissa lanceolata , Eremophila mitchelli and Acacia spp. These patches are usually interspersed within more extensive Whites Ironbark (Eucalyptus white 0 communities. The ground layer is sparse, frequently dominated by Triodia pungens , with other graminoids such as Aristida spp., Paraneurachne muelleri , Heteropogon contortus, and Eriachne mucronata (present regional ecosystem 10.3.10 in mosaic with 10.3.9, Sattler & Williams, 1999). The soils consist of deep, unconsolidated sands, much like dune rises. This vegetation type occurs west and adjacent to the Cainozoic sand sheets described above, and on extensive sand sheets on the ancestral floodplains of the Flinders River (now centred on Torrens Creek). The specimen from Ulva Station was hand- captured and found sheltering under a log. At this locality the habitat was characterised by having a moderate tussock grass, forb and litter layer (>50%), and a discrete mid-storey shrub layer. DISCUSSION Lerista chordae is morph- ologically close to L. dorsalis , to which it would key in Cogger (2000) by virtue of its 4/4 digital formula, moveable eyelid, ear opening same size as nostril and well-developed dorsal stripes. Its 4/4 digital formula, moveable eyelid, 3 supraoculars, 5 supraciliaries, 6 upper labials and 1 8-20 midbody scale rows places it in the L.frosti group of Storr et al. (1999) along with L. dorsalis , L. flammicauda , L. frost i. L. quadrivincula and L. zietzi , and the more broadly defined L. elegans group of Wilson & Knowles (1988). However, none of the species groups within Lerista have been tested genetically so their phylogenetic validity is unknown. Some morphological character states have been used by Greer (Greer etal., 1983, Greer, 1986, 1990a) to redefine some species groups on the basis of synapomorphies, but most remain no more than aids to rapid identification. Work on the evolutionary relationships among species of Lerista is in progress using DNA sequence data and morph- ology (pers. comm. Adam Skinner, SA Museum and University of Adelaide). Preliminary data 130 MEMOIRS OF THE QUEENSLAND MUSEUM from both mitochondrial nuclear sequences consistently support the clade ((L. fragilis , L. chordae ), L. frosti)) but fail to find a close relationship between these three and L. dorsalis. While future work may better resolve relationships, it is clear at the present time that the DNA evidence supports recognition of L. chordae as a distinct species from L. dorsalis , and further suggests that its nearest relatives are more likely to be northern Australian taxa rather than the geographically remote L. dorsalis. In the light of the DNA information obtained so far, the extra phalanx on the fourth finger of L. dorsalis is intriguing. This character separates it from L. chordae as well as all other members of the L. frosti group. No variation was observed in these characters, in contrast to Greer’s (1987, 1990b) findings of up to 9.5% within a species, although our sample sizes are small, as phalanges were not always clearly visible in X-ray photographs (L. chordae n = 9; L. dorsalis n = 26; L.flammicauda n = 4; L. frosti n = 1\L. zietzi n = 7). Any functional significance of such a small difference is unclear. However, the sequence from L. dorsalis to other members of the L. frosti group (0. 2.3.4. 3 — ► 0.2. 3.4.2) fits within the schema of progressive limblessness within Lerista given by Greer ( 1 990b). The type of L. quadrivincula , as a member of the L. frosti morphological group, was examined for this study. This is the only known specimen of this species. As the type description (Storr, 1 990) is brief, a redescription, based on our observ- ations, is included here in the Appendix. No significant inconsistencies between Storr’s description and our observations were noted. A number of potential threats to the populations of L. chordae can be identified. The location of specimens in long unbumt spinifex at Bede Station suggests that the persistence of a high ground and Titter cover is required by this species. However, introduced Buffel Grass ( Cenchrus ciliaris ), which can alter and increase the fire intensity in spinifex communities is invading these ecosystems (Morgan et al., 2002). Furthermore regional ecosystem 10.3.10 and 10.3.9 in the Torrens Creek region has in recent years been heavily targeted for land clearing, with a further 40% (>100,000ha) proposed (Morgan et al., 2002). Cattle grazing in general can cause dramatic changes to ground cover, which may have some impact on fossorial species (Woinarski et al., 200 1 ). Though these threats are in part speculative and may seem exaggerated, the case of Lerista allanae , a species distributed on the eastern edge of the Desert Uplands and now considered probably extinct (Covacevich et al., 1996), suggests that even small reptiles are susceptible to long-term threats of land cover change associated with agriculture. ACKNOWLEDGEMENTS The Tropical Savanna CRC and James Cook University funded the survey of the Desert Uplands bioregion. John Woinarski and Alaric Fisher (Biodiversity Unit, Northern Territory Department of Infrastructure, Planning and Environment) and Jeanette Kemp (Queensland Herbarium) provided additional invaluable assistance throughout the survey. Doug Morrison (Environmental Protection Agency) kindly drafted Figure 3. Steve Peck (CS1RO) generously allowed access to the X-ray machine in his care. Fieldwork by SA Museum staff in the Torrens Creek area was carried out under scientific permit number F 1/00032 1/00/SAA. Patrick Couper (Queensland Museum) provided advice on the manuscript and provided Figure 2. LITERATURE CITED COGGER, H.G 2000. Reptiles and amphibians of Australia. (Reed New Holland: Sydney). COUPER, P.J., AMEY, A.P. & KUTT, A.S. 2002. A new species of Ctenotus (Scincidae) from central Queensland. Memoirs of the Queensland Museum 48( 1 ): 85-9 1 . COVACEVICH. J.A.. COUPER, P.J. & MCDONALD, K.R. 1996. Lerista allanae (Scincidae: Lygosominae): 60 years from exhibition to extinction? Memoirs of the Queensland Museum 39(2): 247-256. GREER, A.E., MCDONALD. K.R. & LAWRIE, B.C. 1983. Three new species of Lerista (Scincidae) from northern Queensland with a diagnosis of the wilkinsi species group. Journal of Herpetology 17(3): 247-255. GREER, A.E. 1986. Diagnosis of the Lerista hipes species-group (Lacertilia: Scincidae), with a description of a new species and an updated diagnosis of the genus. Records of the Western Australian Museum 13(1): 121-127. 1987. Limb reduction in the lizard genus Lerista 1. Variation in the number of phalanges and presacral vertebrae. Journal of Herpetology 21(4): 267-276. 1989. The biology and evolution of Australian lizards. (Surrey Beatty & Sons: Sydney). 1990a. The taxonomic status of Lerista aericeps Storr 1986 with a diagnosis of the Lerista orientals species group. Records of the Western Australian Museum 14(4): 443-448. A NEW SPECIES OF LERISTA 131 1990b. Limb reduction in the scincid lizard genus Lerista 2. Variation in the bone complements of the front and rear limbs and the number of postsacral vertebrae. Journal of Herpetology 24(2): 142-150. HORNER, P. 1992. Skinks of the Northern Territory. (Northern Territory Museum of Arts and Sciences: Darwin). KUTT, A.S. 2003. Patterns in the distribution and composition of the vertebrate fauna of the Desert Uplands Bioregion, Queensland. Unpubl. PhD. thesis, James Cook University, Townsville. MORGAN, G, LORIMER, M., MORRISON, A. & KUTT, A.S. 2002. The conservation of biodiversity in the desert uplands. (Environmental Protection Agency: Townsville). SATTLER, P. & WILLIAMS, R. 1999. The Conservation of Queensland’s regional ecosystems. (Environmental Protection Agency: Brisbane). SHEA, GM. 1991. A replacement name for Lerista concolor (Lacertilia: Scincidae). Records of the Western Australian Museum 15: 289. STORR, G.M. 1990, A new' Lerista (Lacertilia: Scincidae) from the Pilbara. Records of the Western Australian Museum 14(4): 669-670. STORR, GM., SMITH, L.A. & JOHNSTONE, R.E. 1999. Lizards of Western Australia. I Skinks. (Western Australian Museum: Perth). WILSON, S.K. & KNOWLES, D.G 1988. Australia’s reptiles. A photographic reference to the terrestrial reptiles of Australia. (Collins Publishers Australia: Sydney). WOINARSKI, J.C.Z., FENSHAM, R.. WHITEHEAD, P. & FISHER, A. 2001. Background paper 1. A review of changes in status and threatening processes. Pp. In Woinarski, J.C.Z., Whitehead, P. Fisher, A., Fensham, R. & Beggs, K. (ed.) Report to the national land and w r atcr resources audit. Vol (Cooperative Research Centre for the Sustainable Development of Tropical Savannas: Darwin). APPENDIX Redescription of Lerista quadrivincula Shea 1991 (original description by Storr, 1990, as Lerista concolor , a junior subjective homonym of Lygosoma (Rhodona) bipes concolor Werner, 1910, see Shea, 1991). Head scale diagrams are provided in Storr ( 1 990). Measurements. SVL (mm) = 51.13. Proportions (%SVL): AG = 67.40; tail broken; L 1 = 1 1 .32; L2 = 22.00; HL = 11.34, SA = 26.58. Proportions (%L2): LI = 51.47. Proportions (%HL): HW = 63.10; EE = 48.62. Scalation. Rostral crescent-shaped with triangular medial projection between nasals; nasals narrowly separated, nostril placed medially; frontonasal wider than long, saddle-shaped, triangular anteriorly between nasals to meet rostral, slightly concave posteriorly; frontal longer than wide, somewhat coffin-shaped, contacts frontonasal, prefrontals, first 2 supraoculars and frontoparictals; 3 supraoculars, 2nd the largest; supraciliaries 5, 1st, 3rd and 4th project between prefrontal and 1st supraoculars to contact frontal, 1st and 2nd supraocular, 2nd and 3rd supraocular respectively; palpebrals 7; frontoparietals paired and distinct; interparietal distinct, angular anteriorly, rounded posteriorly, overlapped by right parietal; parietals in contact behind interparietal; 4 enlarged nuchals; loreals 2, 1st largest; 2 preoculars, 1 presubocular; 2 postoculars, 1 postsubocular; primary temporal in contact with pretemporal, secondary temporal, 5th and 6th supralabials; secondary temporal in broad contact with parietal, primary temporal and pretemporal; supralabials 6, 4th subocular; infralabials 6, 2 contacting postmental; 2 additional rows of enlarged chin shields; ear opening circular, same size as nostril; midbody scale rows 20; paravertebrals 78; lamellae beneath 4th toe 7; supradigitals above 4th toe 5; lamellae beneath longest toe (3rd) 17; supradigitals above longest toe (3rd) 1 1 . Osteology. Presacral vertebrae 41. Fore-limb phalangeal formula 0.2. 3. 4. 2, hind-limb 0.2.3. 5. 3, Colour Pattern. Ground colour brown. Four longitudinal lines of discontinuous chocolate brown flecks running from nape to base of tail, breaking up into irregular flecking along tail. Chocolate brown dorsolateral band from nasal to base of tail, two half-scale widths on flanks, ill-defined, continuing along tail where it breaks up into irregular flecking merging with dorsal flecks and reticulate pattern ventrally. Head with sparse dark, fine flecks above. Limbs mottled with dark brown above. Tail same colour as body. Ventral surfaces immaculate except for reticulate pattern on tail. Supralabials edged with dark brown, some edging on infralabials also. Lateral scales mottled with dark brown merging with dorsolateral band. 132 MEMOIRS OF THE QUEENSLAND MUSEUM POR TMA CQUARJA NOM. NOV. PRO MACQUARIA BLOME, 2002 (NEMATODA: CHROMADORIDAE). Memoirs of the Queensland Museum 50(2): 132. 2005:- Macquaria Blome, 2002 (free-living marine nematodes) is preoccupied by Macquaria Cuvier in Cuvier & Valenciennes, 1830 (Percichthyidae: Pisces) and has to be replaced (ICZN, 1999. Art. 52, 53, 56,60). The type locality for this Australian sandy beach nematode genus is Port Macquarie, New South Wales and so the junior homonym is here replaced with Portmacquaria . Portmacquaria norn. nov., syn. Macquaria Blome, 2002 nec Macquaria Cuvier in Cuvier & Valenciennes, 1 830. Type species: Portmacquaria chimaira (Blome, 2002) comb. nov. (=Macquaria chimaira Blome, 2002). The genus is monotypic. Acknowledgements I thank Christian F. Kammcrcr, University of Chicago, USA, for making me aware of the homonymy. Literature Cited BLOME, D. 2002. Five new genera of Tree-living marine nematodes from sandy beaches of eastern Australia. Memoirs of the Queensland Museum 48: 29-43, CUVIER. G. & VALENCIENNES. A. 1830. Histoire naturelle des poissons. Tome cinquiemc. Livre cinquieme. Des Sci6noi'des. Hist. Nat. Poiss. i-xxviii + 1-499 + 4 pp. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE, 1999. International Code of Zoological Nomenclature. 4lh Edition, pp. i-xxix + 1-306. (London). Dietrich Blome, Breslauer Strafe 2, D-27616 Lunestedt, Germany (e-mail: dietrich.blome@t-online.de); 15 September 2004. A REVISION OF THE AUSTRALIAN ODACANTHINE GROUND BEETLES, INCLUDING CHECKLISTS FOR AUSTRALIA AND THE PAPUAN SUBREGION. (INSECTA: COLEOPTERA: CARABIDAE) MARTIN BAEHR Baehr, M. 2005 01 10: A revision of the Australian odacanthinc ground beetles, including checklists for Australia and the Papuan subregion (Insecta: Coleoptera: Carabidae). Memoirs of the Queensland Museum 50(2): 133-194. Brisbane. ISSN 0079-8835. The Australian Odacanthinae are revised, except for Giachinoana Baehr, Ophionea Klug (= Casnoidea Castelnau), Porocara Sloane, Renneria Baehr, and the brunnea-group of Dicraspeda Chaudoir. New species taxa described are: Archicolliuris occipitalis , A. splendissimus , Clarencia b rev i ceps, Deipyrus inops , Eudalia at rata, E.femorata , E. latipennis interioris subsp. nov., E. minor , E. obliquiceps punctifrons subsp. now, E. obliquiceps tozeria subsp. nov., E. punctipennis , E. reticulata , Gestroania setipennis , G. storeyi , Myrmecodemus lucai and M. pilosellus. Clarencia angusticollis (Macleay) and Eudalia waterhousei Castelnau are raised from synonymy to full specitic status. Eudalia froggatti Macleay is moved to Gestroania Liebke. Neoeudalia gen. nov., is erected for Eudalia nigra Sloane. Myrmecodemus (Trichodemus) subgen, nov., is erected for Myrmecodemus pilosellus sp. nov. To stabilise nomenclature, lectotypes, and associated paralectotypcs, arc designated for the following taxa: Anasis howittii Castelnau, Clarencia angusticollis (Macleay), C australis (Chaudoir), C. clarensii (Castelnau), Deipyrus palustris (Sloane), Dicraspeda brunneipennis (Sloane), D. obscura (Castelnau), Eudalia castelnaui Sloane, E. latipennis latipennis ( Macleay), E. waterhousei Castelnau, Gestroania fwggatti (Macleay), Myrmecodemus formicoides (Sloane), M. globulicollis (Macleay), and M. riverinae (Sloane). A neotype is designated for Neoeudalia nigra (Sloane). Notes are provided about species for which new material or information is at hand. A key to genera of Odacanthinae currently recorded from Australia is provided. Keys are also provided for the Australian species of Archicolliuris Liebke, Clarencia Sloane, Deipyrus Liebke, Dicraspeda Chaudoir, Eudalia Castelnau, Gestroania Liebke, Myrmecodemus Sloane, Ophionea Klug, and Porocara Sloane. Checklists are provided ol all odacanthine species recorded from Australia and the Papuan subregion, with some information about distribution. Representatives of all Australian genera and all new taxa described in this paper are figured. □ Coleoptera, Carabidae, Odacanthinae, Australia, New Guinea, checklists. Martin Baehr, Zoologische Staatssammlung, Munchhausenstr. 21, D-81247 Miinchen, Germany (e-mail: martin.baehf-@zsm.rnwn.de); received 20 December, 2003. Until recently the Australian odacanthine fauna was believed to be much less spcciose than that of other continents (Moore et ah, 1987). Despite this apparent species paucity, the Australian fauna is surprisingly diverse and appears to include some of the most plesiotypic Odacanthinae. My collecting in far northern and northwestern Australia convinced me that at least some genera arc much more speciose than suspected. Review of many Australian types and of many unidentified Odacanthinae in Australian museums and other collections found a new genus, a new subgenus and several new species. Certain previously synonymised species are returned to separate species rank. Recently described genera and species (Baehr, 1986, 1996a, b, 1999, 2003b, c), revaluation of Australian species (Baehr, 2003c), and new records of widespread Oriental species in Australia (Baehr, 2000) are incorporated in a checklist of the Australian odacanthine species. The brunnea- group of Dicraspeda Chaudoir (Baehr, 2003c), Ophionea Klug (=Casnoidea Castelnau), Porocara Sloane (Baehr, 1986, 1 996b, c), Renneria kamouni Baehr and Giachinoana carinipennis Baehr (Baehr, 1999, 2003b) are not added to herein. As Sloane’s (1910, 1917, 1923) partial keys to the Australian Odacanthinae are outdated, a new key to all known Australian genera is provided. Keys are also given for the species of all genera that include more than one species. Existing keys for Ophionea and Porocara (Baehr, 1986, 1 996b,c) are repeated or rearranged to cover only the Australian species, because no new data are available. A checklist is provided for the Papuan subregion based mainly on the work of 134 MEMOIRS OF THE QUEENSLAND MUSEUM Darlington (1968, 1971) and Baehr (1995, 1996a,b, 1997b, 1998, 2003a,c). Types of newly described species are shared with the respective collections, but holotypes and spare paratypes described from material from DPIM, Mareeba, and also paratypes and duplicates of species from my own collecting have been lodged in Queensland Museum. MATERIAL AND METHODS Altogether c. 900 specimens of Australian Odacanthinae were available for this study. About 80 additional specimens of New Guinean and Oriental Odacanthinae were used for comparison. The male genitalia were removed from specimens soaked for a night in a jar under wet atmosphere, then cleaned for a short while in hot KOH. For examination of the fine punctuation and microreticulation of the surface a high resolution stereomicroscope with up to 64 x magnification was used, supported by a lamp of high intensity giving natural light that could be focussed. For exact definition of the microsculpture such light is preferable, because fibre-optics lights sub- stantially change perception of the surface structures. The habitus photographs were taken with a digital camera using SPOT Advanced for Windows 3.5 and subsequently were worked with Corel PhotoPaint 10. Measurements were taken using a stereo- microscope with an ocular micrometer. Length has been measured from apex of labrum to apex of elytra. Lengths, therefore, may slightly differ from those of other authors. Length of eye includes a small dark coloured ring of ocellae that in some instances is present behind the light area. Length of orbit is taken from posterior margin of eye to ‘neck’ suture. Length of head is the distance from apex of labrum to ‘neck’. Length of pronotum was measured from the most advanced part of base to the most advanced part of apex; width of pronotum at widest part, including those parts of the proepistemum that are visible from above. Length of elytra was taken from the most advanced part of humerus to the most advanced apex of elytra including any apical denticles or spines. ABBREVIATIONS. Collections. ANIC, Australian National Insect Collection, Canberra; BMNH, The Museum of Natural History, London; CBM, Working Collection M. Baehr, Munchen; CFP, Collection S Facchini, Piazcnca; CGT, Collection P. M. Giachino, Torino; CMP, Carnegie Museum, Pittsburgh; CRC, Cooperative Research Centre for Tropical Rainforest Ecology and Management, Cairns; CSM, Collection R. Sciaky, Milano; CTV, Collection L. Toledano, Verona; DEI, Deutsches Entomologisches Institut, Eberswaldc; HNMB, Hungarian National Museum of Natural History, Budapest; MCSN, Museo Civico di Storia Naiurale, Genoa; MCZ, Museum of Comparative Zoology, Cambridge/Mass.; MNHB, Museum fur Naturkunde der Humboldt-Universitat, Berlin; MNHP, Museum National d’Histoirc Naturellc, Paris; NMNHP, National Museum of Natural History, Prague; MV, Museum of Victoria, Melbourne; MDPI, Queensland Department of Primary Industries, Mareeba; QM, Queensland Museum, Brisbane; SAM, South Australian Museum, Adelaide; UQIC, University of Queensland Insect Collection, Brisbane; ZSM, Zoologische Staatssammlung, Munchen. Collectors. Collectors’ names are abbreviated as follows: A. C’alder (AC), A. D. Selby (AS), A.J. Watts (AW), B. Cantrell (BC), .1. Balderson (BJ), Britton & Misko (BM), J. Bugeja (BU), I.C. Cunningham (CC), J. Cardale (CJ), C. Oke (CO), C. Vallis (CV), M. & G. De Baar (DB), D. Cook (DC), S. De Faveri (DF), G. Dickinson (DG), D. Heiner (DH), D.C. F. Rentz (DR), W. Dressier (DW), D.K. Yeates (DY). E.B. Britton (EB), E.C. Dahms (ED), E.D. Edwards (EE), E.G. Matthews (EM), E.S. Nielsen (EN), E.F. Rick (ER), E. Sutton (ES), Earthwatch/Qld. Museum (EW), F.P.Dodd (FD), Fay & Halfpapp (FH), F.G. Satter(FS), F. E. Wilson (FW), J.G. Brooks (GB), G Daniels (GD), G. Hangav (GH), GB. Monteith (GM), H.W. Brown (HB), H.W. Davcy (HD), H.W. Grawes (HG), K. Hyde (HK), H. Mitchell (HM), H. Shepherdson (HS), I.F.B. Common (IC), Ian Naumann (IN), J.D. Brown (JB), J.H. Calaby (JC), J.E. Feehan (JF), J. Hasenpusch (JH), J F. Lawrence (JL). J.H. Taylor (JT), J.W. Schomberg (JS), J.L. Wassell (JW), K.H. Halfpapp (KH), L.P. Kelsey (LK), L.A. Ring (LR), L. Toledano (LT), Lea & Wilson (LW), M. Baehr (MB), M. Mendum (ME), M.S. Moulds (MM), M.S. Upton (MU), N. Gough (NG), N.B. Tindale (NT), O. Hillert (OH), P. Aitken (PA), P.M. Giachino (PG), P.M. Hammond (PH), P. Machacek (PM). P. Zborowski (PZ), R.A. Barrett (RB), R. Kitching (RK), R. Olivieri (RO), R.I. Storey (RS), S. Bily (SB), Storey, Brown & Jacobson (SBJ), S. Feam (SF), S.A. Hogcnhout (SH), S. R. Monteith (SM), T. Gush (TG), T. Weir (TW), V. Framenau (VF), W.D. Dodd ( WD), A. & M. Walford- Huggins (WH), W.J.M. Vestjens (WV). Localities. For New Guinean localities PNG =Papua New Guinea, and IJ means Irian Jaya, the Indonesian part of the island which recently was renamed West Papua. TAXONOMIC PRINCIPLES Male genitalia, which are widely used for the distinction of carabid species, rarely yield good AUSTRALIAN ODACANTHINE GROUND BEETLES 135 distinctive characters in Odacanthinae. Odacanthine aedeagi, except for those of the most highly evolved species, generally lack complex and/or sclerotised internal structures and the folding of the internal sac is quite simple. The most striking differences are in size, overall shape and sometimes in the shape of their apices. Female stylomeres rarely yield characters even for generic differentiation. Nevertheless, distinction of species is reasonably easy, because most species differ in external structural characters that are generally easy to detect. Besides shape and relative size of head, prothorax, and elytra, chaetotaxy and structure of the surface, in particular striation of the elytra, punctuation, microreticulation, and degree of pilosity, are of major importance. Colouration can vary in certain species, but colouration of appendages and pattern of elytra, if present at all, seem to be fairly constant and can be of some value. Recognition of insect subspecies is a matter of opinion, or better, a taxonomic hypothesis, when the actual interruption of gene How between populations is not known. This is especially the case for insular populations. Such decisions are extremely difficult, particularly when the respective taxa cither are able and willing to fly over considerable distances, or are easily trans- ported by natural or human carriers. Classifications, therefore, are based mostly on morphological characters, at least where insects of almost unknown habits and life histories are concerned, such as those in the present paper. 1 use sub- species for taxa that exhibit small morphological differences and are allopatric (or at least allotopic), and I use species for taxa that are more strikingly different and/or are sympatric. Generic concepts in the Australian Odacanthinae are generally well founded, with a few exceptions. Sloane (1917,1 923) who founded the systcmatics of the Australian Odacanthinae, was uncertain about the affiliation of the many differently shaped species of the Dicraspeda- Eudalia- complex and he changed his view several times. Liebke (1938) separated Dicraspeda and Eudalia , but he also split from Dicraspeda the genera Philemonia Liebke and Macrocentra Chaudoir and arranged them at quite different places in his system. Darlington (1968) adopted the present system, combining Philemonia and Macrocentra with Dicraspeda and separating Eudalia. This concept is followed here, but it should be noted that Dicraspeda in its present concept is remarkably heterogeneous and could be dismembered in future (Baehr, 2003c). Comparable problems are faced in Eudalia which is also quite diverse and could be divided into two subgenera or genera. This is more likely when the rather aberrant New Guinean E. anomala Darlington is taken into consideration. Liebke (1931, 1938) subdivided Colliuris De Geer into various subgenera, most of which are retained. Many have been raised to generic rank, but these decisions were not recognised by all authors, or they were adopted to different degrees. As no thorough phylogenetic work has been done on the Colliuris- complex, its division into genera and subgenera is still a matter of opinion. I follow Lorenz (1998). NOMENCLATORIAL NOTE. A number of specimens of previously described species were designated holotype by RJ. Darlington, probably during his stay in Australia in 1956-58. Some of these designations were apparently done without comparing the original descriptions, and in some specimens, even without reading the labels. Some bear written cotype or even topotype labels, and should not have have been designated holotype. It is difficult to imagine why the rules of nomenclature were neglected by a leading taxonomiust in these instances. As a consequence, nomenclatorial decisions of Darlington a priori should be checked very carefully. KEY TO AUSTRALIAN GENERA OF THE ODACANTHINAE 1 . 4th tarsomcrcs deeply excised in middle and markedly bilobate 2 4th tarsomeres not deeply excised in middle and not markedly bilobate 3 2. Head elongate, more or less distinctly triangular; prothorax rather tubular, without distinct lateral sulcus; elytra narrow, upper surface depressed, red with black or blue pattern and white spots (in Australian species) (Fig. 38D) Ophionea Klug Head not elongate, convex; prothorax not tubular, with distinct lateral sulcus; elytra rather short and wide, upper surface not markedly depressed, uniformly black or dark piceous (in Australian species) (Fig. 34D, E) Dicraspeda Chaudoir (part) 3. Head not distinctly narrowed behind eyes; colour of surface brick-red, elytra with serrate, cruciate dark pattern (Fig. 38E) Porocara Sloane Head distinctly narrowed behind eyes; colour of surface different, elytra with different pattern, or unicolourous 4 4. 3rd antennomere very elongate, as long as 4th and 5th together (Fig. 33F,34A) Clarencia Sloane 3rd and 4th antennomeres of about equal length .... 5 5. Elytra elongate, parallel, upper surface remarkably depressed, apex almost transverse, without any excision (Fig.33A) v4nastsCastclnau 136 MEMOIRS OF THE QUEENSLAND MUSEUM Elytra usually shorter and less parallel; when elongate, then upper surface not markedly depressed and apex oblique, usually more or less distinctly excised .... 6 6. Head with distinct longitudinal sulcus and ridge inside of eye . 7 Head without or with indistinct sulcus and ridge inside of eye , 11 7. Odd elytral intervals cariniform, external apices of elytra spiniform (Fig. 37B) Giachinoana Baehr Odd elytral intervals not cariniform. external apices of elytra not spiniform, at most gently angulatc (in Australian species) 8 8. Pronotum with indistinct lateral margin, or margin not medially bordered by a deep sulcus; elytra glossy black with 2 or 4 small white spots (in Australian species) (Fig. 33B.C) Archicolliuris Liebke Pronotum with very conspicuous ridge-like margin, margin medially bordered by a deep sulcus; elytra piceous or black, without white spots 9 9. Elytra deeply punctate-striate in basal third, barely striate in apical two thirds; with deep transverse sulcus in basal third; elytra rather narrow and elongate; lateral margins of pronotum conspicuously sinuate (Fig. 33E) • • • * Basistichus Sloanc Elytra fully striate, or striation becoming gradually weaker towards apex; without or with only shallow' transverse sulcus in basal third, in latter case elytra more depressed and rather wide; lateral margins of pronotum barely sinuate 10 10. Surface with dense, elongate, erect pilosity; elytra with shallow transverse depression near apex, 7th interval tumid in apical third; 3rd antennomcre sparsely setose; tibiae dark with conspicuous yellow' ring (Fig. 38C) Neoeudalia gen. nov. Surface without pilosity; elytra without transverse depression near apex, 7th interval not tumid; 3rd antennomcre glabrous, except for apical setae; tibiae uniformly yellow or dark, w ithout yellow ring (Fig. 34D, F) Dicraspeda Chaudoir 1 1. 3rd antennomere impi lose 12 3rd, and usually also 1st and 2nd antennomeres, pilose. 15 12. Elytral striae impunctate, rather sulcate; head markedly triangular towards base (Fig. 33D) . Aulacolius Sloane Elytral striae punctate, not sulcate; head convex behind eyes 13 1 3. Lateral margin of pronotum medially bordered by a deep sulcus; all odd intervals with a row' of many (>10) erect setae (Fig. 38F) Renneria Baehr Lateral margin of pronotum medially not bordered by a deep sulcus; at most 3rd, 5th and 7th intervals with a row' of fewer (<5) erect setae 14 14. Elytra ampliate, apically considerably widened and externally ungulate or spinose; antenna very elongate, 4th antennomere not pilose (Fig. 36D-F, 37A) • * * • Gestroania Liebke Elytra not ampliate, apically not or little widened and externally not angulatc; antenna shorter. 4th antennomere pilose (Figs 34F, 35, 36A-C) EudaliaC astelnau 15. Elytral striae almost complete; prothorax densely punctate and pilose on disk 16 Elytral striae incomplete, either only basal third striate, or but 2 inner striae complete; prothorax in middle impunctate, either glabrous or coarsely, transversely rugose, impilose on disk (Fig. 37D-F, 38A-B) Myrmecodemus Sloane 16. Lateral margin of pronotum medially bordered by a deep sulcus; elytral striae punctate-striate (Fig. 34B-C) Deipyrus Liebke Lateral margin of pronotum medially not bordered by a deep sulcus; elytral striae only coarsely punctate (Fig. 37C) Lachnothorax Motschulsky Anasis Castelnau, 1867 Anasis Castelnau, 1867: 15; 1868: 101; Csiki, 1932: 1537; Liebke, 1938: 93; Moore et al., 1987: 276; Lorenz, 1998: 420. TYPE SPECIES. Anasis howittii Castelnau, 1867, by monotypy. DIAGNOSIS. Body elongate; elytra very elongate, parallel, dorsally remarkably depressed, highly glossy, impilose; elytral apices almost transverse, not at all sinuate. RELATIONSHIPS. According to shape and structure this is an isolated genus. It may be nearest to Eudalia as an early offshoot of the Eudalia lineage. This suggestion is supported by the S Vic range of the single species (if correct as reported). Only species of Eudalia occur with Anasis in mesic Vic. and in adjacent parts of SE Australia. Anasis howittii Castelnau, 1867 (Figs 30A, 33A) Anasis howittii Castelnau, 1867: 15; 1868: 101; Csiki, 1932: 1537; Liebke, 1938: 93; Moore ct al., 1987: 276; Lorenz, 1998:420. TYPE MATERIAL. LECTOTYPE (here designated): 9 (slightly damaged), Geelong ‘JuT (? uncertain) - Lewis/ Anasis howittii Cvs\J Anasis Howittii/Cast./Type/ Holotype T-17960 (MV). Although the specimen is labeled ‘holotype’, this is questionable, because the description says nothing about the number of examined specimens, only that the species was captured on a flower. Moore et al. ( 1 987) wrote ‘syntypes (possible)’, but did not indicate that multiple specimens were used in the original study. Because this species has apparently never been recaptured, and as it remains enigmatic, never having been mentioned in a subsequent paper or key, I am giving a partial rcdescription. DESCRIPTION. Measurements. Length: 8.1mm; width: 2.35mm. Ratios. Length eye/orbit: 1.0; length/width of head: 1.25; length/width of prothorax: 1 .32; length/width of elytra: 1 .88. AUSTRALIAN ODACANTHINE GROUND BEETLES 137 Colour (Fig. 33A). Upper surface chestnut-brown, elytra slightly lighter than fore body, mouthparts, antennae and legs reddish. Head. Eyes large, laterally protruding, orbits moderately convex. Surface with 2 impressions behind clypeus, a triangular central one on frons, and 2 circular ones between eyes. Medially of eye without any ridge and sulcus. Posterior supra- orbital seta located well behind eye. Mentum with elongate, triangular tooth, with 2 setae behind tooth, submentum with an elongate seta on either side. Glossa and lacinia not examined due to condition of holotype. Antenna elongate, probably just surpassing base of pronotum. Three basal antennomeres glabrous. Surface glossy, impunctate, impilose, without microreticulation. Pronotum. Elongate, lateral margin gently convex in anterior half, slightly sinuate towards base, margin distinct, forming a sharp ridge. No sulcus inside of margin. Disk slightly convex. Proepisternum narrowly visible from above. Punctation of disk dense and coarse. Surface impilose, without microreticulation, glossy. Elytra. Remarkably elongate, dorsally conspicuously depressed, very gently widened towards apex. Humeri indistinct, obliquely rounded, lateral margin very gently convex, apex almost transverse, gently convex, without any sinuation. Striae complete, coarsely punctate, though not impressed, punctac smaller towards apex. Intervals depressed, impunctate. 3rd interval tripunctate, fixed setae in basal third, middle, and apical third. Surface without micro- reticulation, very glossy. Hind wings present. Lower Surface. Thorax with coarse, dense punctuation, impilose, glossy; abdomen impunctate. Metepisternum very narrow, elongate, >3 x as long as wide at apex. Terminal abdominal sternum in female quadrisetose. Legs. Narrow, elongate. Tarsi not lobed, 5th tarsomere with a dense fringe of elongate setae below. Claws small, thick, smooth. Female Genitalia (Fig. 30A). Stylomere 2 very elongate, median margin characteristically angled above middle, with elongate, acute apex, with 3 elongate ventrolateral ensiform setae almost completely on the ventral surface, a rather small dorsomedian ensiform seta about middle of stylomere. and a single short nematiform seta arising from a groove below apical third. Base of stylomere 1 with 6 elongate ensiform setae. Lateral plate with dense fringe of stiff nematiform setae at apical rim. DISTRIBUTION. Type locality only. Archicolliuris Liebke, 1931 Archicolliuris Liebke, 1931: 291; 1938: 61; Csiki, 1932:1525; Lorenz, 1998:418. Colliuris De Geer, 1774: 79; Darlington, 1968: 204; Moore et al„ 1987: 277. TYPE SPECIES. Casnonia bimaculata Kollar & Redtenbacher, 1 842, by original designation. DIAGNOSIS. Elongate; head posteriorly tri- angular; pronotum more or less elongate, rather tubular; elytra elongate, dorsally depressed. Head and pronotum without additional setae; elytra with setae on 3rd or 3rd, 5th and 7th intervals. Surface generally glabrous, glossy; elytra usually with 2 or 4 light spots. Archicolliuris par (Darlington, 1968) Colliuris par Darlington, 1968: 206; Moore et al., 1987: 277. Archicolliuris par (Darlington); Lorenz. 1998: 418. TYPES. HOLOTYPE and 5 PARATYPES: Hollandia, July-Sept. 1944, P.J. Darlington (MCZ) (not seen). DIAGNOSIS. Relatively small, black, quadrimaculate; pronotum moderately elongate; elytra with moderately deep transverse sulcus; surface of head and pronotum with fine though distinct microreticulation. SUPPLEMENTARY DESCRIPTION. Measurements (Table 1). DISCUSSION. This New Guinean species was recorded from the northern tip of Cape York Peninsula by Darlington (1968). I have not examined type material, but have seen a specimen from New Guinea that agrees well with the description and is distinct from A. splendissimus sp. nov. from Cape York Peninsula. As I have not seen any genuine Australian specimen of A. par , Darlington’s record probably refers to A. splendissimus and A. par does not occur in Australia. DISTRIBUTION. ? extreme tip of Cape York Peninsula; New Guinea, New Britain. RELATIONSHIPS. This species is closely related to A. splendissimus sp. nov. Archicolliuris splendissimus sp. nov. (Figs 1, 30B, 33B) ETYMOLOGY. Refers to the highly glossy surface. MATERIAL. HOLOTYPE: (J, Lockerbie, 0- 31.iii.64, I.F.B. Common & M.S. Upton/ Colliuris par Darlington, det. T.A. Weir 2000 (ANIC). PARATYPES: 9, ‘Eclectus’ Iron Range, Qld 12°45’46”S 143°17’10”E, 10 June 1995, 138 MEMOIRS OF THE QUEENSLAND MUSEUM 20m, G Daniels, mv lamp (UQIC); 9 , Iron Range, Cape York Pen.. 11-1 7.V.1968, G Monteith (QM); 9 , same data but 28.iv-4.v.l968 (CBM). DIAGNOSIS. Relatively small, black, quadri- maculate: head short, wide; pronotum short, globose; elytra with deep transverse sulcus in basal third; surface without microreticulation, remarkably glossy. DESCRIPTION. Measurements (Table 1). Colour (Fig. 33B). Black, elytra with 4 small circular to slightly elongate light yellow spots, the anterior, larger ones situated in 5th interval, the posterior ones in 5th and 6th intervals. Mandible dark reddish with blackish tip, palpi black with dark reddish basal palpomeres, antennae black with 3 dark reddish basal antennomeres. Basal half of femora reddish, apical half black, tibiae dark piceous, tarsi black. Head . Short, very wide, markedly rhomboidal. Neck very narrow, with deep transverse impression. Eyes large, protruding, though not much separated from orbits which are very gently convex, though remarkably oblique. Surface with 2 deep impressions behind clypeus, frons in middle with a horseshoe like impression, also the anterior supraorbital seta in deep, circular impression. Medially eye with a shallow sulcus, but without ridge. Posterior supraorbital seta in front of posterior margin of eye. Mentum with short, triangular tooth, with 2 setae behind tooth, also submentum with an elongate seta on either side. Apex of glossa transverse, laterally oblique, with 2 elongate median and 2 short lateral setae. Paraglossae free, narrow, glossy. Lacinia elongate, interior margin with a fringe of few spines. Antenna short, not attaining base of pronotum. Three basal antennomeres glabrous. Surface very glossy, impunctate and impilose, without microreticulation. Pronotum. Short, laterally and dorsally convex, near apex and base with deep transverse impressions. Apex not bordered, lateral margin complete though indistinct, base with thick border, median line very shallow. A single marginal seta in front of middle. Proepistemum narrowly visible from above. Disk impunctate, except near apex and base. Surface without microreticulation, highly glossy. Elytra . Short, wide, subrectangular, gently widened towards apex, surface moderately convex. Base oblique, humeri distinct, obtusely rounded, lateral margin gently incised at basal fourth, apex slightly oblique, gently excised, external apical angles distinct, though obtuse. FIG 1. Archicolliuris splendissimus sp. nov. Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). Base narrowly margined towards 4th stria, apex distinctly margined. Surface in anterior third with a deep, irregularly transverse impression, disk humped in front of impression. 7th and 8th striae near apex conspicuously tumid. Striae incomplete, beginning behind base, external striae not attaining apex. Striae not impressed, in basal half coarsely punctate, becoming very inconspicuous towards apex. Intervals depressed, impunctate. Scutellar stria elongate, consisting of about 10 punctures. 1st interval with 2 setae in basal third, 3rd and 5th intervals multipunctate. Surface without microreticulation, very glossy. Hind wings present. Lower Surface. Proepistemum and prostemum coarsely punctate in basal half and near sternal suture, proepimeron and mesothorax similarly punctate. Abdomen impunctate. Whole lower surface impilose and very glossy. Metepistemum narrow and elongate almost 3 x as long as wide at apex. Terminal abdominal sternum in male bisetose, in female quadrisetose, and with short pilosity in middle of apex. Legs. Narrow, elongate. Tarsi not lobed, 5th tarsomere with a dense fringe of elongate setae below. Claws smooth. Male anterior tarsus biseriately squamose at apical half of 1st tarsomere and at 2nd and 3rd tarsomeres. Male Genitalia (Fig. 1). Terminal abdominal stemite in middle incised. Genitalia medium sized. Genital ring strongly sclerotised, elongate, slightly asymmetric, parallel, shortly narrowed to the obtusely rounded apex. Aedeagus compact, depressed, laterally remarkably sinuate, sinuation even somewhat angulate, lower surface very gently bisinuate. Apex short, gently AUSTRALIAN ODACANTHINE GROUND BEETLES 139 TABLE 1. Measurements and ratios of all Australian species of Archicolliuris . N length (mm) length eye/orbit 1/w head 1/w prothorax 1/w elytra ' par 1 6.6 1.36 1.18 1.92 1.60 splendissimus 2 6. 1-6.9 1.42-1.48 1.15 1.46-1.57 1.66-1.72 occipitalis 2 9.5-10.5 0.50-0.51 1.69-1.73 2.03-2.08 1.75-1.77 upturned, barely knobbed, slightly turned to right, incision at right side moderately deep. Orificium short. Folding of internal sac very simple. Parameresof rather similar shape, though left paramere much larger than right one, left one with very gently convex apex, right one with wide, rounded apex. Female Genitalia (Fig. 30 A). Stylomere 2 comparatively elongate, laterally evenly curved, with acute apex. With 2 large ventrolateral ensiform setae, a rather large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 8-9 elongate ensiform setae. Lateral plate with dense fringe of stiff, elongate nematiform setae at apical rim. Variation. Very little variation noted. DISTRIBUTION. Northern half of Cape York Peninsula. Archicolliuris occipitalis sp. nov. (Figs 30C, 33C) ETYMOLOGY. Refers to the very elongate occiput. MATERIAL. HOLOTYPE: 9 , NE Qld, Roaring Meg Ck, 6km W Cape Tribulation, 22 April 1983, GB. Monteith, D.K. Yeates/QM Berlesate No. 536. 16.05S 145.24E, Rainforest 710m, moss (QMT 99168). PARATYPE: 1 9, Mt Lewis, Via Julatten, N Qld. 3,500-4,000’ 27-28.XI.1965. GM JClarencia sp. det. B.P. Moore 74 (CBM). DIAGNOSIS. Comparatively large; occiput very elongate; prothorax tubular, elongate, bimaculate or indistinctly quadrimaculate elytra; pronotum with dense transverse striolation. DESCRIPTION. Measurements (Table 1). Colour (Fig. 33C). Black, elytra with 2 small, distinct though irregularly shaped orange spots in basal third on 4th-6th intervals, and with an indistinct small spot in basal third that is composed of 2 narrow reddish lines on 4th and 5th intervals. This spot may be almost completely reduced. Mouthparts and 2 basal antennomeres light reddish, median antennomeres very slightly darker. Basal 2/5 or 1/2 of femora light reddish, apical 1/2 black, though knees and tibiae piceous, tarsi reddish to reddish piceous. Lower surface black, abdomen in middle and near apex also laterally, reddish to piceous. Head. Narrow, very elongate. Neck very narrow, with deep transverse impression. Eyes relatively small, laterally protruding, slightly separated from orbits which are very elongate, and gently convex. Surface with 2 deep irregular impressions behind clypeus that combine to a horseshoe like impression in middle of frons. Medially of eye with a shallow sulcus, but without ridge. Posterior supraorbital seta located far behind posterior margin of eye. Mentum with short, though markedly acute, triangular tooth, with 2 setae behind tooth, also submentum with an elongate seta and a short seta on either side. Apex of glossa transverse, with 2 elongate median and 2 shorter lateral setae. Paraglossae free, narrow, surpassing glossy. Lacinia elongate, interior margin with a fringe of rather few spines. Antenna elongate and remarkably thin, slightly surpassing base of pronotum. Three basal antennomeres glabrous. Surface moderately glossy, impunctate, impilose; basal halt with extremely fine and highly superficial microreticulation composed of very transverse meshes and lines. Immediately at base surface covered with transverse sulci. Pronotum. Elongate, tubular, laterally convex, near apex and base with very shallow transverse impressions. Apex bordered, lateral margin complete though indistinct, almost straight, base with thick border, median line very shallow. A single marginal seta situated slightly in front of middle. Proepistemum well visible from above behind middle. Disk with very dense and remarkably coarse transverse sulci. Surface apparently impunctate, with microreticulation at least within sulci, remarkably rugose. Elytra. Moderately elongate, conspicuously widened in apical half, surface moderately convex. Base very oblique, humeri indistinct, very widely rounded, lateral margin distinctly incised at basal fourth, apex very oblique, gently excised, external apical angles distinct, though obtuse, sutural angles produced, slightly disjoined. Base margined towards 4th stria, apex distinctly margined. Surface in anterior third with a deep, irregularly transverse impression, disk gently tumid in front of impression. Striae al most 140 MEMOIRS OF THE QUEENSLAND MUSEUM complete, originating near base, attaining apex, but becoming very inconspicuous towards apex. Striae not impressed, in basal half coarsely punctate. Intervals generally depressed, impunctate, though 5th and 6th intervals narrow and convex within the area of the basal transverse impression. Scutellar stria moderately elongate, consisting of 6-8 punctures. 1 st interval unisetose in basal third, 3rd interval polysetose, 5th interval with 2-3 setae in basal half. Surface without microreticulation, very glossy. Hind wings present. Lower Surface. Proepistemum, prostemum, and mesepistemum coarsely punctate. Metathorax and abdomen impunctate. Whole lower surface impilose, with very fine microreticulation. Metepistemum narrow and elongate, c. 3 x as long as wide at apex. Terminal abdominal sternum in female quadrisetose and with short pilosity in middle of apex. Legs. Very narrow and elongate. Tarsi not lobed, 5th tarsomere with a dense fringe of elongate setae below. Claws large, smooth. Male Genitalia. Unknown. Female Genitalia (Fig. 30C). Stylomere 2 fairly elongate, laterally evenly curved, with acute apex. With 2 elongate ventrolateral ensiform setae, a rather large dorsomedian ensiform seta situated in middle of stylomere, and a single short nematifonn seta arising from a groove in apical third. Base of stylomere 1 with 9-10 moderately elongate ensiform setae. Lateral plate with dense fringe of elongate, stiff nematifonn setae at apical rim. Variation. Little variation noted due to limited material. One specimen almost completely lacks the anterior elytral spot. DISTRIBUTION. Base of Cape York Peninsula in rainforest at high altitude. Hence, this might be an arboreal rather than hygrophilous species. RELATIONSHIPS. This species is distantly related to Australian and New Guinean species of Archicolliuris , and in shape and structure is more similar to certain Oriental species. KEY TO THE AUSTRALIAN SPECIES OF ARCHICOLLIURIS I . Head very elongate, orbit c. 3 X as long as eye; pronotum with extremely dense and coarse transverse wrinkles; elytra with anterior light spot indistinct or absent, though when present, not circular; body length >9mm occipitalis sp. nov. Head much shorter, orbit <1.5 x as long as eye; pronotum without or with weak transverse wrinkles; elytra with anterior light spot distinct, circular; body length <7mm 2 2. Head and pronotum with faint though distinct microreticulation; pronotum less orbicular; elytra with less deep transverse sulcus par Darlington Head and pronotum without any traces of micro- reticulation, highly glossy; pronotum remarkably orbicular; elytra with deep transverse sulcus. * splendissimus sp. nov. Aulacolius Sloane, 1923 Autacolius Sloane, 1923: 32; Csiki, 1932: 1537; Licbke, 1938: 94; Moore ct al., 1987: 276; Lorenz, 1998: 420. TYPE SPECIES. Aulacolius triondinatus Sloane, 1 923, by monotypy. DIAGNOSIS. Elytral striae impunctate though complete, deep and sulcate; body compact; head remarkably elongate, triangular. Aulacolius triordinatus Sloane, 1923 (Figs 2, 30D, 33D) Aulacolius triordinatus Sloane, 1923: 32; Csiki, 1932: 1537; Licbke, 1938: 94; Moore et al., 1987: 276; Lorenz, 1998: 420. MATERIAL. Holotype: 9, GF. Hill Darwin, NT/Type/ Aulacolius triordinatus SI. Type/HOLOTYPE A. triordinatus SI., P.D. (ANIC). New records (13 ex.): NT: South Alligator R., 19.12.1999, MB (CBM); Crocodile I., HS (SAM). Qld: Momington 1. Mission, 12.5.1963, 15.5.1963, 23.5.1963, 5.1963, PA & NT (SAM); Stewart R, WD (SAM). DIAGNOSIS. Head markedly triangular; pronotum short, dorsally convex, with sharp, ridge-shaped lateral borders; elytra short, compact, with complete, deeply impressed, sulcate striae and yellow-spotted apex; several marginal seta on prothorax; numerous setae on intervals 3, 5, and 7; surface absolutely glabrous, shining. SUPPLEMENTARY DESCRIPTION. Male Genitalia (Fig. 2). Terminal abdominal stemite in middle incised. Genitalia comparatively large. Genital ring fairly elongate, barely asymmetric, rather parallel, shortly narrowed to the narrow, acute, triangular apex. Aedeagus very slender and elongate, depressed, laterally barely sinuate, lower surface only immediately near base concave, in apical three quarters gently convex. Apex elongate, very depressed, straight, not knobbed, but remarkably spoon-shaped, slightly turned to right, incision at right side deeper than at left side. Folding of internal sac very simple. Parameres of similar shape, though left paramere much larger than right one both with wide, rounded apex. AUSTRALIAN ODACANTHINE GROUND BEETLES 141 FIG. 2. Aulacolius triordinatus Sloanc. Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). Female Genitalia (Fig. 30D). Stylomere 2 comparatively elongate, laterally evenly curved, with acute apex. With 2 small ventrolateral ensiform setae, a large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 7-8 elongate ensiform setae. Lateral plate with dense fringe of stiff nematiform setae at apical rim. DISTRIBUTION. Moore et al. (1987) gave distribution as northern NT. New records extend its range to Momington Island in the Gulf of Carpentaria and lower Cape York Peninsula. COLLECTING CIRCUMSTANCES. One specimen collected at light. Basistichus Sloane, 1917 Basistickus Sloane. 1917: 415; 1923: 30; Csiki, 1932: 1535; Licbke, 1938: 81; Darlington, 1968: 208; Moore et al., 1987: 276; Lorenz, 1998: 420. TYPE SPECIES. Odacantha micans Macleay, 1864, by original designation. DIAGNOSIS. Elytra elongate, parallel, with unequal striation. Basistichus micans (Macleay, 1864) (Figs 3, 30E, 33E) Odacantha micans Macleay, 1864: 107. Basistichus micans , Sloanc, 1917: 415; Csiki, 1932: 1535; Licbke, 1938: 81; Darlington, 1968: 208; Moore et al., 1987: 276; Lorenz, 1998: 420. MATERIAL. SYNTYPES: 3 ex., in poor condition, ‘Port Denison’, (see Moore et al., 1987: 276) (AN1C). NEW RECORDS (55 ex.): Qld: 15km S Marlborough, 21.L1982, MB (CBM); 20km N Biggenden, 2211982, MB (CBM); Undara, 12.ii.2000, SB (CBM); Qld 01/31, Me Leod River, 12km NW Mt Carbine, 12.iv.2001, MB (CBM); 3km E of Lockerbie, Cape York, 30.i.-4.ii.l975, GM (QM); Odacantha micans McLeay jun. Port Denison (MV); Casnonia micans Sloane, Cooktown (MV); Kuranda, GB (MV); Mutchilba, xii.1936, AS (MV); Cairns, 1 1 .i.50, CO (MV); Townsville, 8.xi.02, FD (MV); C. micans Macl. Rockhampton (MV); 70km SW Greenvale, 8-15.xii.1995, AW (SAM); Einasleigh R. via Mt Surprise, 6-711980, RS (MDPI); 11km WSW of Petford, 17. i. 1987, 3-4.iv.1988, RS (MDPI); Tolga, 25. i. 1985, JB (MDPI); 7km NE of Tolga, ii.1989, RS (MDPI); Pouth Ck via Geoigetown. 81 1 980, RS (MDPI); Marecba, l.ii.1979, KH (MDPI); 27km E Forsyth. 29.xii. 1 977, RS (MDPI ); 12km E Georgetown, 4.xii. 1 979, RS, JB (MDPI); 6km SE of Mareeba, MDPI, FIT. Site 36. 16. xii. -1511991, 28.iii.-19.iv. 1991, vi.-xi. 1991 . SF (MDPI); 21km E of Marccba, 2111991, RS (CBM, MDPI); Cairns, 3/50, GB (ANIC); Davies Creek, 10/49 (ANIC); Mareeba-Atherton Rd, 9.vii.65 (ANIC); Cooktown, 1/71, GB (ANIC); 11.45S, 142.35E, 1 leathlands. 24-281. 1993, PZ (ANIC); Bamaga. 28.iii.64. IC, MU (ANIC); Cardstone, 111.1966, HK (ANIC); 17.27S, 145.29E. nr The Crater, 18km N of Ravenshoc, 28-29.xi.1981, BJ (ANIC). - NT: Kakadu NP, Cooinda. 22.-25.iii. 1993, LT (CBM); 12.52S, 132.50E, Koongarra 6-10.iii.73, MU (ANIC). - WA: 14.45S, 125.47E, 10km NW by N of Mining Camp, Mitchell Plateau, 1 1-17.V.83, IN, CJ, DR, BJ (ANIC). According to Moore et al. (1987) and T. Weir (pers. comm.) the types are in very poor condition. As the species is easily distinguished by its external morphology, borrowing the fragile remnants for examination was not considered necessary. DIAGNOSIS. Head short, wide, with very large, strongly protruding eyes, with a distinct sulcus and ridge inside of eyes; prothorax elongate, with sharp, sinuate lateral margin; elytra narrow, parallel, dorsally convex, deeply striate at base, with a conspicuous transverse impression in basal third, completely glabrous in apical two thirds. FIG. 3. Basistichus micans (Macleay). Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). 142 MEMOIRS OF THE QUEENSLAND MUSEUM SUPPLEMENTARY DESCRIPTION. Male Genitalia (Fig. 3). Terminal abdominal stemite incised in middle. Genital ring fairly elongate, gently asymmetric, with short basal plate, shortly narrowed to the rather narrow, obtuse apex. Aedeagus rather slender and elongate, laterally little sinuate, lower surface only near base concave, in apical half almost straight. Apex moderately short, very gently knobbed, straight, gently turned to right, and with a very slight notch at the right side. Folding of internal sac very simple. Parameres of fairly dissimilar shape, left paramere much larger than right one and with straight or even slightly concave upper margin, right one short, with obtusely angulate apex. Female Genitalia (Fig. 30E). Stylomere 2 comparatively elongate, laterally moderately curved, with acute apex. With 2 rather narrow, elongate ventrolateral ensiform setae, a rather large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 6-7 moderately elongate ensiform setae. Lateral plate with moderately dense fringe of fairly elongate, stiff nematiform setae at apical rim. DISTRIBUTION. E Qld, N NT, N WA; also in New Guinea. Most labeled specimens were collected at light, some at a considerable distance from water. A small series was collected on surface of ground. Probably a litter-inhabiting species. Clarencia Sloane, 1917 Clarencia Sloane. 1917: 413; Sloane, 1923: 30; Csiki, 1932: 1535; Liebke, 1938: 81; Darlington, 1968: 209; Moore ct al., 1987: 276; Lorenz, 1998: 420. TYPE SPECIES. Casnonia aliena Pascoe, 1860, by original designation. DIAGNOSIS. 3rd antennomere very elongate. Head triangular; prothorax elongate; elytra elongate, dorsally depressed, with a large, light spot in apical half. Clarencia aliena (Pascoe, 1860) (Figs 4, 30F) Casnonia aliena Pascoe, 1860: 39; Chaudoir, 1872: 405; Sloane, 1 890: 644. Clarencia aliena , Sloane. 1917: 413; Csiki, 1932: 1536; Liebke, 1938: 82; Moore et al., 1987: 276; Lorenz, 1998: 420. Casnonia australis Chaudoir. 1862: 277; Moore ct al., 1987: 276; Lorenz, 1998: 420. Casnonia clarensii Castelnau, 1867: 14; 1868: 100; Chaudoir, 1872: 405; Moore et al., 1987: 276; Lorenz, 1998: 420. MATERIAL. HOLOTYPE: 9, Typ dCasnonia aliena Pascoe Type/Moreton Bay/Pascoe Coll J Casnonia aliena Pasc. (BMNH). Syntypes of australis : 9, Australis Chaud/Ex Musaeo Chaudoii/aliena Pascoe Australie, Melbourne (MNHP). 1 9, Ex Musaeo Chaudoir/t///™; Pascoe Australie, Melbourne (MNHP). Syntypes of clarensii : <3, Clarence River Coll. Caste 1 na u H o 1 otypus Casnonia (?) clarensii Castelnau, 1867 (MCSN). NEW RECORDS (14 EX.): Narrabeen. 29.xii.1983, GH (HNMB); Bateman’s Bay, 22.L1963, IC. MU (AN1C); NSW (ANIC). Kilcoy, 10.iv.30 (ANIC); Cooloolabbin Dam, 1 0km W Yandina, l.ii.1997. DB (ANIC); 25 mis. W of Tullv, 8.iii.l964, IC & MU (ANIC); 40km W Tully, 3 1 .v. 1 97 1 . ER (ANIC); Rockhampton, 20.ii. 1 942, CV, ES (QM); 22 D 02’S 148°03’E, Moranbah, 3km S Bendee, 24.-25.iii.2000, GM & SM (CBM. QM); Qld3, L. Broadwater, 35km SSW Dalby, 16.-1 7.xii. 1998, MB (CBM); Casnonia aliena Pascoe (MV). DISCUSSION. This species was described by Chaudoir (1862) as Casnonia australis and by Castelnau (1867) as C. clarensii. Chaudoir ( 1 872) recognised that both names refer to the same species. Sloane (1890), also synonymised C. angusticollis Macleay, 1888, and was followed by all later authors. Examination of the types and comparison of material from SE Qld and NE NT reveals that Macleay ’s name refers to a separate species. It is uncertain whether the Melbourne type locality of C. australis is correct as no modern material is from that far south. As the determination label of the ‘type’ specimen of C. aliena in BMNH was written by Pascoe, this specimen is accepted as the holotype. The single original specimen of C. clarensii bears a printed label holotype that was attached to the specimen recently. DIAGNOSIS. Head long; prothorax long, with almost complete transverse striolation; elytra with more deeply excised, but not decidedly quadridentate apex; aedeagus large relative to body size. SUPPLEMENTARY DESCRIPTION. Measure- ments (Table 2). Male Genitalia (Fig. 4). Very large in comparison to other species of this genus. Genital ring elongate, fairly parallel, slightly asymmetric, shortly narrowed to the wide, obtuse apex. Aedeagus slender and elongate, laterally barely sinuate, lower surface evenly but very gently concave. Apex short, slightly knobbed, suddenly turned to right, and with a distinct notch at the right side. Folding of internal sac very simple. Parameres of similar shape, though left paramere much larger than right one, and with a less sclerotised area along upper margin. AUSTRALIAN ODACANTHINE GROUND BEETLES 143 Female Genitalia (Fig. 30F). Stylomere 2 comparatively elongate, laterally evenly curved, with acute apex. With 3 elongate ventrolateral ensiform setae, a rather large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 6-8 elongate ensiform setae that are characteristically curved towards apex. Lateral plate with dense fringe of stiff, elongate setae at apical rim. DISTRIBUTION. Coastal eastern Australia from Cairns to S NSW, and perhaps Vic. COLLECTING CIRCUMSTANCES. Specimens collected by G. Monteith and myself were at light, the latter near a reed-bordered lagoon. On the basis of body shape, this should be a hygro- philous, probably reed-inhabiting, species. Clarencia angusticollis (Macleay, 1888) (Figs 5, 30G) Casnonia angusticollis Maclcay, 1888: 446; Sloane 1890: 644. Clarencia angusticollis, Moore et al., 1987: 277; Lorenz, 1998:420. MATERIAL. LECTOTYPE (here designated): 9 (?, damaged). N WA/SYNTYPE/Cos«o«/Vj angusticollis Macl., Kings Sound N WA (ANIC-MMS). PARALECTOTYPES: 2 (sex?, very damaged), N WA (ANIC-MMS). NEW RECORDS (1 1 ex.): Momington 1. Mission, 12.v. 1963, PA & NT (SAM); St Margarets Ck, S of T’vill dClarencia aliena (Pascoe) det. R.I. Storey 1988 (MDPI); Normanton, 3.v. & 5.v. 1963, PA & NT (SAM): Cape Bedford/ Clarencia aliena Chd. 20 1 . Andr. (MNHB). Goose Lagoon, 16.1 OS 136.15E, 11km SW by S of Borroloola, 17.iv.1976, JF (ANIC, CBM). FIG. 5. Clarencia angusticollis (Macleay). Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). DIAGNOSIS. This species differs from C. aliena (Pascoe) in its much shorter, more oval-shaped and far less rugose pronotum, the shorter basal part of the head, and the much smaller and slenderer aedeagus. Hence, synonymy with C. aliena is unjustified and the name is reinstated. From more similar C. breviceps sp. nov. it differs by longer prothorax, more deeply excised apex of elytra, and narrower aedeagus with almost straight lower surface. For better comparison, measurements and ratios are given below and the male genitalia are described and figured for the first time. SUPPLEMENTARY DESCRIPTION.Afeawe- ments (Table 2). Male Genitalia (Fig. 5). Much smaller in comparison to C. aliena. Genital ring fairly elongate, barely asymmetric, shortly narrowed to the rather narrow, obtuse apex. Aedeagus very slender and elongate, laterally barely sinuate, lower surface only near base concave, in apical half almost straight. Apex moderately short, gently knobbed, slightly turned up, turned to right, and with a slight notch at the right side. Folding of internal sac very simple. Parameres of fairly dissimilar shape, left paramere much larger than right one and with almost transverse apex, right one with obtusely angulate apex. Female Genitalia (Fig. 30G). Stylomere 2 comparatively elongate, laterally evenly curved, with acute apex. With 4 elongate ventrolateral ensiform setae, a rather large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 8-9 elongate ensiform setae that 144 MEMOIRS OF THE QUEENSLAND MUSEUM are characteristically curved towards apex. Lateral plate with dense fringe of stiff setae at apical rim. DISTRIBUTION. Far N Australia from N Qld to the Kimberley, WA. COLLECTING CIRCUMSTANCES. According to labels, collected at light. Its body form indicates a hygrophilous, reed-inhabiting species. Clarencia breviceps sp. nov. (Figs 6, 30H, 33F) ETYMOLOGY. Refers to the relatively short head. TYPE MATERIAL. HOLOTYPE: 6, 12.36S 132.52E Magela Creek. NT 1km NNW of Mudginbarrv HS. 25.V.73, Matthews & Upton (ANIC). PARATYPES: 1 d, 1 9, same data (ANIC, CBM); 1 1 .52; prothorax almost completely transversely striolate; apex of elytra more deeply excised, external angles considerably projecting aliena (Pascoe) Head shorter and wider, ratio length/width of head < 1 .49; prothorax barely or only near base transversely striolate; apex of elytra less excised, external angles little projecting 3 3. Prothorax slightly shorter as a rule, ratio length/width 1.52-1.59; apex of elytra more excised, outer apical angles more advanced angusticollis (Macleay) Prothorax slightly longer as a rule, ratio length/width 1 .58-1 .63; apex of elytra less excised, outer apical angles less advanced breviceps sp. nov. Deipyrus Liebke, 1938 Deipyrus Liebke, 1938: 104; Csiki, 1932: 1542; Moore et al., 1987: 276; Lorenz, 1998: 420. TYPE SPECIES. Lachnothorax palustris Sloane, 1910, by original designation. DIAGNOSIS. Easily distinguished by combination of pilose surface, complete elytral striation, very distinct ‘neck’, conspicuous lateral sulcus and ridge on prothorax, presence of a yellow spot right at apex of either elytron, and presence of 2 dorsal nematiform setae on female stylomere 2. NOTE: Although the genus was described by Liebke in 1938, the name had already been used by Csiki (1932) in his catalogue, but without giving the citation. Obviously, Liebke must have used this name before, either in printed form, or in litteris. Deipyrus palustris (Sloane, 1910) (Figs 7, 301, 34B) Lachnothorax palustris Sloane, 1910: 396; 1917: 414. Myrmecodemus palustris , Sloane, 1923: 33. Deipyrus palustris, Liebke, 1938: 105; Csiki, 1932: 1542; Moore et al., 1987: 276; Lorenz, 1998: 420. MATERIAL. LECTOTYPE (here designated): 2, Caims/Cotype/1 1556 Lachnothorax palustris Sin., Queensland, Cotype (SAM). PARALECTOTYPES. 1 d, 1 2, Cairns distr., E. Allen/Cotype (SAM). NEW RECORDS (24 ex.): Qld: Cairns dist. E. Allen/ Lachnothorax palustris Shi. Id. by A.M. Lea (MV): Janie Ck. S ofMapoon Mission, Wei, 30.xi.1983, WH (CBM); KurandaColl. Lemoult (IRSBN); Stewart R. i.-ii. 1927, N1 (SAM); Cairns /Lachnothorax palustris Sin. (SAM); Caims/French Coll. Kraatz (DEI). - NT: Darwin, HG (MV); ? (unreadable) Saltpan Roper R. Coll. ? FIG 7. Deipyrus palustris (Sloane). Male genitalia: aedeagus, paramercs and genital ring (scale 0.25mm). (unreadable)/ Lachnothorax palustris SI .( 1 9 1 0) Id. by T.G Sloane (MV); Groote Eylandt, NT (SAM); Adelaide R., HB (SAM); Roper R., NT (SAM); Roper River, 3.iii.l6 (DEI). DIAGNOSIS. Like D. inops sp. nov. but: Legs reddish; antenna reddish; eyes convex, laterally projecting, composed of glossy ommatidia of about equal size. SUPPLEMENTARY DESCRIPTION. Measurements (Table 3). Male Genitalia (Fig. 7). Terminal abdominal stemite in middle incised. Genital ring fairly elongate, triangular, almost symmetric, with convex basal plate, regularly narrowed to the narrow, triangular apex. Aedeagus slender and elongate, laterally sinuate, lower surface in basal half gently concave, in apical even slightly convex. Orificium very elongate. Apex rather wide, moderately short, very gently knobbed, straight, gently turned to right, and with a short, distinct notch at the right side. Folding of internal sac very simple. Parameres short, of fairly dissimilar shape, left paramere much larger than right one and with convex apex, right one short, with obtusely angulate apex. Female Genitalia (Fig. 301). Stylomere 2 large, though comparatively short and stout, laterally moderately curved, with fairly acute apex. With 2 small ventrolateral ensiform setae, a medium-sized dorsomedian ensiform seta situated about in middle of stylomere, and 2 short nematiform setae arising from a groove in apical third. Base of stylomere 1 with 6-7 elongate, slender ensiform setae. Lateral plate with dense fringe of elongate, stiff nematiform setae at apical rim. AUSTRALIAN ODACANTHINE GROUND BEETLES 147 DISTRIBUTION. NE Qld, N NT. COLLECTING CIRCUMSTANCES. Little known. One specimen was collected on a salt pan, and since Moore ct al. (1987) also record it as living on ‘salt marsh’, this may be a halophile species, generally or facultatively. Deipyrus inops sp. nov. (Fig. 30J, 34C) ETYMOLOGY. Refers to the small, remarkably depressed eyes. MATERIAL. HOLOTYPE: 9, Australia C. Oke Collection/ \1yrnwc()demus sp. near palustris SI. det. B.P. Moore (MV). PARATYPE: 9 , same data (CBM). DIAGNOSIS. Like D. palustris (Sloane) but: legs dark; antenna dark; eyes remarkably depressed, laterally not at all projecting, composed of conspicuously rough ommatidia of very unequal size. DESCRIPTION. Measurements (Table 3). Colour (Fig. 34C). Surface, including mouthparts, antennae and legs black. Apex of elytra with an inconspicuous, ill defined reddish-orange spot that extends from about 3rd interval to external apical margin and broadly meets the apex. Head . Rather narrow, remarkably oval. Neck very narrow, with deep transverse impression. Eyes of moderate size, though absolutely depressed and laterally not protruding, not separated from orbits which are elongate and rather convex. Colour of eye characteristically dark grey in middle, with a slightly lighter margin. Surface of eyes remarkably rugose, in particular in middle where the ommatidia are conspicuously smaller than at margin. Medially of eye with a fairly deep sulcus that extends to behind middle of eye, but without ridge. Posterior supraorbital seta located well behind posterior margin of eye, but seta difficult to detect within the elongate pilosity of surface. Clypeus on either side with 2 additional setae behind the anterior one. Mentum with elongate, acute, triangular tooth, with 2 setae behind tooth, submentum with several elongate setae on either side. Apex of glossa transverse, with 2 elongate median and 2 shorter lateral setae. Paraglossae free, narrow, surpassing glossy. Lacinia elongate, interior margin with a dense fringe of spines. Antenna moderately elongate, almost attaining base of pronotum, pilose from 1st antennomere. Surface glossy, without microreticulation, though rather dense and coarsely punctate, with very elongate, hirsute pilosity that is inclined anteriorly. Also lateral margin and lower surface of head pilose. Pronotum. Short and very convex, almost hemispherical, near base with shallow transverse impression. Apex not bordered, lateral margin complete, slightly ridge-like, convex, with distinct, rather wide sulcus medially. Base with thick border. Median line shallow. Marginal seta(e) not visible within the elongate pilosity. Proepistemum broadly visible from above. Disk rather densely and coarsely punctate, near apex and base punctuation dense and rugose. Surface without microreticulation, glossy, with very elongate, hirsute, erect pilosity. Elytra . Rather short and wide, somewhat rectangular, laterally gently convex, barely widened in apical half, surface gently convex. Base gently oblique, humeri fairly distinct but rounded, lateral evenly convex and rounded towards sutural angle without any indication of lateral apical angles. Apex, therefore, remarkably convex. Base margined towards position of 4th stria, apex very narrowly margined. Surface in anterior third in middle with very shallow, transverse impression. All striae complete, running from base to apex. Striae slightly impressed, coarsely punctate in anterior halt, punctures becoming inconspicuous posteriorly, in apical half striae only linear. Intervals almost depressed, with rather dense, irregular, about biseriate punctuation. Scutellar stria elongate, consisting of about 8 large punctures. No fixed setae visible within the elongate pilosity. Surface without microreticulation, glossy, but with dense, elongate, hirsute, pilosity that is inclined posteriorly. Hind wings present. Lower Surface. Thorax densely and coarsely punctate and with elongate, hirsute pilosity. Abdomen with finer punctures and shorter, slightly denser pilosity. Metepisternum moderately elongate, c. 2 x as long as wide at apex. Terminal abdominal sternum in female tri- or quadrisetose on either side. Legs. Of moderate size. Tarsi not lobed, pilose on upper surface, 5th tarsomere with a dense fringe of elongate setae below. Claws large, smooth. Squamosity of male anterior tarsus unknown. Male Genitalia . Unknown. Female Genitalia (Fig. 30J). Stylomere 2 large, though comparatively short and stout, laterally moderately curved, with fairly acute apex. With 2 small ventrolateral ensiform setae, a medium- sized dorsomedian ensiform seta situated about 148 MEMOIRS OF THE QUEENSLAND MUSEUM in middle of stylomere, and 2 short nematiform setae arising from a groove in apical third. Base of stylomere 1 with 6 elongate, slender ensiform setae. Lateral plate with dense fringe of elongate, stiff nematiform setae at apical rim. Variation. Due to limited material very little variation noted. DISTRIBUTION. ‘Australia’. KEY TO AUSTRALIAN SPECIES OF DEIPYRUS 1 . Eyes convex, laterally well projecting; antennae and legs reddish to light brown, tibiae with yellow median ring. . palustris (Sloane) Eyes depressed, not projecting; antennae and legs uniformly black inops sp. nov. Dicraspeda Chaudoir, 1862 Dicraspeda Chaudoir, 1862: 300; Sloane, 1923: 30; Csiki, 1932: 1536; Liebke, 1938: 88; Darlington, 1968: 210; Moore et al., 1987: 274; Bachr. 1996a: 138; 1997b: 30: 1998: 174; 1999: 116; 2000: 11; 2003b: 101; 2003c: 251; Lorenz, 1998: 420. TYPE SPECIES. Dicraspeda brunnea Chaudoir, 1 862, by monotypy. DIAGNOSIS. Distinct sulcus and ridge inside the eye; distinct sulcus inside the prothoracic margin; elytra impilose (except for fixed setae at 3rd interval), with elytral apex slightly excised and not bearing angulate or even spinose external angles. DISCUSSION. For a time this genus was conftised with the related Eudalia , and even Sloane ( 1 9 17. 1 923) was not sure to which genus the quite differently shaped Australian species should be referred. Dicraspeda obscura (Castelnau). for example, was referred to Arame Andrewes by Sloane (1923). Those species that are today combined to form Dicraspeda , are remarkably different in their external shape and structure. When considering the species that occur in New Guinea, the problem becomes even more difficult, because the former genera Philemonia Liebke and Macrocentra Chaudoir have to be taken into consideration. Today these are included in Dicraspeda sensu lato , but certainly they again deviate in shape and structure. Some of these problems are discussed by Baehr (1996a, 1997b, 1998, 1999, 2000 2003b, 2003c). Many species that have described from New Guinea fall in the former ‘ Philemonia ’ (Baehr, 1996a, 1997b, 1998), some new Australian records of New Guinean species (Baehr, 2000), and the brunnea-g roup’ of species was revised (Baehr, 2003c). No further taxonomic information about the species of the latter group will be given. DISTRIBUTION. E and N Australia, New Guinea, Bismarck Archipelago, Solomon Islands, New Hebrides, Moluccan Islands, Greater and Lesser Sunda Islands, Philippines. Dicraspeda brunnea Chaudoir, 1862 Dicraspeda brunnea Chaudoir, 1862: 300; Sloane, 1923: 31, Csiki. 1932: 1536; Liebke, 1938: 89; Darlington, 1968: 211; Moore et al 1987: 274; Baehr, 1996a: 138; 1998: 174; 2003c: 251; Lorenz, 1998: 420. DISCUSSION. This species was described from Sulawesi and is widely distributed from southernmost Thailand and Indonesia through Sulawesi and the Philippines (Baehr, 1998, 2003c). It was recorded from New Guinea by Darlington (1968) and from N Australia by Moore et al. (1987). Baehr (2003c) noted that these records are probably erroneous, because in New Guinea certain species occur that are different from brunnea , and moreover, I have not seen brunnea from New Guinea or Australia. For New Guinea, Darlington’s records probably refer to either D. nigripes Baehr, D. obsoleta Baehr or D. papuensis Baehr. For Australia, Moore et al’s ( 1 987) records probably refer either to D. sublaevis (Macleay) that was synonymised with D. brunnea by Sloane (1923) but reinstated by Baehr (2003c), or D. nitida Sloane or D. glabrata Baehr. Therefore, D. brunnea probably does not belong to the Australian fauna. Dicraspeda sublaevis (Macleay, 1888) (Fig. 34D) Eudalia sublaevis Macleay, 1888: 448; Sloane, 1917: 418. Dicraspeda sublaevis (as D. brunnea Chaudoir), Sloane, 1923: 31; Csiki, 1932: 1537; Moore et al., 1987: 274; Lorenz, 1998: 420. Dicraspeda sublaevis , Baehr, 2003c: 253. NEW RECORDS (37 ex.): QLD: 10km S Georgetown, 30.xii.1979, RS, JB/ Dicraspeda brunnea Chaudoir Det. R.I. Storey 1988 (CBM. MDPI); Elizabeth Ck., Wrotham Park Stn., via Chillagoe, 6.xii.l990, DG (MDPI); Sellheim, xii/42. )B! Dicraspeda brunnea C’hd. 1493. (ANIC); Mary Creek, 16.33°S 145.12°E, 4-5.xii.1968, BM (ANIC, CBM); 13.58°S 143.1 1 °E, Mt White, 12.1.1994. PZ, EE (ANIC); Mt Carbine, 511964, GM (QM0. - NT: Humpty Doo, 6km E, 9.ii.-4.iii.l987, RS, GB (CBM, MDPI. QM); 12.52S 132.50E, Koongarra, 6-10.iii.93, MU (ANIC); Katherine env. 10.- 1412004, PM (CBM); Horn Islet., Pellcw Group, 1 5-3 l.i. 1968, 1 5-2 l.ii. 1968, BC7 Dicraspeda sublaevis Macl. det. B.P. Moore 1974 (CBM, QM). - WA: Kununnurra, 22.xii.91 -611992, RS (CBM, MDPI). Most specimens were collected at light. AUSTRALIAN ODACANTHINE GROUND BEETLES 149 DISCUSSION. Although synonymised with D. brunnea for a long period, Baehr (2003c) demon- strated that it is a well characterised, separate species. DISTRIBUTION. The new records extend the range into N Qld, but only to the western slope of Great Dividing Range. Dicraspeda nitida (Sloane, 1917) Dicraspeda glabrata Baehr, 2003c Dicraspeda glabrata Baehr, 2003c: 255. NEW MATERIAL. None. Dicraspeda brunneipennis (Sloane, 1917) (Figs 8, 30K) Eudalia brunneipennis Sloane. 1917: 420. Dicraspeda brunneipennis Csiki, 1932: 1537; Liebke, 1938: 89; Moore et al., 1987: 275; Lorenz, 1998: 420. Eudalia nitida Sloane, 1917: 420 Dicraspeda nitida , Csiki, 1932: 1537; Moore et al, 1987: 275; Lorenz, 1998: 420; Baehr, 2003c: 253. NEW RECORDS (67 ex ): OLD: Mt Lewis, via Julatten, 29.xii.1979, RS, N GIDi c rasped a nitida SI. det. B.P. Moore‘79 (MDPI); Cape Tribulation, Daintree area, 15-16.xii.1978, RSI Dicraspeda nitida SI. det. B.P. Moore' 86 (MDPI); Cape Tribulation, 24.-29.xii. 1980, RS, NG (MDPI); Cow Bay, N of Daintree, 25.i.-7.ii.l984, CC (MDPI); 15km WSW of South Johnstone, 19.1.1986, JH (MDPI); 45km NE of Cooktown, 23.xii.1979, RS (MDPI): Bloomfield Range via Cooktown, 24.xii.1979, RS (MDPI); Cape Flattery, 23.-28.xii.1989, RS (CBM, MDPI); Cardstone, 4-1 6.i. 1966. KH (ANIC); 15.47°S 145.14°E, Shiptons Flat, 1 7- 1 9.x. 1 980. TW (ANIC); 16.03S to 16.05°S 145.28°E, Cape tribulation area, 21-28.iii.1984, AC, TW(ANIC); 16.19°S 145.24°E, 12km S of Daintree, 27.xi.1981, BJ (ANIC): 15,04°S 145.07°E, Mt Webb N.P., 27-30.iv.1981, AC, JF (ANIC): 15.29°S 145.16°E, Mt Cook N.P., 10-12.V.1981, AC, JF (ANIC): Cairns. ii.50,JB (ANIC); 12.43°S 143.18°E, 1 lkmENEof Mt Tozer, ll-16.vii.1986, TW, AC (ANIC); Shute Harbour, 3.iii.64, 23.iv.64, 1C, MS (ANIC); Barron R., Cairns, 1 5.i. 1993, BU (ANIC); Lake Placid disL3.ii. 1995, BU (ANIC); 15.28°S 1 45. 1 5°E. Cooktown, Walker's Bay, 2911995, LR (ANIC); 11.45°S 142.35°E, Heathlands, 2211992, TW, IN (ANIC. CBM): 11 41°S 142.42°E, 14km ENE Heathlands, 8.xii.l992. PZ & WD (ANIC, CBM); 1 6.03°S to 16.08°S 145.28°E, Cape Tribulation area, l-ll.v.1992, JL (ANIC): 13.58°S 143.11°E, Mt White, 12. i. 1994, PZ, EE (ANIC); Rossville env. 25.-27.xii.2003, PM (CBM); Green Hill, Thursdav I., 21.5.2003, GM/I0°35'S 142°13'E, 80m (QMB); Horn I. 2.5km W of Horned Hill, 19.5.2003, GM/10°36'S 142°18’E, 50m (CBM, QMB); Hammond I., 1 km W village, 20.5.2003, GB/10°35’S 142°13’E, 50m (CBM, QMB); Nelly Bay, Magnetic I. 12.1997, SF (QMB). DISTRIBUTION. Widely distributed in NE Qld, E of Great Dividing Range, including S Torres Strait Islands. Most specimens collected at light, a few ‘in rainforest', some in ‘gallery forest litter’ and ‘vine scrub litter’. MATERIAL. LECTOTYPE (here designated): <3, Cairns (K) Dodd 04 5 /Eudalia brunneipennis SI from Kuranda. Cotype/H OLOTYTE £, brunneipennis SI. PJD (ANIC). NEW RECORDS (6 ex.): Qld: Noah Creek, 16°08’S 145°25’E, 27.vii.1993, HM, RK (CRC); Bellenden Kcr Range, Cableway Base Stn, 100m, 17.x.-9.xi.l981, EW (CBM, QM); Cape Tribulation, Daintree area, 15-16.xii.1978, RS /Dicraspeda sp. det. B.P. Moore 1979 (MDPI). DIAGNOSIS. Distinguished from those species bearing a narrow marginal pronotal sulcus and rather convex elytra by prothorax impunctate, elytra non-microreticulate, with barely excised apical margin and always distinctly lighter than the fore body; 4th tarsomeres not deeply excised. SUPPLEMENTARY DESCRIPTION. Male Genitalia (Fig. 8). Terminal abdominal stemite in middle incised. Genital ring comparatively wide, rather triangular, barely asymmetric, with short, acute, triangular apex. Aedeagus small, elongate, fairly depressed, laterally moderately sinuate, lower surface very gently concave. Orilicum moderately elongate. Apex short, wide, almost straight, slightly knobbed and spoon-shaped, moderately turned to right, with shallow incisions at both sides. Folding of internal sac rather simple. Parameres of moderately dissimilar shape, left paramere much larger than right one, both with wide, obtusely rounded apex. Female Genitalia (Fig. 30K). Stylomerc 2 comparatively elongate, laterally evenly curved, with acute apex, with 3 large ventrolateral ensiform setae, a large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomerc 1 with c. 6 ensiform setae of decreasing size, the median ones longest and conspicuously curved. Lateral plate with dense fringe of ensiform setae at apical rim. TABLE 3. Measurements and ratios of both Australian species ot Deipyrus. N length (mm) length eye/orbit 1/w head 1/w prothorax l/w elytra inops 2 8. 3-8. 5 0.71-0.73 1.58-1.59 1.14-1.16 1.47-1.48 palustris 6 7. 3-8.4 0.59-0.67 1.36-1.43 1.14-1.21 1.53-1.58 | DISTRIBUTION. Wet Tropics of NE Qld. The Bellenden Ker and Noah Creek specimens were collected by insecticide fogging in rain forest. 150 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 8. Dicraspeda brunneipennis (Sloane). Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). Dicraspeda dubia (Gestro, 1879) Odacantha dubia Gestro, 1879: 558. Philemonia dubia , Csiki, 1932: 1536 ; Liebke, 1938: 83. Dicraspeda dubia , Darlington, 1968: 212; Moore ct al.. 1987: 275; Baehr, 1996a: 138; 1997b: 30; 1998: 174; 2000: 11; Lorenz, 1998: 420. DIAGNOSIS. Distinguished from all species bearing a narrow marginal pronotal sulcus and convex elytra by impunctate prothorax, microreticulate elytra with deeply excised apical margin, and not deeply excised 4th tarsomeres. DISTRIBUTION. This New Guinean species was recorded in N Qld (Bamaga) by Darlington ( 1 968), a record repeated by Moore et al. (1 987). I have not seen D. dubia from Australia and since the related D. longiloba (Liebke), is recorded from N Qld (Baehr, 2000) I am not sure whether the D. dubia record is genuine. Dicraspeda longiloba (Liebke, 1938) Philemonia longiloba Liebke, 1938: 83. Dicraspeda longiloba , Darlington, 1968: 212; Lorenz, 1998: 420; Baehr, 2000: 11. DIAGNOSIS. 4th tarsomeres very deeply excised. DISTRIBUTION. New Guinea and Australia (Baehr, 2000). Dicraspeda obscura (Castelnau, 1867) (Figs 9, 30L, 34E) Casnonia obscura Castelnau, 1867: 14; 1868: 100; Chaudoir, 1872: 407; Gestro, 1875: 851. Eudalia obscura , Sloane, 1917: 418. Arame obscura, Sloane, 1923: 31. Dicraspeda obscura , Csiki, 1932: 1537; Liebke, 1938: 89; Moore et al., 1987: 275; Lorenz, 1998: 420. MATERIAL. LECTOTYPE (here designated): d, Rockhampton Coll. Castelnau/Esempl. tipico Coll. Castelnau/ofocwra CasUCasn. obscura Cast. (Castelnau ’s handwriting)/SYNTYPUS Casnonia obscura Castelnau, 1867 (MCSN). PARALECTOTYPES: I d, 2 9 9, same data (MCSN). NEW RECORDS (77 ex ): Qld: 15km N Marlborough, MB (CBM); 20km N Biggenden, MB (CBM); Cairns, Whitfield Rd, 28.L1974, WH (CBM, CMP); Mt Lewis, 21.U976, WH (CBM. CMP); Moa I. Torres Straits, JS (SAM); Casnonia obscura (Cast.)/Casnonia CS Rockm (MV); Mackay, (MV); Cairns, i.52, JB (MV); Cairns, i. 1956, CO (MV); Casnonia obscura Cast. /Mackay (MV); Mackay (MV); Iron Range, 4.v.l975,mm (MV); Old Bushman Beach, 20km N Townsville, 26-29.ii.1998, AW (SAM); Kuranda/Griftith Collection (SAM); Tolga, i. 1 980, NG, JB /Dicraspeda obscura (Cast.) det. R.I. Storey 1988 (MDPI); Tolga. 7.iii. 1983. 231.1986, JB (MDPI, QM); 7km NE ofTolga. ii. 1 988, RS, DF (MDPI); Morehead R. N of Laura. 201.1990, FH (MDPI); 15km WSW of South Johnstone, 24.xii.1985, FH (MDPI); Walkamin, 8-15.iii.1985, JB (MDPI); Townsville, JT (ANIC); Cairns, iii.51, GB ( ANIC); Archers Ck. iv.74. GB (ANIC); Mt Spec, i.75, GB (ANIC); 3 mi. W of Mossman, 14.iii.64, 1C, MU (ANIC); Eungella N.P., 2400 ft. 2.iii.64, IC, MU (ANIC); Yeppoon, 26-29.xii.64. IC, MU (ANIC); 16.47°S, 145.22°E, 24km N by W of Mareeba, 24-25.xi.1981, BJ (ANIC); Lansdown Station, 19.40°S, 146.51°E, 7km S of Woodstock. 161.74, RB (ANIC). - NT: Goose Lagoon, 16.1 OS, 136.15E, 11 km SW by S of Borrooloola, 17.iv. 1976, JF (ANIC); McArthur River, 16.27°S, 136.05E, 48km SW by S of Borrooloola, 13.iv.1976, JF (ANIC); 12.52S, 132.50°E, Koongarra, 6-10.iii.73, MU (ANIC); 12.47°S, 132.51°E, 19km NE by E of Mt Cahill, 16.xii.1972, MU (ANIC); Ferguson R., 14.19°S, 131.50°E, 25.vi.1968, ME (ANIC); Tindal, 14.31°S, 13222°E, l-20.xii. 1867, WV(ANIC); 16°4rS 135°44'E, Cape Crawford, 17-19.iv.2004,GM, DC, 1 1621 (QMB).- WA: 14.45°S, 125.47°E, 1 0km NW by N of Mining Camp, Mitchell Plateau, ll.v.83, IN, CJ (ANIC); 14.49°S, 125.50°E, Mining Camp. Mitchell Plat. 9-19.V.1983, IN, CJ, DR, BJ ( ANIC); 14.25°S, 126.38°E,CALM Site 13/4, 12km S of Kalumburu Mission, 7-1 l.vi. 1988, TW (ANIC). DIAGNOSIS. Pronotum coarsely punctate. SUPPLEMENTARY DESCRIPTION. Male Genitalia (Fig. 9). Terminal abdominal stemite in middle deeply incised. Genital ring comparatively narrow, elongate, parallel, slightly asymmetric, with short, acute, triangular apex. Aedeagus small, moderately elongate, fairly depressed, laterally moderately sinuate, lower surface very gently concave. Orificum very elongate. Apex short, wide, almost straight, knobbed and slightly spoon-shaped, moderately turned to right, with shallow incisions at both sides. Folding of internal sac rather simple. Parameres of very dissimilar shape, left paramere AUSTRALIAN ODACANTHINE GROUND BEETLES 151 much larger than right one, stout, with wide, obtusely transverse apex, right paramere narrow, with convex apex. Female Genitalia (Fig. 30L). Stylomere 2 comparatively elongate, laterally evenly curved, with acute apex. With 2 large ventrolateral ensiform setae, a large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 5-6 stout ensiform setae of decreasing size. Lateral plate with dense fringe of very elongate stiff setae at apical rim. DISTRIBUTION. E Qld, Torres Strait, N NT, and N WA. Most specimens were collected at light. It is uncertain whether this is an hygrophilous species or leaf litter species independent of water. KEY TO AUSTRALIAN SPECIES OF DICRASPEDA 0.25mm). Eudalia Castelnau, 1867 Eudalia Castelnau, 1867: 16; 1868: 102; Sloane, 1917: 415; 1923: 30; Csiki, 1932: 1542; Darlington, 1968: 214; Moore et al., 1987: 273; Lorenz, 1998: 421; Baehr, 1999: 116; 2003b: 101. 1. Marginal pronotal sulcus wide; elytra short and wide, depressed (Fig. 34D) 2 Marginal pronotal sulcus narrow; elytra longer and narrower, rather convex (Fig. 34E) 4 2. Striae deeply impressed, intervals clearly convex, even near apex sublaevis (Macleay) Striae not or barely impressed, intervals depressed, at leastnearapex 3 3. Surface of elytra in basal third without perceptible transverse impression, with superficial though distinct microreticulation nitida (Sloane) Surface of elytra in basal third with distinct transverse impression, at least in basal half without perceptible microreticulation glabrata Baehr 4. Whole pronotum densely and very coarsely punctate; 4th tarsomeres of anterior and median tarsi barely excised obscura (Castelnau). Pronotum punctate only near base and apex, punctuation rather fine; 4th tarsomeres of anterior and median tarsi considerably excised 5 5. Excision of 4th tarsomeres of all tarsi very deep, > Va of length of 4th tarsomcrc longiloba (Liebke) Excision of 4th tarsomeres less deep, c. Vz of length of 4th tarsomere, in metatarsus excision shallow 6 6. Elytra without traces of microreticulation, apical margin little excised, external apical angles little projecting; head longer and narrower, eyes barely surpassing curvature of orbit; elytra always distinctly lighter than fore body bnmneipennis (Sloane) Elytra with fine microreticulation, apical margin deeply excised, lateral apical angles projecting; head shorter and wider, eyes distinctly surpassing curvature of orbit; elytra little lighter than fore body .... dubia (Gestro) TYPE SPECIES. Odacantha latipennis Macleay, 1 864, by original designation. DIAGNOSIS. Distinct ridge and sulcus medially of eye absent; deep sulcus inside of the marginal border of pronotum absent; full elytral striation; striae punctate; apex of elytra not or barely excised. DISCUSSION. Eudalia seems to be a genus of convenience which includes quite differently shaped and structured species that are combined more by plesiomorphic than by apomorphic characters. The single New Guinean species, E. anomala Darlington, deviates even more and is fairly similar to the Oriental Andre wesia Liebke. Two well separated lineages in Australia are 1) the obliquiceps- lineage that comprises rather elongate, impilose or scarcely pilose species with smaller, less protruding eyes (C. obliquiceps , C. minor , C. castelnaui , C. reticulata , C. at rata, C. femorata ), and 2) the macleayi-Ymeage that comprises short, compact, densely pilose species with large, protruding eyes and short, remarkably convex orbits (£. macleay i, E latipennis , E. punctipennis , E. waterhousei). In some character states E. nigra Sloane differs from Eudalia and resembles Dicraspeda. Hence, it is removed from Eudalia , but due to structural differences between it and all other members of Dicraspeda , it is given the status of a separate genus. 152 MEMOIRS OF THE QUEENSLAND MUSEUM DISTRIBUTION. Apart from the aberrant New Guinean E. anomala , that may merit the erection of a separate genus, Eudalia is confined to Australia. Eudalia obliquiceps Sloane, 1917 (Figs 10-12, 30M, 34F, 35A) Eudalia obliquiceps Sloane, 1917: 4 IS; Csiki, 1932: 1542; Liebke, 1938: 106; Moore etal., 1987: 274; Lorenz, 1998: 421. Dicraspeda obliquiceps , Sloane, 1923: 31. DIAGNOSIS. Colour uniformly black; elytra without microreticulation, with extremely fine, microscopic pilosity; 3rd and 5th interval with setiferous punctures; legs conspicuously bicoloured. DISCUSSION. Moore et al. (1987) record this species from Cairns and the type locality (Laura). New material extends the range through Qld, NT and WA. The material also reveals differences between the type and other specimens from northwestern Qld, NT, northernmost WA, Iron Range in Cape York Peninsula, and specimens attributed to E. obliquiceps from other localities in northern Qld and northern NSW. Hence, specimens from certain localities in northern Qld and northern NSW that differ more substantially, are described as separate species, whereas specimens from Iron Range and those from NT and northwestern Australia, respectively, are described as separate subspecies of E. obliquiceps . This procedure has been chosen, because available representatives of the latter populations are quite uniform in shape and degree of microreticulation, but differ in minor characters of external morphology (degree of punctuation, shape of pronotum), and in shape of male aedeagus. Eudalia obliquiceps obliquiceps Sloane, 1917 (Figs 10, 30M) Eudalia obliquiceps Sloane, 1917: 418. All citations mentioned above under the species refer to the nominate subspecies. MATERIAL. HOLOTYPE: 4 elongate ventrolateral ensiform setae, stylomere 1 with fringe of > 9 extremely elongate setae at apical rim ( Fig. 30N,P) 6 3. Head short and wide with comparatively large eyes; pronotum short and wide (Table 4), lateral margins considerably rounded, conspicuously sinuate near basal angles, dorsal surface rather convex 4 Head narrower with slightly smaller eyes; pronotum narrow and elongate (Table 4), lateral margins almost parallel, not sinuate near basal angles, dorsal surface more depressed 5 4. Body size larger, >7.7mm; pronotum anteriorly more extensively punctate; aedeagus less delicate, apex less upturned, less markedly turned to right side, without a distinct lateral notch; left paramere convex at apex (Fig. 12) obliquiceps punctifrons subsp. nov. Body size smaller, <7.3min; pronotum anteriorly barely punctate; aedeagus delicate, apex decidedly upturned, markedly turned to right side and; left paramere transverse at apex (Fig. 13) mz/iorsp.nov. 5. Head barely punctate; pronotum punctate only in basal half; elytra generally slightly longer (Table 4); aedeagus longer and narrower, lower surface less curved, apex 166 MEMOIRS OF THE QUEENSLAND MUSEUM decidedly upturned, both parameres longer (Fig, 11). obliquiceps tozeria subsp. nov. Head more extensively punctate; pronotum almost completely punctate; elytra generally slightly shorter (sec Tab. 4); aedeagus shorter and stouter, lower surface evenly curved, apex barely upturned, both parameres shorter ( Fig. 10) obliquiceps obliquiceps Sloane 6. Pronotum laterally very- convex, little longer than wide, margin anteriorly with several short setae; 5th stria with > 5 setiferous punctures; elytra with dense, rugose, distinct microreticulation castelnaui Sloane Pronotum laterally little convex, considerably longer than wide, margin anteriorly without additional setae; 5th stria without or with 2-3 setiferous punctures only; elytra with much more superficial microreticulation 7 7. 5th stria with 2-3 setiferous punctures, towards apex striae still perceptibly punctate; microreticulation of elytra in basal half indistinct, less developed than in apical half; eyes slightly larger and more protruding, ratio length of cye/length of orbit c. 1 .45; aedeagus with shorter and wider apex and w ith deep excision on right side behind apex (Fig. 14) reticulata sp. nov. 5th stria without setiferous punctures, towards apex striae barely punctate though distinctly impressed; microrcticulation of elytra in basal half about as distinct as in apical half; eyes slightly smaller and less protruding, ratio length of eye/lcngth of orbit < 1 .30; aedeagus with longer and narrower apex and with shallow' excision on right side behind apex (Fig. 15). . 8 8. Legs uniformly dark; elytra wider, posteriorly distinctly widened, ratio 1/w <1.68; intervals more depressed, barely convex towards apex; striae more coarsely punctate, microreticulation of intervals distinct * . atrata sp. nov. Legs dark but upper surface of femora contrastingly light reddish; elytra narrow'er, almost parallel, ratio 1/w 1 .76; intervals convex throughout; striae less coarsely punctate, microreticulation of intervals more superficial * femorata sp. nov. 9. Apex of elytra more or less widely yellow and micro- reticulation of elytra distinct 10 Elytra unicolourous or when apex indistinctly yellow, microrcticulation of elytra very faint or absent . ... 11 10. Body size larger, length usually > 9mm; eyes slightly smaller (Table 4); pronotum densely and regularly punctate; yellow' apical margin of elytra generally narrower, less produced along lateral margins, and less distinct; microrcticulation distinct, surface dull; lower surface of aedeagus more sinuate, apex less knobbed (Fig. 18) latipennis latipennis (Macleay) Body size smaller, length < 8.5mm; eyes slightly larger (Table 4); pronotum less densely and regularly punctate, anteriorly with some impunctate areas; yellow apical margin of elytra generally wider, more produced along lateral margins, and more distinct; microreticulation superficial, surface glossy; lower surface of aedeagus almost straight, apex markedly knobbed (Fig. 19). latipennis interioris subsp. nov. 11. Elytra completely black with slight greenish tinge; microreticulation of elytra conspicuous, rugose, punctuation of intervals regular, barely transversely confluent; head with impunctate areas in middle ♦ macleavi Bates Elytra blackish or piceous without greenish tinge, sometimes with inconspicuous, narrow, ill defined light apex; microrcticulation of elytra faint or absent, punctuation of intervals more irregular, at least in parts transversely confluent; head with dense and regular punctuation, without impunctate areas 12 12. Intervals with conspicuously confluent punctures which form irregular transverse sulci (Fig. 39A); micro- rcticulation of elytra extremely superficial or completely absent; aedeagus short and stout, with short, wide apex (Fig. 20) waterhousei Castelnau Punctuation of intervals not much confluent, elytra much more regularly punctate; microrcticulation of elytra superficial though distinct (Fig. 38F); aedeagus longer and more delicate, with longer, narrower apex (Fig. 21 ). punctipennis sp. nov. Gestroania Liebke, 1938 Gestroania Liebke, 1938: 89; Csiki, 1932: 1537; Moore et al., 1987: 275; Lorenz, 1998: 420. TYPE SPECIES. Casnonia amplipennis Gestro, 1875, by monotypy. DIAGNOSIS. Antenna elongate; elytra ampliate, depressed, with sharply angulate or even spinose external apices; 3rd, 5th, and 7th intervals setose; pronotum elongate, sharply margined, dorsally very rugosely punctuate. Pronotum. DISCUSSION. Although the genus was described by Liebke in 1938, the name had already been used by Csiki (1932) in his catalogue, but without mentioning the citation. The genus is similar to Giachinoana Baehr, but is distinguished from the latter by absence of the frontal furrow and ridge near the eye, and by odd intervals not definitely carinate. Gestroania amplipennis (Gestro 1875) (Figs 3 IE, 36D) Casnonia amplipennis Gestro, 1875: 853. Eudalia amplipennis Sloane, 1917: 413. Gestroania amplipennis Liebke, 1938: 89; Csiki, 1932: 1537; Moore et al., 1987: 275; Lorenz, 1998: 420. MATERIAL. HOLOTYPE: 9, Swan River Coll. CastcUwu/Typus/ amplipennis Gestro/HOLOTYPE Casnonia amplipennis Gestro, \815/Gestroania amplipennis Gestro Det. M. Liebke. Hamburg (MCSN). New records (1 ex.): WA: Nicol Bay/Ex Musaeo H.W. Bates, 1892 (MNHP). DIAGNOSIS. Distinguished from GJ'roggatti by conspicuously bicoloured femora, dark antenna, and more coarsely crenulate elytral striae; and from G. setipennis and G. storeyi by the lower number of setiferous punctures on the odd intervals. AUSTRALIAN ODACANTHINE GROUND BEETLES 167 SUPPLEMENTARY DESCRIPTION. Measurements (Table 5). Male Genitalia. Unknown. Female Genitalia (Fig. 3 IE). Stylomere 2 moderately elongate, laterally moderately curved, with acute apex. With 2 medium-sized ventrolateral ensiform setae, a arge dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 7 moderately elongate ensiform setae. Lateral plate with fairly dense fringe of moderately elongate, stiff nematiform setae at apical rim. DISTRIBUTION. Southern half of WA from Nicol Bay to Perth. Gestroania froggatti (Macleay, 1888) (Figs 23, 3 IF, 36E) Eudalia froggatti Macleay, 1888: 448: Sloane, 1917: 417; Csiki, 1932: 1542; Liebke, 1938: 106; Moore etaL 1987: 274; Lorenz, 1998:421. Dicraspedci froggatti, Sloane, 1923: 3 1 . MATERIAL. LECTOTYPE (here designated): ? sex (damaged), NW Austr./SYNTYPE/£wt/tf//u froggatti , Macl. Barrior Range N WA. (ANIC-MMS). New records (2 ex.): WA: Kununurra, 22.xii. 1991 -511992, RS (CBM, MDPI). DIAGNOSIS. Distinguished from G. ampli- pennis (Gestro) by completely dark femora, yellow antenna, and less coarsely crenulate elytral striae; and from both, G setipennis sp. nov. and G storevi sp. nov. by the lower number of setiferous punctures on the odd intervals. SUPPLEMENTARY DESCRIPTION. Measurements (Table 5). Male Genitalia (Fig. 23). Terminal abdominal sternite in middle incised. Genital ring moderately wide, triangular, barely asymmetric, with narrow, triangular apex. Aedeagus slender and elongate, moderately depressed, laterally little sinuate, lower surface concave near base, almost straight in apical half. Orificum short. Apex short, wide, very slightly upturned and knobbed, slightly turned to right, barely incised at right side. Folding of internal sac simple. Parameres of moderately dissimilar shape, left paramere much larger than right one, left paramere stout with almost transverse apex, right paramere with narrow apex. Female Genitalia (Fig. 3 IF). Stylomere 2 moderately elongate, laterally moderately curved, with acute apex. With 3 medium-sized FIG. 23. Gestroania froggatti (Macleay). Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). ventrolateral ensiform setae, a large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 7 moderately elongate ensiform setae. Lateral plate with fairly dense fringe of moderately elongate, stiff nematiform setae at apical rim. DISCUSSION. By comparison with the type of G. amplipennis Gestro it became evident that Eudalia froggatti Macleay is extremely closely related to this species from which it differs only in minor character states. At the same time, both species are so different in certain remarkable morphological characters from all other known species of Eudalia that maintenance of Gestroania as a separate genus is well justified. Moore et al. (1987) record Eudalia froggatti only from the type locality. New records now enlarge the range from the southern margin of the Kimberley to its northeastern margin. Never- theless, this seems still an extremely rare species. DISTRIBUTION. Kimberley Division, WA. Both non type specimens captured at light. Gestroania setipennis sp. nov. (Figs 24, 31G, 36F) ETYMOLOGY. Refers to the multisetosc odd elytral intervals. MATERIAL. HOLOTYPE: 1 d, Greenvale, 70km SW, 22-30.xi.1995, A.J. Watts (SAM). PARATYPES: 1 d, Split Rock, 1 4km S of Laura, Nth Old, 23-26. vi. 1 975, GM (QM): 1 9 , Nth Qld, Pouth Ck. via Georgetown, 8.i. 1 980, RS (CBM); 1 d, Windsor T'land, via Mt Gamet/Qld, lO.ii. 1978, RS /Dicraspeda sp. n. det. B.P. Moore 1978 (MDPI); 1 9, 11.01E 136.45E, Rimbija I. Wessel islands NT, 3-14.ii.1977, TW (ANIC). 168 MEMOIRS OF THE QUEENSLAND MUSEUM DIAGNOSIS. Distinguished from G. storeyi by completely dark femora, much more coarsely crenulate elytral striae, only angulate but aspinose external elytral apices, and not microreticulate head; and from G. setipennis and G. amplipennis by the larger number of setiferous punctures on the odd intervals. DESCRIPTION. Measurements (Table 5). Colour (Fig. 36F). Surface black, labrum and mouth parts reddish to dark yellow, antennae completely yellow. Femora completely dark, tibiae and tarsi yellowish. Head . Fairly wide. Neck narrow, with con- spicuous transverse impression. Eyes large, laterally protruding, slightly separated from orbits which are about half as long as eyes and gently convex. Behind clypeus with a linear, fairly deep groove. Medially of eye with a slight sulcus, but without distinct ridge. Posterior supraorbital seta located very shortly behind posterior margin of eye. Mentum with elongate, acute, triangular tooth, with 2 pairs of elongate setae behind tooth, submentum with 2 very elongate setae on either side. Apex of glossa transverse, with 2 elongate median and 2 shorter lateral setae. Paraglossae free, narrow, surpassing glossy. Lacinia elongate, interior margin with a sparse fringe of spines. Antenna very narrow and elongate, surpassing base of pronotum by about 3 antennomeres, pilose from middle of 4th antennomere. Surface without microreticulation, or with extremely superficial remnants only, impilose and impunctate, with some elongate wrinkles only near eyes, highly glossy. Pronotum. Comparatively short (in genus), laterally gently convex, widest in front of apical third, dorsally moderately depressed, near apex with shallow triangular impression. Apex and base not bordered, lateral margin complete, slightly ridge-like, in anterior half except for the immediate apex, oblique and almost straight, towards base very slightly diverging, without sulcus medially of margin. Median line shallow. A single marginal seta situated just behind apical third. Proepisternum narrowly visible from above in middle. Disk with extreme dense and coarse, remarkably rugose punctuation. Surface almost devoid of microreticulation, impilose, despite the coriaceous microstructure moderately glossy. Elytra. Moderately elongate, towards apex considerably w-idened, disk fairly depressed, not raised towards suture. Base oblique, humeri widely rounded, lateral margin oblique in anterior half, very faintly incised at basal third, gently convex and almost evenly rounded towards lateral apical angle. Lateral apical angle angulate, not spinose, apex moderately excised, oblique and evenly concave, sutural angle angulate. Base narrowly margined to position of 4th stria, apex coarsely margined. Surface without any transverse impression. All striae complete and deeply impressed, coarsely punctate-crenulate, punctures become smaller posteriorly, but are still well perceptible at apex. Intervals convex throughout, though not carinate. Scutellar stria elongate, consisting of 9-10 punctures. 3rd, 5th, and 7th intervals each with more than 4 (usually 6-8) setiferous punctures, other intervals asetose, setae light brown, elongate, erect. Intervals impunctate, with coarse, transverse, distinct, though somewhat superficial microreticulation, surface gently glossy. Hind wings fully developed. Lower Surface. Proepisternum, prosternum, and lateral parts of mesothorax and metathorax including metasternum with dense, very coarse punctures. Middle of metastemum and abdomen impunctate, finely microreticulate, abdomen also TABLE 5. Measurements and ratios of the Australian species of Gestroania. N length length 1/w 1 u 1/w (mm) eye/orbit head prothorax elytra amplipennis 2 7.8-8. 1 2.05-2.15 1.03-1.05 1.26-1.32 1.64-1.70 froggatti 2 7.4-8. 0 1.90-1.95 1.04-1.05 1.25-1.31 1.59-1.60 setipennis 4 8.2-8. 6 1. 9-2.0 1.00-1.04 1.18-1.23 1.60-1.63 storeyi 6 8.3-9. 1 1.75-2.0 1.03-1.05 1.27-1.34 1.65-1.70 AUSTRALIAN ODACANTHINE GROUND BEETLES 169 slightly strigose. Metepistemum elongate, >2.5 x as long as wide at apex. Terminal abdominal sternum in male bisetose, in female quadrisetose and shortly pilose. Legs. Narrow and elongate. Tibiae sulcate on upper surface, 2 basal tarsomeres sulcate on outer and inner surfaces, though indictinctly so on protarsus. Tarsi not lobed, impilose on upper surface, 5th tarsomere with a fringe of setae below. Claws large, smooth. 1 st - 3rd tarsomeres of male anterior tarsus with sparse, slightly asymmetric squamosity. Male Genitalia (Fig. 24). Terminal abdominal stemite in middle gently incised. Genital ring fairly narrow and elongate, moderately triangular, barely asymmetric, with narrow, triangular apex. Aedeagus slender and elongate, moderately depressed, laterally little sinuate, lower surface gently concave throughout. Orificum short. Apex short, fairly wide, not upturned nor knobbed, turned to right, moderately bi-incised at right side. Folding of internal sac simple. Parameres of moderately dissimilar shape, left paramere much larger than right one, left paramere stout with gently rounded apex, right paramere with narrow apex. Female Genitalia (Fig. 31G). Stylomere 2 moderately elongate, laterally moderately curved, with acute apex. With 3 medium-sized ventrolateral ensiform setae, a large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 6 moderately elongate ensiform setae. Lateral plate with fairly dense fringe ot moderately elongate, stiff nematiform setae at apical rim. Variation. Little variation noted. DISTRIBUTION. N Qld, Arnhem Land, NT. RELATIONSHIPS. Although in shape and size this species is extremely similar to G. amplipennis and G. froggatti, the multisetose intervals demonstrate a closer relationship to G. storeyi than to either species. Gestroania storeyi sp. nov. (Figs 25, 31H, 37A) ETYMOLOGY. For Ross Storey. MATERIAL. HOLOTYPE: d, N WA, Kununurra, 22.xii.l991-5.i.l992, R. Storey / Dicraspeda spp. det. B.P. Moore 1992 (QMT 99171). PARATYPES: 4 9 9, same data (CBM, MDPI); 1 9, Katherine Gorge, Jan. 27/1977 N.T., mm ! Dicraspeda sp. n. det. B.P. Moore 1978 (MDPI). DIAGNOSIS. Distinguished from G. setipennis by conspicuously bicoloured femora, much less coarsely crenulate elytral striae, spinose external elytral apices, and microreticulate head; and from both, G. setipennis and G. amplipennis by the larger number of setiferous punctures on the odd intervals. DESCRIPTION. Measurements (Table 5). Colour (Fig. 37A). Surface black, labrum, and mouth parts reddish to dark yellow, antenna piceous, apical antennomeres usually slightly lighter. Femora in basal half yellow, in apical half contrastingly black, tibiae except for base and apex, yellow, tarsi infiiscate. Head. Fairly wide. Neck narrow, with conspicuous transverse impression. Eyes large, laterally protruding, slightly separated from orbits which are about half as long as eyes and gently convex. Behind clypeus with wide, fairly deep, circular groove the bottom of which may be somewhat striolate. Medially of eye with a slight sulcus that extends to about middle of eye, but without distinct ridge. Posterior supraorbital seta located very shortly behind posterior margin of eye. Mentum with elongate, acute, triangular tooth, with 2 pairs of elongate setae behind tooth, submentum with 2 very elongate setae on either side. Apex of glossa transverse, with 2 elongate median and 2 shorter lateral setae. Paraglossae free, narrow, surpassing glossy. Lacinia elongate, interior margin with a sparse fringe of spines. Antenna very narrow and elongate, surpassing base of pronotum by about 3 antennomeres, pilose from middle of 4th antennomere. Surface with fine though distinct, isodiametric micro- reticulation, impilose and impunctate, with some transverse wrinkles only on middle of frons, moderately dull. Pronotum. Comparatively elongate, laterally gently convex, widest at apical third, dorsally depressed, near apex with shallow triangular impression. Apex and base not bordered, lateral margin complete, slightly ridge-like, in anterior half" very gently convex, towards base slightly diverging, without sulcus medially of margin. Median line shallow. A single marginal seta situated just in front of middle. Proepistemum narrowly visible from above in middle. Disk with extreme dense and coarse, rugose punctuation, punctures tend to form irregular transverse sulci. Surface also with fine, isodiametric micro- reticulation, impilose, dull. Elytra. Comparatively elongate, towards apex considerably widened, disk depressed, slightly 170 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 25. Gestroania storeyi sp. nov. Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). raised towards suture. Base oblique, humeri widely rounded, lateral margin oblique in anterior half, very faintly incised at basal third, gently convex and almost evenly rounded towards lateral apical angle. Lateral apical angle spinose, apex deeply excised, oblique and evenly concave, sutural angle shortly spinose. Base narrowly margined to position of 4th stria, apex coarsely margined. Surface without any trans- verse impression, but in apical half moderately raised towards suture. All striae complete and deeply impressed, finely punctate-crenulate, though punctures become smaller posteriorly and diminish in front of apex. Intervals convex throughout, though not carinate. Scutellar stria elongate, consisting of 9-10 punctures. 3rd, 5th, and 7th intervals each with more than 4 (usually 6-8) setiferous punctures, other intervals asetose, setae light brown, elongate, erect. Intervals impunctate, with very coarse, slightly transverse microreticulation, surface remarkably dull. Hind wings fully developed. Lower Surface . Proepistemum, prostemum, and lateral parts of mesothorax with dense, very coarse punctures. Mesostemum, metathorax, and abdomen impunctate, densely microreticulate, and abdomen also strigose. Metepisternum elongate, >2.5 x as long as wide at apex. Terminal abdominal sternum in male bisetose, in female quadrisetose and shortly pilose. Legs. Narrow and elongate. Tibiae sulcate on upper surface, 2 basal tarsomeres sulcate on outer and inner surfaces. Tarsi not lobed, impilose on upper surface, 5th tarsomere with a fringe of setae below. Claws large, smooth. 1st - 3rd tarsomeres of male anterior tarsus with sparse, slightly asymmetric squamosity. Male Genitalia (Fig. 25). Terminal abdominal stemite in middle gently incised. Genital ring moderately wide, triangular, barely asymmetric, with narrow, triangular apex. Aedeagus slender and elongate, moderately depressed, laterally little sinuate, lower surface gently concave throughout. Orificum short. Apex short, fairly wide, not upturned nor knobbed, turned to right, moderately incised at right side. Folding of internal sac simple. Parameres of moderately dissimilar shape, left paramere much larger than right one, left paramere stout with almost transverse apex, right paramere with widely rounded apex. Female Genitalia (Fig. 31H). Stylomere 2 moderately elongate, laterally moderately curved, with acute apex. With 3 medium-sized ventrolateral ensiform setae, a rather large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 5-6 moderately elongate ensiform setae. Lateral plate with fairly dense fringe of moderately elongate, stiff nematiform setae at apical rim. Variation. Slight differences noted in shape of pronotum, and in extent of punctuation on head, pronotum, and elytral striae. The single male also is definitely smaller and has larger eyes. DISTRIBUTION. N NT, NE WA. KEY TO SPECIES OF GESTROANIA 1. 3rd, 5th, and 7th intervals each with more than 4 setiferous punctures 2 3rd interval at most 4-setose. 5th 2-3-setose, 7th unisetose, rarely unilaterally bisetose 3 2. Elytra longer and narrower, ratio 1/w > 1 .65, external and sutural angles distinctly spinose, punctuation of striae finer (Fig. 37A); pronotum longer, anteriorly more incurved; head regularly and completely microreticulate; basal half of femora yellow, apical half contrastingly black; aedeagus (Fig. 24) storeyi sp. nov. Elytra shorter and wider, ratio l/w <1.63, external and sutural angles only shortly angulate, punctuation of striae coarse (Fig. 36F); pronotum shorter, anteriorly little incurved; head not or barely microreticulate; femora unicolourous dark; aedeagus (Fig. 23). setipennis sp. nov. 3. Femora dark, coxae, tibiae and tarsi dark yellowish (Fig. 36E); orbits slightly longer, ratio eyc/orbil <1.95; striae of elytra less coarsely punctate, especially in apical half; apex of aedeagus very slightly knobbed (Fig. 22). Jroggatti (Macleay) Legs light yellow, only apical tip of femora dark (Fig. 36B); orbits slightly shorter, ratio eyc/orbit >2.05; striae of elytra very coarsely punctate, even in apical half; aedeagus unknown amplipennis (Gestro) AUSTRALIAN ODACANTHINE GROUND BEETLES 171 Giachinoana Baehr, 2003b Giachinoana Baehr, 2003b: 100. TYPE SPECIES. Giachinoana carinipennis Baehr, 2003b, by original designation. DIAGNOSIS. Distinguished from most Australian genera except Gestroania by the sharply angulate external apices of the elytra; distinguished from Gestroania by the carinate odd intervals and the distinct sulcus and ridge inside of eye. Giachinoana carinipennis Baehr, 2003 (Fig. 311, 37B) SUPPLEMENTARY DESCRIPTION. Female Genitalia (Fig. 31H). Stylomere 2 moderately elongate, laterally moderately curved, with acute apex. With 3 medium-sized ventrolateral ensiform setae, a rather large dorsomedian ensiform seta situated about in middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base ot stylomere 1 with 5-6 comparatively short ensiform setae. Lateral plate with fairly dense fringe of moderately elongate, stiff nematiform setae at apical rim. Variation. Very little variation noted, neither in size nor in shape and surface structure. DISTRIBUTION. N NT, NE WA. Sampled at light. NEW RECORDS. Type species (8 ex.): WA: Kununurra, 22.xii. 1991-6. i. 1992, RS (CBM, MDPI, QM). Lachnothorax Motschulsky, 1862 Lachnothorax Motschulsky, 1862: 48; Sloanc, 1917: 414; 1923: 33; Csiki, 1932: 1542; Licbke, 1938: 103; Darlington, 1968: 214; Lorenz, 1998: 420; Baehr, 1996d: 2 ; 2000 : 11 . TYPE SPECIES. Lachnothorax higuttatus Motschulsky, 1862, by monotypy. DIAGNOSIS. Upper and lower surfaces densely pilose; head short, rounded, lacking a sulcus and ridge medially of eyes; elytra short, con- spicuously bimaculate, with coarsely punctate though not impressed striae. AUSTRALIAN SPECIES. Lachnothorax tokkia Gestro 1875 (Fig. 37C). Myrmecodemus Sloane, 1923 Myrmecodemus Sloane, 1923: 33; Csiki, 1932: 1543; Liebke, 1938: 106; Moore ct al., 1987: 275; Lorenz, 1998: 420. TYPE SPECIES. Casnonia riverinae Sloane, 1890, by original designation. DIAGNOSIS. Head globose, with comparatively small, though convex eyes; prothorax globose; elytra short, wide, bearing a transverse im- pression across basal half; striation incomplete; proepistemum and prostemum fused; elongate, erect setae on head, lower surface of prothorax, and on disk and along the margins of pronotum; , rows of elongate, erect setae on odd intervals, DISCUSSION. Until now this genus comprised M. fomicoides (Sloane), Af. globulicollis (Macleay) and M. riverinae (Sloane). M. globulicollis and M. riverinae are very closely related, and share with M. formicoides the glabrous, impilose surface of head, pronotum, and elytra (apart from fixed setae), the smooth surface of pronotum, and the absence of striae in the apical two thirds of the elytra. One of the new species described herein, although similar in shape and certain other features, differs in all mentioned characters common to the three species. Therefore, for this unique species Myrmecodemus (Trichodemus) subgen. nov. is erected. Myrmecodemus (Myrmecodemus) s. str. Myrmecodemus Sloane, 1923: 33. TYPE SPECIES. Casnonia riverinae Sloane, 1890, by original designation. DIAGNOSIS. See Sloane (1923: 33). Characters of subgeneric value are: surface of head, pronotum, and elytra (apart from fixed setae) impilose, surface of pronotum smooth, without any trace of transverse wrinkles, and apical two thirds of elytra not striate. DISTRIBUTION. N NT and WA, S inland NSW, N Vic. Myrmecodemus (M.) globulicollis (Macleay, 1888) (Fig. 37D) Casnonia globulicollis Macleay, 1888: 447. Lachnothorax globulicollis. Sloane, 1910: 397 ; 1917:413. Myrmecodemus globulicollis, Sloane, 1923: 33, Csiki. 1932: 1543; Liebke, 1938: 107; Moore ct al.. 1987: 275; Lorenz, 1998: 420. MATERIAL. LECTOTYPE (here designated): 6 (damaged), N WA/ HOLOTYPE/ Casnonia globulicollis , Macl. Barrior Range N WA (ANIC-MMS). NEW RECORDS (5 ex.): NT: Tindal, 14.3 IS, 132.22E, 1-20. xii. 1967, WV (ANIC). - WA: Kununurra, 22.xii. 1 99 1 -6.i. 1 992, RS (CBM, MDPI); Fitzroy Crossing, 220 mi. E of Broome, 25.ii.68, EM (ANIC). 172 MEMOIRS OF THE QUEENSLAND MUSEUM DIAGNOSIS. Distinguished from the other species, except for M. riverinae (Sloane), by large size and bright red prothorax. From the latter species it is distinguished by purplish black colour and quadrimaculate pattern of the elytra. DISTRIBUTION. N NT, Kimberley. New records were collected at light. Myrmecodemus (M.) riverinae (Sloane, 1 890) (Fig. 37E) Casnonia riverinae Sloane, 1890: 643. Lachnothorax riverinae, Sloane, 1910: 397; 1917: 413. Myrmecodemus riverinae , Sloane. 1923: 33, Csiki, 1932: 1543; Liebke, 1938: 107; Moore ct al., 1987: 275; Lorenz, 1998:420. MATERIAL. LECTOTYPE (here designated): cJ, E.P. 7218/? HOLOTYPE (ANIC). PARALECTOTYPES: 2 9 9, 2 (?sex, damaged), on same card, E.P. 72 18 / Myrmecodemus riverinae SI. Id. by T.G Sloane (ANIC). NEW RECORDS (7 ex.): NSW: N. S. Wales/Cawowa riverinae SI. Id. By A.M. Lea (CBM, MV); Casnonia riverinae SI, NSW (MV). - VIC: Inglewood, 27.xii.35, CO (MV). DISCUSSION. Moore et al. (1987) noted the 5 type specimens as located in the SAM, Adelaide. This is a lapse because the specimens are indeed in ANIC and were labeled by Sloane himself. The ‘Holotype’ label is insignificant, because Sloane in his description did not give any information about type designations, and the label was attached later to the specimen. Therefore the specimen bearing the Holotype label is here designated Lectotype. DIAGNOSIS. Easily distinguished from the other species, except for M. globulicollis (Macleay), by large size and bright red prothorax. From the latter species it is distinguished by glossy blue colour and semilunar pattern of the elytra. DISTRIBUTION. S central NSW, N Vic. Myrmecodemus (M.) formicoides (Sloane, 1910) (Figs 26, 32A, 37F) Lachnothorax formicoides Sloane, 1910: 397; 1917:413. Myrmecodemus formicoides , Sloane, 1923: 33, Csiki, 1932: 1543; Liebke, 1938: 108; Moore et al.. 1987: 275; Lorenz. 1998:420. MATERIAL. LECTOTYPE (here designated): 9, Port Darwin Dodd .03 / Myrmecodemus fonnicoides SI. Id. by T.G Sloane/HOLOTYPE Myrmecodemus formicoides SI. PJD (ANIC). PARALECTOTYPE: 1 9 (immat.), Port Darwin Dodd .03 / Myrmecodemus fonnicoides SI. Id. by T.G Sloane (ANIC). NEW RECORDS (4 ex.): NT: NT1, 3km E Humpty Doo, 3.viii.l995, MB (CBM); Humpty Doo, 6km E 9.ii-6.iii. 1987, RS /Myrmecodemus fonnicoides SI. AW-H det. 1987 (MDPI); Oenpelli, from P. Cahill leg. 6.xii.l8 (MV); 12.52S, 132.50E, Koongarra, 6-10.iii.73, MU (ANIC). DIAGNOSIS. Small; pronotum dark, glabrous; distinguished from M, lucai by shorter, wider elytra, reddish subhumeral fascia, circular apical spot, shorter eye, and apical part of aedeagus more upturned and more decidedly spoon-shaped. SUPPLEMENTARY DESCRIPTION. Measurements (Table 6). Male Genitalia (Fig. 26). Terminal abdominal stemite in middle incised. Genital ring fairly elongate, slightly asymmetric, gently triangular, shortly narrowed to the narrow, asymmetrically triangular apex. Aedeagus fairly slender and elongate, depressed, laterally barely sinuate, lower surface conspicuously bisinuate, upturned towards apex. Orificum elongate. Apex elongate, depressed, straight, not knobbed, but decidedly spoon-shaped, very slightly turned to right, incision at right side less deep than at left side. Folding of internal sac simple. Parameres of dissimilar shape, left paramere much larger and also longer than right one, right paramere short and high, both with wide, rounded apex. Female Genitalia (Fig. 32A). Stylomere 2 comparatively short and stout, laterally evenly curved, with acute apex. With 2 small ventrolateral ensiform setae, a fairly small dorsomedian ensiform seta situated above middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 4 short but very stout ensiform setae. Lateral plate with dense fringe of stiff setae at apical rim. DISCUSSION. Darlington designated one specimen ‘Holotype’, although Sloane gave no indication about type designations in his description. Thus, Darlington’s designation is invalid according to the rules of ICZN and the specimen is here designated Lectotype. Moore et al. ( 1 987) noted the species only from the type locality. In spite of the new records , this seems to be an extremely rare species. DISTRIBUTION. NT from W Arnhem Land to Darwin. One specimen from a Barber trap near a small pool in Tropical Savannah Woodland. Probably a hygrophilous, ground-living species. RELATIONSHIPS. This species is closely related to M. lucai and might be taken for the same species, though certain differences in shape and colouration contradict this supposition. Both AUSTRALIAN ODACANTHINE GROUND BEETLES 173 FIG 26. Myrmecodemus formicoides (Sloane). Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). species share the same range, although M. lucai probably is more widely distributed to the west. At Oenpelli both species were collected together, but even the specimens mounted together on the same card arc immediately distinguished. Myrmecodemus (M.) lucai sp. nov. (Figs 27, 32B, 38A) ETYMOLOGY. For Luca Toledano, collector of the main series. MATERIAL. HOLOTYPE: d, NT, Gregory N.P. Timber Creek 30-31.xii.96 at light/al lume Leg. L. Toledano, R. Olivieri (WAM). PARATYPES: 2 <3, 9 9 9, same data (ANIC, CBM, CFP, CSM, CTV); 1 9 (damaged), NT Gregory N.P. Victoria Crossing 1-2197 at light/al lume, LT, RO (CTV); 1 d, N.T. Kakadu N.P. 22-25.iii.93 Cooinda at light/al lume, LT (CBM); 1 9, Australie/Collection E. Rousseau (IRSBN); 2 S6> Oenpili, N.T. from P. Cahill leg. 6.xii.l8 (MV); 1 9, 15.3 IS 143.55E GPS 5km SE Hann River bridge, 1511994, at light, PZ & EE (ANIC). DIAGNOSIS. Small; pronotum dark; surface glabrous; distinguished from M. formicoides by longer and narrower elytra, indistinct or even absent subhumeral fascia, elongate apical spot, larger eye, and apical part of aedeagus barely upturned and less decidedly spoon-shaped. DESCRIPTION. Measurements (Table 6). Colour (Fig. 38A). Surface blackish, very dark piceous on centre of head and sometimes on basal part of elytra. Labrum, mandibles, palpi, and 3 basal antennomeres more or less dark reddish, rest of antenna dark. Apical spot of elytra light yellow, sharply delimited, square, meeting apical margin. Also lateral margin narrowly yellowish in apical part. Subhumeral fascia absent or very inconspicuous, area of fascia more or less dark piceous, as sometimes the whole base of elytra. Base of elytral epipleura dark, apical 3 A light yellow. Legs light yellow with contrastingly dark knees. Head. Wide, rhomboidal. Neck remarkably narrow. Eyes comparatively small, though laterally protruding, slightly separated from orbits which are by far longer than eyes and gently convex. Behind clypeal suture laterally with shallow, oblique impression, and in middle with more or less distinct v-shaped impression. Medially of eye without any indication of a sulcus. Posterior supraorbital seta located slightly behind posterior margin of eye. Behind posterior seta with 2 additional pores and setae, one of which is situated laterally. Mentum with elongate, triangular, at apex slightly obtuse tooth, with 2 setae behind tooth, submentuni with 2 elongate setae on either side. Glossa short, apex transverse, with 2 elongate setae. Paraglossae free, narrow, barely surpassing glossy. Lacinia elongate, interior margin with a sparse fringe of spines. Antenna elongate, surpassing base of pronotum by at least 2 antennomeres, sparsely pilose from 2nd antennomere. Surface highly glossy, without microreticulation, glabrous, impunctate. Pronotum. Short, somewhat globose, laterally convex, dorsal ly very convex, near apex with shallow, near base with deep and wide, transverse impression. Base thickly bordered, lateral margin complete, slightly ridge-like, gently convex, towards base slightly diverging, without sulcus medially of margin. Median line barely recognisable. A well developed marginal pore and seta situated in apical fifth near margin, disk with some additional elongate erect setae. Proepisternum and proepimeron well visible from above. Disk impunctate (except for elongate setae), without microreticulation, glossy. Elytra. Rather short and wide, somewhat square, widest about in middle, dorsal surface moderately convex. Base moderately oblique, humeri rounded, lateral margin gently and almost evenly convex, but very faintly incised at basal third. Apex wide, lateral apical angles well indicated but obtuse, apex oblique and laterally well excised. Base narrowly margined halfway to scutellum, lateral margin moderately wide, apex coarsely margined. Surface in anterior third with distinct, transverse impression. Striae almost completely absent, only odd intervals marked by rows of conspicuous punctures and elongate erect setae. Surface at base slightly uneven, as odd intervals are faintly raised. Scutellar stria 174 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 27. Myrmecodemus lucai sp. nov. Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). barely recognisable. Surface impunctate (except for elongate setae), very glossy, disk without microreticulation, but with very superficial, isodiametric microreticulation along lateral margin and in lateral parts of subhumeral grooves that give the surface a somewhat silky appearance. Hind wings present. Lower Surface. Proepisternum not separated from prostemum. Both impunctate, except for some elongate setae on prostemum, and without microreticulation. Proepimeron with extremely rugose, ridge-like punctuation, apical rim of mesothorax with very coarse punctuation, rest of mesothorax, metathorax and abdomen glabrous, without microreticulation, very glossy. Metepisternum elongate, c. 2.5 X as long as wide at 'apex. Abdominal sterna quadri- or 6-sctose, terminal abdominal sternum in male bisetose and glabrous, in female quadrisetose, with short adpressed pilosity. Legs. Narrow and elongate. Tarsi not lobed, densely pilose on upper surface, 5th tarsomere with a dense fringe of elongate setae below. Claws large, smooth. 2nd-3rd tarsomeres of male anterior tarsus with sparse, asymmetric squamosity. Male Genitalia (Fig. 27). Terminal abdominal sternite in middle incised. Genital ring fairly elongate, slightly asymmetric, gently triangular, shortly narrowed to the narrow, acute, triangular apex. Aedeagus fairly slender and elongate, depressed, laterally barely sinuate, lower surface only immediately near base concave, in apical three quarters gently convex. Orificum elongate. Apex elongate, depressed, straight, not knobbed, but moderately spoon-shaped, very slightly turned to right, incision at right side barely deeper than at left side. Folding of internal sac simple. Parameres of dissimilar shape, left paramere much larger and also longer than right one, right paramere short and high, both with wide, rounded apex. Female Genitalia (Fig. 32B). Stylomere 2 comparatively short and stout, laterally evenly curved, with acute apex. With 2 small ventrolateral ensiform setae, a fairly small dorsomedian ensiform seta situated above middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 4-5 short but stout ensiform setae. Lateral plate with dense fringe of stiff setae at apical rim. Variation. Rather little variation noted in shape of pronotum and distinctness of colouration of subhumeral fascia. DISTRIBUTION. N Australia from Cape York Peninsula to NTAVA border. Recently caught specimens taken at light, close to river beds or lagoons. Probably a ground-living, hygrophilous species. RELATIONSHIPS. This species is very closely related to M. formicoides with which it apparently shares part of its range. Myrmecodemus (Trichodemus) subgen. nov. TYPE SPECIES. Myrmecodemus pilosellus sp. nov. DIAGNOSIS. Surface of head densely punctuate, with elongate, depressed, anteriorly inclined pilosity; pronotum bearing dense, coarse, transverse wrinkles in middle; elytra with remarkably deep impression in anterior third and with a likewise conspicuous boss on either side in front of that impression, surface within im- pression and along suture striate and altogether coarsely punctate and pilose; rest of elytra (apart from fixed setae) impilose. Myrmecodemus (T.) pilosellus sp. nov. (Figs 28, 32C, 38B) ETYMOLOGY. Refers to the dense pilosity of the head. MATERIAL. HOLOTYPE: 6, 13.27S 142.42E, Qld, Mungkan Kandju N.P., Langi Lagoon, 29.vi.-5.vii. 1998, T.A. Weir fANIC). PARATYPES: 1 d, Stewart R. Q., WD/J.7710 Lachnothorax ? formicoides Sin Qld (SAM); 1 9, 13.59S 143.33E Silver Plains, Cape York Pen. Qld 1 .iv. 1 965, JW / Myrmecodemus ?? T.A. Weir 1 993 ( ANIC); 1 9, DAOS 142.40E Qld 2km N Rokeby, 16.xii.1993 at light, PZ (ANIC); 1 9, Nth Qld 17.1.2000 Laura, SB (CBM). AUSTRALIAN ODACANTHINE GROUND BEETLES 175 TABLE 6. Measurements and ratios of species of the formicoides- group and Myrmecodemus ( Trichodemus). N length (mm) length eye/orbit 1/w head 1/w prothorax 1/w elytra 1 fi formicoides 3 4.75-4.90 0.62-0.65 1.30-1.33 1.33-1.34 1.45-1.47 lucai 6 5. 1-5.4 0.73-0.83 1.26-1.31 1.29-1.34 1.50-1.54 | pilosellus 4 4. 5-4. 8 0.51-0.56 1.20-1.24 1.17-1.21 1.45-1.48 i DIAGNOSIS. Head densely punctuate, pilose; pronotum rugose; elytra apically bistriate, in parts pilose. DESCRIPTION. Measurements (Table 6). Colour (Fig. 38B). Black, elytra with a triangular dirty yellow to light reddish subhumcral fascia and a similarly coloured transverse subapical fascia. Apex behind fascia more or less dark piceous. Base of elytral epipleura dark, becoming yellow towards apex. Labrum, mandibles, and palpi dark piceous with slightly lighter borders. Basal antennomere dark reddish to piceous, 3 following antennomeres darker, almost blackish, rest of antenna again slightly lighter, dark reddish to piceous. Coxae piceous, femora basally light yellow or reddish, apical half (profemur) or apical third (meso- and metafemur) blackish, tibiae light yellow or reddish with the very base and apex dark, tarsi dark. Lower surface black to dark piceous. Head. Wide and short, laterally and dorsally very convex. Neck remarkably narrow. Eyes small, though laterally conspicuously protruding, well separated from orbits which are by far longer than eyes and very convex. Behind clypeal suture laterally with shallow, oblique impression, and in middle with shallow v-shaped impression. Medially of eye without any indication of a sulcus. Posterior supraorbital seta located just behind posterior margin of eye. Shortly behind posterior seta with an additional elongate setae, and another far behind near the ‘neck’, setae sometimes difficult to recognise within the adpressed setosity. Apex of labrum transverse, 6-setose, setae elongate. Mentum with elongate, triangular, at apex slightly obtuse tooth, with 2 setae behind tooth, submentum with 2 very elongate setae on either side. Glossa short, apex transverse, with 2 elongate setae. Paraglossae free, narrow, not surpassing glossy. Lacinia elongate, interior margin with a sparse fringe of spines. Penultimate palpomere of labial palpus with 2 very elongate setae. Antenna moderately elongate, just surpassing base of pronotum. sparsely pilose from 1st antennomere, densely pilose from middle of 3rd antennomere. Surface of labrum, clypeus and middle of frons behind clypeus glabrous, apart from 2 setae just behind clypeal suture, glossy. Labrum with about isodiametric, superficial micro- reticulation, rest of surface without microreticulation, though with dense and somewhat rugose punctuation and covered by elongate hairs that are inclined and adpressed anteriorly. Pronotum. Short, somewhat globose, laterally convex, dorsally very convex, near apex with shallow, near base with deep and wide, transverse impression. Base thickly bordered, lateral margin complete, coarse slightly ridge-like, gently convex, towards base slightly diverging, with a shallow sulcus medially of margin in anterior two thirds. Median line distinct, moderately impressed. A well developed marginal pore and seta situated in apical fourth near margin, disk with some additional, very elongate erect setae and several shorter setae. Proepistemum and proepimeron well visible from above. Disk in middle with coarse, very rugose, transverse sulci, impunctate (except for elongate setae), without microreticulation. Apex and base with coarse, rugose punctuation. Elytra. Short and wide, subquadrate, widest about midlength, dorsal surface convex, very uneven. Base little oblique, humeri distinct though rounded, lateral margin gently and almost evenly convex, but very faintly incised at basal third. Apex wide, lateral apical angles well indicated but obtuse, apex oblique and laterally well excised, incurved towards suture. Base coarsely margined close to scutellum, lateral margin extremely narrow throughout, apex finely margined. Surface in anterior third with very deep, transverse impression that is laterally enlarged to form a triangle. In front of impression surface raised to 2 conspicuous humps, behind impression surface evenly convex. Striae only recognisable within the subbasal transverse impression, along suture in posterior two thirds, and at apex, marked by extremely coarse, rugose punctures and dense, moderately elongate, yellow setosity. In other parts of surface striae completely absent, only odd intervals marked by rows of conspicuous punctures and elongate erect setae. Apart from transverse subbasal impression, 2 inner striae, and apex, surface impunctate, impilose (except for elongate setae), without microreticulation, and very glossy. 176 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 28. Myrmecodemus pilosellus sp. nov. Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). Scutellar stria barely recognisable. As a summary; surface of elytra remarkably uneven and differently structured. Hind wings present. Lower Surface. Proepisternum not clearly separated from prostemum, a very inconspicuous, superficial suture only visible in basal half. Both, proepisternum and prosternum impunctate, except for few elongate setae on prostemum, and without microreticulation. Proepimeron with rugose punctuation, mesothorax and metathorax impunctate, without microreticulation, very sparsely setose. Abdomen glabrous, without microreticulation, very glossy. Metepisternum elongate, c. 2.5 x as long as wide at apex. Abdominal sterna quadrisetose in middle, and with a row of 4-6 shorter setae laterally on either side. Terminal abdominal sternum in male bisetose and glabrous, in female quadrisetose, with short adpressed pilosity. Legs. Narrow and elongate. Tarsi not lobed, densely pilose on upper surface, 5th tarsomere with a dense fringe of elongate setae below. Claws large, smooth. 2nd-3rd tarsomeres of male anterior tarsus with sparse, asymmetric squamosity. Male Genitalia (Fig. 28). Terminal abdominal stemite in middle incised. Genital ring fairly elongate, slightly asymmetric, in basal two thirds almost parallel, shortly narrowed to the wide, obtuse apex. Aedeagus fairly slender and elongate, very depressed, laterally barely sinuate, whole lower surface gently convex, even near base barely concave. Orificum moderately elongate. Apex short, wide, depressed, straight, not knobbed, but moderately spoon-shaped, very slightly turned to right, incision at right side slightly deeper than at left side. Folding of internal sac simple. Parameres of dissimilar shape, left paramere much larger and also longer than right one, with almost transverse apex, right paramere very short and high, with wide, rounded apex. Female Genitalia (Fig. 32C). Stylomere 2 comparatively short and stout, laterally little curved, with acute apex. Apparently without ventrolateral ensiform setae, with a small dorsomedian ensiform seta situated above middle of stylomere, and with a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 4 elongate, stout ensiform setae. Lateral plate with moderately dense fringe of elongate, stiff setae at apical rim. Variation. Little variation noted due to scarce material. One specimen has colouration of light areas on elytra and legs light reddish rather than light yellow, perhaps because teneral or effected by preservation. DISTRIBUTION. Lower Cape York Peninsula. Holotype collected at light in open forest. KEY TO AUSTRALIAN SPECIES OF MYRMECODEMUS 1. Head, pronotum, and elytra (except for fixed setae) glabrous, impilosc; pronotum without transverse wrinkles; elytra without striae in apical two thirds. Myrmecodemus s. str 2 Head densely punctate and pilose; pronotum with coarse transverse wrinkles in middle; elytra with very deep, punctate and pilose impressions in basal third, with 2 punctate and pilose striae along suture. Trichodemus subgen. nov pilosellus sp. nov. 2. Pronotum bright red; body size larger, length >7mm . . 3 Pronotum black or piccous; body size smaller, length <5. 5mm 4 3. Elytra quadrimaculate, with red posthumcral and apical maculae on purplish-black background. globulicollis (Macleay) Elytra with a red fascia in apical half, ground colour of elytra bluish-black riverinae (Sloanc) 4. Elytra shorter and wider, ratio length/width e. 1.45; subhumeral fascia in the basal impression distinct, pale reddish, subapical yellow spot circular, removed from apical margin; lateral margin of elytra wider, completely and conspicuously yellow (Fig. 37F); eye shorter in comparison to orbit, ratio length eyc/orbil c. 0.65; apex of aedeagus more upturned and more decidedly spoon- shaped (Fig. 25) . . . formicoides ( Sloane) Elytra longer and narrower, ratio length/width >1.50; subhumeral fascia in the basal impression absent or extremely indistinct, subapical yellow spot elongate, widely meeting apical margin; lateral margin of elytra narrower, not completely and also less conspicuously yellow (Fig. 38A); eye longer in comparison to orbit, ratio length cye/orbit >0.70; apex of aedeagus barely upturned and less decidedly spoon-shaped (Fig. 26). lucai sp. nov. AUSTRALIAN ODACANTHINE GROUND BEETLES 177 Neoeudalia gen. nov. TYPE SPECIES. Eudalia nigra Sloane, 1910. DIAGNOSIS. Dense, erect surface pilosity; a shallow transverse impression near apex of elytra, tumid 7th interval in apical third, barely excised apical margin of elytra, sparsely setose 3rd antennomere. DESCRIPTION. Head large, with large, laterally markedly protruding eyes; labrum anteriorly straight; mandibles elongate; both palpi impilose, maxillary palpus also asetose; terminal palpomeres apically transversely cut; mentum with a large triangular, apically slightly incised tooth; glossa at apex almost transverse, with 2 elongate median and 2 much shorter lateral setae; paraglossae membranous, slightly surpassing and largely separated from glossa; inner margin of lacinia markedly dentate; mentum with 2 elongate setae, submentum with 4 elongate setae, lower surface of head below eyes with an elongate seta on either side; antenna elongate, 2 basal antennomeres impilose, 3rd antennomere with few elongate setae below apical setae, antenna more densely pilose from mid of 4th antennomere, basal antennomere with a single seta; medially of eye with a sulcus which is laterally bordered by a costa; neck well separated from head; surface of head sparsely but coarsely punctate, and with elongate, erect setosity; posterior supraorbital seta difficult to dis- tinguish; prothorax with complete but not sharply raised lateral borders; proepisterna laterally projecting beyond lateral margin and visible from above; mediad of lateral border with well developed sulcus that is medially bordered by an edge; margin and sulcus with a very dense fringe of elongate setae; disk densely punctate near base, with elongate, erect setosity; elytra elongate, not spinose nor sinuate at apex, with one transverse sulcus in anterior third, and another shortly in front of apex; striae complete, coarsely punctate; intervals convex, punctate, 7th interval tumid in apical third; surface with dense, elongate, erect setosity; legs slender; upper surface of tarsi impilose, claws simple; terminal abdominal stemite in male quadrisetose; aedeagus with short, asymmetric, somewhat knobbed apex, and with simple folding of internal sac. RELATIONSHIPS. Because of certain structural similarities, Neoeudalia is most probably related to Dicraspeda , Basistichus , Gestroania and Giachinoana , though in some characters it is more plesiomorphic than these genera. Probably, Neoeudalia takes a somewhat intermediate position between Eudalia and Dicraspeda as enumerated below, and it could be regarded a plesiotypic, basic branch of the Dicraspeda complex. Characters o/’Neoeudalia shared with Eudalia. Disk ofpronotum densely and coarsely punctate Elytral striae coarsely punctate Apex of elytra not excised nor spined Characters o/Neoeudalia shared with Dicraspeda Costa and sulcus inside of eye well developed Lateral channel of pronotum present Characters peculiar to Neoeudalia. Surface with remarkably elongate, erect setosity Head with a pair of elongate setae below eyes Lateral margin of pronotum not costate, sulcus very coarsely punctate, margin with remarkably dense and elongate fringe of setae Elytra with a transverse impression near apex 7th interval tumid in apical third Terminal abdominal stemite in male quadrisetose Legs very conspicuously bicoloured DISTRIBUTION. S central NSW, N Vic, S central Qld, and N NT and WA adjacent to border. Neoeudalia nigra (Sloane, 1900) (Figs 29, 32D, 38C) Eudalia nigra Sloane, 1900: 581; 1917: 417; Csiki, 1932: 1542; Liebke, 1938: 106; Moore ct al., 1987: 274; Lorenz, 1998:421. Dicraspeda nigra Sloane, 1923: 31 . MATERIAL. NEOTYPE (here designated): 9, C.H ./ Eudalia nigra SI. Topotype/ HOLOTYPE E. nigra SI. PJD (ANIC). NEW RECORDS (12 ex.): Vic: Hattali, CO (CBM, MV). - Qld: Eudalia nigra SI. Qld (MV); Rockhampton, AL/J.7706 Eudalia nigra Sloane (SAM); Eudalia nigra SI. (SAM): 623/Dawson R. (CBM, SAM); 7km NE ofToIga, xii.1988, RS & DF (MDPI); Pinnarendi Stn 60km W of Mt Garnet, 7,ii.I989. DH (QM); Musgrave, 5/72, GB (ANIC). - NT: 12.06S, 133.04E, Cooper Creek, 1 9km E by S of Mt Borradaile, 3 1 .v.73, EM (ANIC); 1.6km E on Warrawarange Rd., off Stuart Hwy, 10km S. McMillans Rd Darwin, 24.vii.1979, LK (ANIC). WA: Kununurra, 22.xii.1991-6.U992, RS (CBM, MDPI). - ?: CCXI (SAM). DIAGNOSIS. Antennae completely black; femora yellow; tibiae with contrastingly black apex, black, yellow ringed. SUPPLEMENTARY DESCRIPTION. Measure- ments (6 ex.). Length, 8. 6-9. 8mm; width, 3. 1-3. 55mm. Ratios: length eye/orbit, 2.0; lengthAvidth of head, 0.92-0.96; length/width of prothorax, 1.14-1.15; length/width of elytra, 1.63-1.66. 178 MEMOIRS OF THE QUEENSLAND MUSEUM Colour (Fig. 38C). Black, only labrum and mouthparts reddish. Antennae black. Femora yellow, apex contrastingly black. All tibiae black, with a contrasting yellow ring below middle. Tarsi black. Head . Eyes very large, markedly protruding, orbits short, moderately convex. Surface with 2 impressions behind clypeus, medially of eye with distinct ridge and sulcus that attain about the middle of eye. Posterior supraorbital seta located shortly behind eye, but difficult to distinguish within the elongate, erect pilosity. Antenna elongate, surpassing base of pronotum by c. 1.5 antennomeres. Two basal antennomeres glabrous, 3rd antennomere with some additional elongate setae below the ring of apical setae. Punctuation of surface coarse, though sparse, pilosity remarkably elongate, erect. Micro- reticulation absent, surface glossy. Pronotum. Rather short, compact, laterally gently convex, dorsal surface slightly convex. Lateral margin little convex, indistinct, very uneven, not ridge-like. Proepistemum widely visible from above. Sulcus inside margin wide and deep, covered by very coarse punctures and by an extremely dense fringe of elongate setae. Part of this fringe is characteristically directed laterally. Punctuation coarse, dense in basal half, anteriorly sparse. Microreticulation absent, surface glossy. Elytra . Moderately elongate, laterally slightly convex, dorsally depressed. In basal third with a distinct, transverse impression which becomes indistinct towards lateral margin, and with a minor though still distinct transverse depression shortly in front of apex. 7th interval tumid in apical third. Striae complete, well impressed and coarsely punctate till apex. Intervals convex, irregularly punctate. Apex convex, oblique, not or barely sinuate. Whole surface covered with dense, elongate, erect, grey hairs, lateral margin with conspicuous fringe of elongate hairs. Microreticulation absent, surface glossy. Hind wings present. Lower Surface. With moderately dense, on thorax also coarse to very coarse punctuation, impilose, glossy. Metepistemum very elongate >3 X as long as wide at apex, terminal abdominal sternum in female polysetose. Legs. Of average size. Tarsi not lobed, 5th tarsomere with a dense fringe of elongate setae below. Claws smooth. Male Genitalia (Fig. 29). Terminal abdominal stemite in middle slightly incised. Genital ring fairly elongate, moderately triangular, barely FIG 29. Neoeudalia nigra (Sloane). Male genitalia: aedeagus, parameres and genital ring (scale 0.25mm). asymmetric, suddenly narrowed to the narrow, very acute, triangular apex. Aedeagus compar- atively large, slender and elongate, moderately depressed, laterally barely sinuate, whole lower surface concave. Orificum short. Apex short, fairly wide, straight, gently knobbed, markedly turned to right, gently incised at right side. Folding of internal sac simple. Parameres of moderately dissimilar shape, left paramere much larger than right one, with rounded apex, right paramere elongate, with narrower, more angulately rounded apex. Female Genitalia (Fig. 32D). Stylomere 2 large, though comparatively short and stout, laterally little curved, with short, moderately acute apex. With 3 medium-sized ventrolateral ensiform setae, a small dorsomedian ensiform seta situated above middle of stylomere, and a single short nematiform seta arising from a groove in apical third. Base of stylomere 1 with 6-7 slender, elongate ensiform setae. Lateral plate with moderately dense fringe of elongate, stiff setae at apical rim. Variation. Little variation noted due to limited material. As some of the available specimens are old and damaged, little can be said about variation. DISCUSSION. Apparently the type(s) of this species is/are lost. One specimen was labeled ‘Holotype’by Darlington, though it is a topotype which is written on the determination label. Nevertheless, this specimen was labeled by T.G. Sloane himself and as it corresponds with the description, it is heredesignated neotype. DISTRIBUTION. S central NSW, N Vic, SE and NE Qld, N NT and WA adjacent to border. In Qld this species was only recorded in dry areas west AUSTRALIAN ODACANTHINE GROUND BEETLES 179 FIG. 30. Female stylomeres 1 and 2 (scales: 0.1mm). A, Anasis howittii Castelnau. B, Archicolliuris splendissimus sp. nov. C, Archicolliuris occipitalis sp. nov. D, Aulacolius triordinatus Sloane. E, Basistichus micans (Macleay). F. Clarencia aliena (Pascoe). G, Clarencia angust icollis (Macleay). H, Clarencia breviceps sp. nov. I, Deipyrus palustris (Sloane). J, Deipyrus inops sp. nov. K, Dicraspeda brunneipennis (Sloane). L, Dicraspeda obscura (Castelnau). M, Eudalia obliquiceps obliquiceps Sloane. N, Eudalia reticulata sp. nov. O, Eudalia atrata sp. nov. P, Eudalia castelnaui Sloane. ofthe Great Dividing Range. Sloane recorded the from N Qld, NT, and NW Australia were species ‘under loose bark of a Red Gum tree, captured at light. Probably a ground-dwelling having taken refuge from flood water', and ‘on species on the banks of rivers and lagoons, the muddy bank of Houlaghan’s Creek, ... a single example under a stick'. All specimens 180 MEMOIRS OF THE QUEENSLAND MUSEUM Ophionea Klug, 1821 (Fig. 38D) Ophionea Klug, 1821: 298; Sloane, 1917: 415; 1923: 30; Csiki, 1932: 1534; Licbke, 1938: 79; Darlington, 1968: 207; Moore et al.. 1987: 277; Lorenz, 1998: 417. Ophionaea Esehscholtz. 1829: 5. Casnoidea Castelnau, 1834: 40; Csiki, 1932: 1534; Darlington, 1968: 207; Moore et al., 1987: 277; Baehr, 1996b: 1044; Lorenz, 1998: 417. TYPE SPECIES. Cicindela cyanocephala Fabricius, 1798 (= Attelabus indicus Thunberg, 1784), by subsequent designation. DISCUSSION. The citation of Darlington (1968), Moore et al. (1987) and others of Ophionea as being first described by Esehscholtz (1829) is incorrect, as had been stated by Lorenz (1998). Bousquet (in Lobl & Smetana 2003) noted: 'The genus-group name Ophionea was first validated by Klug in 1821. Not only did he describe in length the genus but he included 3 available species: Attelabus pennsylvanicus Linnaeus, Cicindela cyanocephala Fabricius, and Attelabus surinamensis Linnaeus. The type species is Cicindela cyanocephala Fabricius, 1798 (= Attelabus indicus Thunberg, 1784), by subsequent designation of Hope ( 1838: 104). it is quite clear that Esehscholtz did not describe a new genus-group taxon under the name Ophionea but that he simply used Klug’s name under a different spelling’. The type species of Casnoidea is Cicindela cyanocephala Fabricius, 1798 (= Attelabus indicus Thunberg, 1784), by original designation. KEY TO AUSTRALIAN SPECIES OF OPHIONEA 1. Base of elytra reddish; prothorax coarsely punctate puncticollis Sloane Base of elytra dark; prothorax not coarsely punctate . . 2 2. Prothorax lacking lateral setae; each elytron with 2 small white spots in anterior third and near apex; dark basal fascia interrupted in middle indica (Thunberg) Prothorax with one pair of lateral setae; each elytron with one large white spot only near apex; dark basal fascia not interrupted in middle 3 3. Orbits markedly convex, eyes laterally markedly protruded; prothorax c. 1 .5 x as long as wide, widest in middle or in front of it; antenna short, median anten- nomeres<2 X as long as wide . . . . thouzeti Castelnau Orbits less convex or almost regularly oblique, eyes laterally less protruded; prothorax >1.67 X as long as wide, widest clearly behind middle; antenna elongate, median antennomeres at least 3 x as long as wide. . . 4 4. Surface of elytra not microreticulate, glossy; light subapical spot small, clearly surpassing posterior border of dark fascia; prothorax >2 x as long as wide . ♦ . australica Baehr Surface of elytra perceptibly microreticulate; light subapical spot very elongate, ending at posterior margin of dark fascia; prothorax <2 x as long as wide storeyi Baehr Porocara Sloane, 1917 (Fig. 38E) Porocara Sloane, 1917: 415; 1923: 30; Licbkc, 1938: 91; Baehr, 1986: 718; 1996c: 254; Moore et al., 1987: 273; Lorenz, 1998: 421. TYPE SPECIES. Pomcara punctata Sloane, 1917, by monotypy. KEY TO SPECIES OF POROCARA 1 . Surface of pronotum and posterior part of head impunctate or nearly so; postmedian vitta of elytra narrow, conspicuously serrate and sinuate; apex of aedcagus virtually not thickened glabrata Baehr Surface of pronotum and posterior part of head coarsely punctate; postmedian vitta of elytra wide, less serrate and sinuate: apex ofaedeagus more or less thickened . 2 2. Aedcagus smaller and shorter, length < 1 .8mm, apex not much upturned and barely curved to right: elytra shorter, laterally less regularly rounded; basal part of lateral borders of pronotum abruptly sinuate 3 Aedcagus larger and longer, length >2. 2mm, apex markedly upturned and curved; elytra longer, laterally regularly rounded; basal part of lateral borders of pronotum not abruptly sinuate 6 3. In fully coloured specimens prothorax, vertex, and most of lower surface black; in tcneral specimens at least part of proepipleurae and base of head darker than remaining parts; elytra markedly square, with accentuate humerus, lateral margins almost parallel; microsculpture of elytra very conspicuous, at 10 X magnification seemingly isodiametric; lower surface of aedeagus gently curved, apex elongate, not turned down. . . . nigricollis Baehr Even in fully coloured specimens prothorax, vertex, and most of lower surface not darker than remainder; elytra not square, with less accentuate humerus, lateral margins perceptibly convex: microsculpture of elytra less conspicuous, at 10 X magnification distinctly transverse; lower surface of aedeagus gently or markedly curved, apex short or elongate, but when elongate, then apex turned down or aedeagus markedly curved 4 4. Pronotum more densely and regularly punctate, basal part of lateral borders less abruptly sinuate; aedcagus compact, apical part at orificum (as seen from below) distinctly widened, lower surface gently or markedly concave, but when gently concave, then apex short. . . 5 Pronotum less densely and regularly punctate, basal part of lateral borders abruptly sinuate; aedeagus delicate, apical part at orificum (as seen from below) barely widened, lower surface gently concave, apex elongate punctata kimherleyana Baehr 5. Elytra shorter, more square, ratio l/vv1.62; lower surface of aedeagus markedly concave, apex longer. AUSTRALIAN ODACANTHINE GROUND BEETLES 181 distinctly club-shaped, pointed down punctata arnhemensis Baehr 6. Pronotum densely punctate; basal lobe of acdcagus less curved, apex more upturned though less asymmetric, lower surface of acdcagus barely striate. . ulrichi Baehr Pronotum more sparsely punctate; basal lobe of acdcagus more curved, apex less upturned though more asymmetric, lower surface of acdcagus conspicuously striate * occidentals Baehr Renneria Baehr, 1999 Renneria Baehr, 1999: 1 16. TYPE SPECIES. Renneria kamouni Baehr, 1999, by original designation. Renneria kamouni Baehr, 1999 (Fig. 38F) NEW RECORDS (6 ex.): NT: Tindal, 14.3 IS, 132.22E, 1-20. xii. 1967, WV (ANIC). - WA: Kununurra, 22.xii.1991-6.U992, RS (CBM, MDPI, QM). DISTRIBUTION. N NT, N WA adjacent to NT. Taken at light. THE AUSTRALIAN ODACANTHINE FAUNA The Australian odacanthine fauna includes 16 genera, 50 species, 5 subspecies and 2 doubtfully recorded species that most probably should be excluded from the Australian fauna (see checklist). Ophionea in die a, which is widely distributed in SE Asia, but not native to Australia has been accidentally introduced by ship. Numbers of indigenous taxa per state are: Vic - 5; ACT - 1; NSW - 7; Qld - 32; NT - 23; WA - 21; unknown - 1 . Therefore, Australian Odacanthinae are predominately tropical, are numerous in the wet tropics, but barely extend to the south or into the drier parts of the continent. They are most speciose in NE Qld, still quite speciose in N NT and N WA, becoming rarer in NSW, Vic and the ACT, and apparently completely absent from SA and Tas. In Vic Anasis howittii was recorded from near the coast, but has never been recaptured, so the record might be doubtful. Neoeudalia nigra and Myrm ecodem us riverinae were recently recorded from NW Vic. In WA south of the Great Sandy Desert, Gestroania amplipennis (Gestro), is known from the SW, Porocara occidentals and Eudalia waterhousei occur in the midwest between the Ashburton and Murchison Rivers. In NT only Eudalia waterhousei occurs in the interior, all others being restricted to the coastal North. In Qld only Eudalia latipennis interioris and Neoeudalia nigra , occur inland. Similarly, in NSW Neoeudalia nigra and Myrmecodemus riverinae , occur inland, i.e. west of Great Dividing Range, with the other five recorded species either in the northeastern part of the state or in the Australian Alps. In the south, only the streams of the Australian Alps harbour an odacanthine fauna worthy of mention (Framenau, Manderbach & Baehr, 2002), but the few species occurring there can be quite numerous in suitable areas. The range of Deipyrus inops , is so far completely unknown, although I suspect it might occur somewhere in the far north. It can be seen from the lists of examined material, that quite a large number of species, or even all species of certain genera, arc either rare, or are rarely collected, for whatever reasons. This particularly applies to species of Anasis , Archicolliuris , Gestroania , Giachinoana , Lachnothorax , Myrmecodemus , Neoeudalia and Renneria , but also to certain species of other genera. This may be due to either very restricted occurrence, very remote ranges, or to scarcity due to ecological factors. Indeed, only a few species are apparently common, which means: they have been collected either at many localities, or, if their range is more restricted, they are at least common where they occur. Examples of widespread and locally common species are BasStichus micans , Eudalia latipennis , E. waterhousei , Neoeudalia nigra , Ophionea thouzeti , whereas species like Deipyrus palustr is , Eudalia at rat a, E. castelnaui , E. macleayi , E. punctipennS and the members of Porocara are generally numerous where they occur. For most species, however, we do not yet know enough about distribution and habits to make any reliable decision. Nevertheless, some distribution patterns seem evident. In particular in the tropical northern half of Australia two patterns seem to exist: one comprising species of true wet tropical affinities that almost exclusively occur in wet NE Qld, in rain forest, either at the edge of rivers and lagoons (e.g. Archicolliuris spp., Dicraspeda brunnei- pennis , D. dubia , D. glabrata , D. longiloba , D. nitida , D. obscura , Lachnothorax tokkia ); and another group of species that inhabit savannah country or even semiarid areas and extend through the tropical belt of northern Australia, but do not occur east of the Great Dividing Range (e.g. Dicraspeda sublaevis , Eudalia punctipennis, E. waterhousei , Gestroania setipennis, G. storeyi , Giachinoana carinipennis , Myrmecodemus formicoides , M. globulicollis , M. lucai , Porocara 182 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3 1 . Female stylomeres 1 and 2 (scales 0. 1 mm). A, Eudalia macleayi Bates. B, Eudalia latipennis latipennis (Macleay). C, Eudalia n aterhousei Castelnau. D, Eudalia punctipennis sp. nov. E, Gestroania amplipennis (Gestro). F, Gestroania J'roggatti (Macleay). G, Gestroania setipennis sp. nov. H, Gestroania storeyi sp. nov. I, Giachinoana carinipennis Baehr. spp., Renneria kamouni). The ranges of the latter species are usually much more extensive and may extend from N NT to NW Australia, and some- times also to N Qld W of the Great Dividing Range. Neoeudalia nigra , however, has an unusual inland distribution pattern, extending from semiarid country in N Vic and S inland NSW to inland Qld and far N and NW Australia. In Qld it extends to mid-eastern Qld, but there only to the dry country belt that extends along the Tropic of Capricorn to the coast. With respect to habits, the Australian odacanthine fauna is unique in that most species apparently inhabit sand or gravel banks of rivers and brooks. Plant climbing, True’ odacanthines, on the other hand are much rarer in Australia compared with the faunas of South Asia, tropical Africa, and South America where they are common. This is easily understood, because wet, swampy habitats and rain forests are scarce in Australia compared with the other large continents. Since all Australian species are winged, they commonly come to light, either directly to the lamp or running about at the margins of the cone of light, though generally not before absolute darkness. It is clear that good flight ability is a prerequisite for beetles living on the margins of water bodies which regularly dry out, as is the case in Australian regions which are semiarid or have a long dry season. Very little is known about diet, feeding strategies, and life history of any Australian odacanthine species. I observed Eudalia and Porocara hunting on river banks at night and devouring larvae and adults of mayflies, but to my knowledge, no other information about diet has been recorded. I collected Eudalia I. latipennis , E. punctipennis , and E. waterhousei on fine sand and silt, whereas species of Porocara were mainly running amongst pebbles AUSTRALIAN ODACANTHINE GROUND BEETLES 183 FIG. 32. Female stylomeres 1 and 2 (scales: 0.1mm). A, Myrmecodemus formicoides (Sloane). B, Myrmecodemus lucai sp. nov. C, Myrmecodemus pilosellus sp. nov. D, Ncoeudalia nigra (Sloane). and coarse gravel. Porocara species are extremely fast runners. Ophionea species are true reed climbers as are many Archicolliuris , though apparently not the Australian species that inhabit rainforest, although we do not know where they live there. Except for the species of the Australian Alps (Eudalia castelnaui and E. macleayi)> almost all species seem to occur at low altitudes, although one species of Archicolliuris at least has been captured above 700m. Apart from these sparse details little is known about the other genera. Surprisingly many species occur some distance from water which may suggest rather a life in litter or even on low vegetation independent of open water bodies. Moore (1965) described the larvae of Eudalia macleayi Bates and noted their high similarity to non-Australian Colliuris. No other larvae of Australian Odacanthinae have been described. COMPOSITION OF THE FAUNA The Australian Odacanthinae includes very differently shaped species, with different ecological requirements, inhabiting remarkably diverse habitats. No other continent harbours, at the same time, highly evolved, elongate, long-necked reed-climbing species like those of Ophionea , Archicolliuris , and Clarencia; compact, short-headed, fast running, ground-living, nocturnal species like those of Porocara and Eudalia ; and extremely setose, globular species like those of Deipyrus and Myrmecodemus. Although the diversity of the Australian Odacanthinae is rather high, it is evident that highly developed rccd-dwelling species are comparatively rare, whereas primitive ground-living species (and genera) are numerous and speciose. Thus the Australian odacanthine fauna in general is plesiotypic, including probably the most basal extant odacanthine of all, Porocara. The strictly nocturnal Porocara occur with cicindclines (Megacephala s.l. spp., in particular species from the ripicolous crucigera- group), chlaeniines ( Chlaenius s.l. spp.) and brachinines ( Pheropsophus spp.) between pebbles, gravel, and sand on the banks of rivers in the Far North. This is perhaps one of the most basic habitats for tropical ground beetles. The serrate-cruciate, red and black colouration of the elytra and the very light colour of the legs are common colour patterns in nocturnal species living in such habitats. This colour pattern probably is well adapted to the reddish sand, as well as to the twilight. Porocara specimens running in the night appear like ghosts whisking along, or speaking more biologically, they resemble the fast running tiger beetles or lycosid spiders that occur in the same habitats. So, body shape and colour pattern in Porocara may also act as a sort of mimicry, without knowing which model Porocara imitates: aggressive spiders or tiger beetles, bombarding Pheropsophus , or distasteful Chlaenius. Grundmannius dispar Basilewsky, a South African chlaeniine beetle, has exactly the same body shape, colour pattern of the elytra (serrate-cruciate reddish-and-black), bulky head and prothorax, elongate legs and elongate mandibles, as Australian species of Porocara. Such similarities seem to corroborate the basic structure of the fauna of the sandy or gravely river banks in tropical, seasonally wet, or even semiarid country. Apart from colouration, certain species of Eudalia are also fairly similar in shape and structure to Porocara. These do not match our common concept of odacanthines, but also seem to represent a very basic stock within the subfamily. The species of the E. latipennis- waterhousei-Uneagc in particular are similar in their bulky body shape to Porocara , and they live in exactly similar habitats — sand and gravel of river banks in wet to dry savannah country or in 184 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 33. Habitus of Australian Odacanthinac (length in brackets). A, Anasis homttii Castelnau (8.1mm). B, Archicolliuris splendissimus sp. nov. (6.9mm). C, Archicolliuris occipitalis sp. nov. (10.5mm). D, Aulacolius triordinatus Sloane (7.6mm). E, Basistichus micans (Macleay) (6.8mm). F, Clarencia breviceps sp. nov. (7.9mm). semidesert. Dicraspeda , Neoeudalia , Anasis , Renneria and even Gestroania and Giachinoana , although the latter three combine species with several apomorpic features, are probably derived from an EudaliaAike stock, and most probably their habits are rather similar to those of Eudalia (ground-living at river banks). Some Australian genera are outstanding in certain morphological characters and their systematic position is not easily fixed. Habits of Aulacolius , Basistichus , Deipyrus and Myrmeco- demus are very little known, although probably they also live on the ground beside water. They all seem to be descended from primitive ancestors, having acquired certain more highly evolved characters. The oriental Lachnothorax , which also includes primitive, ground-living species, should also be added to this group. HAUSTRALIAN ODACANTHINE GROUND BEETLES 185 E FIG. 34. Habitus of Australian Odacanthinae (length in brackets). A, Clarencia quadridens Darlington (11.3mm). B, Deipyrus palustris (Sloane) (8.0mm). C, Deipyrus inops sp. nov. (8.5mm). D, Dicraspeda sublaevis (Macleay) (5.9mm). E, Dicraspeda obscura (Castelnau) (6.8mm). F, Eudalia obliquiceps tozeria subsp. nov. (8.2mm). Only three Australian genera belong to the highly evolved odacanthine lineage that matches our idea of typical odacanthines. Archicolliuris and Clarencia do not reach the extreme degree of adaptation in the structure of their tarsi to life on grass and reeds. However, Ophionea , with deeply excised 4th tarsomeres, dense clothing with elongate hairs on the lower surface of tarsi, and conspicuous, probably aposematic, red-and-black colouration, form the most evolved group within the subfamily. 186 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 35. Habitus of Australian Odacanthinae (length in brackets). A, Eudalia obliquiceps punctifrons subsp. nov. (8.5mm). B, Eudalia minor sp. nov. (6.9mm). C, Eudalia reticulata sp. nov. (8.6mm). D, Eudalia atrata sp. nov. (9.7mm). E, Eudalia femorata sp. nov. (9.8mm). F, Eudalia latipennis latipennis (Macleay) (8.4mm). Archicolliuris and Ophionea , however, are widespread in the Oriental Region and are recent immigrants from the north that reached Australia during, or perhaps even after, the Glacial Period ( Archicolliuris ), or at least not before the meeting of the Australian plate with the SE Asian insular belt in late Miocene. The Australian-New Guinean Clarencia is probably descendend from a ‘Co/Z/wm’ -like ancestor that may have also evolved during this period. HAUSTRALIAN ODACANTHINE GROUND BEETLES 187 FIG. 36. Habitus of Australian Odacanthinae (length in brackets). A, Eudalia latipennis interiors subsp. nov. (8.0mm). B, Eudalia waterhousei Castelnau (9. 1 mm). C, Eudalia punctipennis sp. nov. (8.7mm). D, Gestroania amplipennis (Gestro) (8.1mm). E, Gestmania froggatti (Macleay) (8.0mm). F, Gestroania setipennis, sp. nov. (8.4mm). Lachnothorax tokkia and Dicraspeda that in certain aspects is more evolved than any longiloba are recent immigrants to Australia and Australian members of the genus, the latter species, with its deeply excised 4th All other Australian genera are indigenous, tarsomercs, belongs to a group of New Guinean systematically rather isolated, and may have centred species within the diverse Dicraspeda originated in this continent. Considering the 188 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 37. Habitus of Australian Odacanthinae (length in brackets). A, Gestroania storeyi sp. nov. (8.8mm). B, Giachinoana carinipennis Baehr (7.1mm). C, Lachnothorax tokkia Gestro (5.6mm). D, Myrmecodemus globulicollis (Macleay) (7.0mm). E, Myrmecodemus riverinae (Sloane) (6.9mm). F, Myrmecodemus formicoides (Sloane) (4.9mm). remarkable plesiotypic structure of several genera, these may be remnants of a very old, basal odacan thine stock, or they may even represent the original stock. This would mean that Australia was where the evolution of odacanthines started. In this context, and if Porocara is the most plesiotypic genus, it should be kept in mind that the habitats where species of Porocara occur, namely banks of rivers in tropical wet and dry savannah, are very old ones, generally containing a very old fauna. Among other Carabidae, this Australian river bank fauna includes several plesiotypic genera, e.g. Megacephala (sensu lato) of Cicindelinae, Perileptus of Trechinae, Tasmanitachoides of Bembidiinae, Loxandrus of HAUSTRALIAN ODACANTHINE GROUND BEETLES 189 FIG 38. Habitus of Australian Odacanthinae (length in brackets). A, Myrmecodemus lucai sp. nov. (5.3mm). B, Myrmecodemus pilosellus sp. nov. (4.6mm). C, Neoeudalia nigra (Sloane) (9.8mm). D, Ophionea thouzeti Castelnau (6.6mm ). E, Porocara punctata kimberleyana Baehr (8.2mm). F, Renneria kamouni Baehr (6.8mm). Pterostichinac and others. In spite of the radical short-term substrate changes that occur periodically due to floods, this seems to be a very stable habitat that has perhaps not changed significantly in its structure through much of the Tertiary. The Australian odacanthine fauna is surprisingly speciose which is not expected in a continent that is exceptionally dry. The fauna is also surprisingly diverse in morphology and habits, and seems to include the most primitive existing odacanthines of all. Superimposed on 190 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 39. Structure of elytra. A, Eudalia waterhousei Castelnau. B, Eudalia punctipennis sp. nov. this ancient fauna are some ‘modern' groups that immigrated more recently from the Oriental Region, but have not yet spread beyond the wet northern tropical and eastern subtropical parts ol Australia where they remain restricted to rain forest or swampy areas. CHECKLIST OF AUSTRALIAN - ODACANTHINAE This checklist was added to update Moore et al. (1987) which is doubled from 25 to 50 species with 5 additional subspecies (Baehr, 1986, 1996b,c, 1999, 2000, 2003b, c; Framenau et al. 2002; this paper) and additional 2 species whose occurrence in Australia is doubtful. Oriental Ophionea indica Thunberg was accidentally introduced to WA by ship (Baehr, 1996b). For ease of use the checklists are alphabetical, and the most important generic synonyms are added. Colliuris has been divided into genera according to the systematic list of extant Carabidae of Lorenz (1998). References to the recorded range of the species are given as states, but with added N (northern), S (southern), W (western), E (eastern) where necessary. The ranges are compiled from information taken from Darlington (1968), Moore et al. (1987), from my papers mentioned above, and from my own collecting and determining experience. The only species of which I have not seen any Australian specimens are Archicolliuris par (Darlington) and Discraspeda dubia (Gestro), both New Guinean species that are claimed to occur in Australia at the northern tip of Cape York Peninsula. Genus Anasis Castelnau, 1867 howittii Castelnau, 1 867 SVic Genus Archicolliuris Liebke, 1931 occipitalis sp. nov N Qld [ par Darlington, 1 968 N Qld] splendissimus sp. nov N Qld Genus Aulacolius Sloane, 1923 triordinatus Sloane, 1 923 N Qld, N NT Genus Basistiehus Sloane, 1917 m/ccws(Macleay, 1864) .... NQld, NNT, NWA Genus Clarencia Sloane, 1917 a//e/?a(Pascoe, 1860) NENSW,E.Qld angw5//co//w(Maclcay, 1888) . NQld, NNT, NWA breviceps sp. nov NNT quadridens Darlington, 1968 NQld HAUSTRALIAN ODACANTHINE GROUND BEETLES 191 Genus Deipyrus Liebke, 1938 inops sp. nov ‘Australia’ palustris {Sloane, 1910) NQld Genus Dicraspeda Chaudoir, 1862 Macrocentra Chaudoir, 1 869 Loxocara Sloanc, 1907 Philemonia Liebke, 1938 [brunnea Chaudoir, 1 862)] [N Qld] brunneipennis (Sloane, 1917) N Qld dubia (Gestro, 1 879) N Qld glabrata Baehr, 2003 N Qld longiloba (Liebke, 1 938) N Qld nitida (Sloane, 1917) N Qld obscura (Castelnau, 1867) . . . NQld, N NT, NWA sublaevis( Macleay, 1888) . . . NQld, N NT, NWA Genus Eudalia Castelnau, 1867 atrata sp. nov CE NSW castelnaui Sloane, 1910 E Vic, S NSW femorata sp.nov CENSW latipennis (Macleay, 1 864) Qld /. latipennis (Macleay, 1 864) N Qld /. interioris subsp. nov SW Qld macleay i Bates, 1871 E Vic, ACT, S. NSW minor sp.nov NW Qld obliquiceps Sloane, 1917. . . . NQld, N NT, NWA o. obliquiceps Sloane, 1917 N Qld o. punctifrons subsp. nov. . . NW Qld, N NT, NWA o. tozeria subsp. nov NE Qld punctipennis sp. nov NW NT, NWA reticulata sp. nov N Qld waterhousei Castelnau, 1 867 NT, C WA Genus Gestroania Liebke, 1938 amplipennis (Gestro, 1875) SWA froggatti (Macleay, 1 888) NWA setipennis sp. nov N Qld, N NT storeyi sp. nov N NT, NWA Genus Giachinoana Baehr, 2003 carinipennis Baehr, 2003 N NT, N WA Genus Lachnothorax Motschulsky, 1862 Lasiocolliuris Liebke, 1931 tokkia Gestro, 1875 NQld Genus Myrmecodemus Sloane, 1923 Subgenus Myrmecodemus Sloane s. str. formicoides (Sloane, 1910) NNT globulicollis (Macleay, 1888) NNT, NWA lucai sp. nov N NT riverinae (Sloane, 1890) N VIC, SC NSW Subgenus Trichodemus subgen nov. pilosellus sp. nov N Qld Genus Neoeudalia gen nov. nigra (Sloane, 1900) N VIC, SC NSW, Qld, N NT, N WA Genus Ophionea Klug, 1821 Casnoidea Castelnau, 1 834 Subgenus Opltionea s. str. australica (Baehr, 1996) .... NQld, NNT, NWA [/W/ca (Thunberg, 1784)] .... [WA (introduced)] puncticollis Sloane, 1923 N Qld, N NT storeyi (Baehr, 1 996) N Qld, N WA thouzeti Castelnau, 1 867 E Qld, N NT Genus Porocara Sloane, 1917 glabrata Baehr, 1 986 N WA nigricollis Baehr, 1 986 N NT occidentalis Baehr, 1986 NWA punctata Sloane, 1917 N Qld, N NT, NWA p. punctata Sloane, 1917 N Qld p. arnhemensis Baehr, 1 996 N NT /;. Idmberleyana Baehr, 1986 .... NW NT, NWA ulrichi Baehr, 1996 N NT, N W A Genus Renneria Baehr, 1999 kamouni Baehr, 1999 N NT, N W A CHECKLIST OF ODACANTHINAE FROM NEW GUINEA, NEW BRITAIN, SOLOMON ISLANDS AND NEW HEBRIDES Darlington (1968, 1971) enumerated 9 genera (if Philemonia and Macrocentra are regarded as subgenera of Dicraspeda) and 21 species from New Guinea. Occurrences of Andrewesia apicalis , Ophionea indica , O. nigrofasciata, and Dicraspeda brunnea are doubtful because they have not been confirmed by specimens. Andrewesia apicalis , Ophionea indica , and O. nigrofasciata are Oriental species the occurrence of which was doubted by Darlington (1968, 1971) though the last two occur on Sulawesi (Baehr, 2003a). The doubtful status of Dicraspeda brunnea is discussed above. 10 genera with 28 species are recorded from New Guinea, 5 from New Britain, 3 from Solomon Islands (of which Dicraspeda inermis and Ophionea brandti are apparently endemic), and Dicraspeda hebridarum is endemic to the New Hebrides. Archicolliuris par , Basistichus micans , Clarencia quadridens , Dicraspeda dubia , Dicraspeda longiloba , Lachnothorax tokkia , Ophionea puncticollis , and Ophionea thouzeti are common to New Guinea and Australia. However, as demonstrated in Baehr (2000) and in the present paper, these faunas are not yet 192 MEMOIRS OF THE QUEENSLAND MUSEUM adequately recorded, and additional shared species will no doubt be found. Species in common between Australia and New Guinea are either Australian elements that are only found in southern New Guinea in environments similar to those of northern Australia ( Basistichus micans , Clarencia quadridens , Ophionea puncticoUis , O. thouzeti ), or they are northern elements originated in New Guinea ( Archicolliuris par , Dicraspeda dubici, D. longiloba), or even more widely through the Oriental region ( Lachnothorax tokkia). The checklist shows how little is known of the odacanthine fauna of W New Guinea (Irian Jaya) which is neglected in Darlington (1968) because at that time almost none had been collected there. Since then the situation has improved through the efforts of recent collectors (Baehr, 1995, 1996a, b, 1997b. 1998, 2003c), but the fauna is still far from documented. Little can be said about how well the faunas of the Bismarck Archipelago, Solomon Islands, and New Hebrides are documented, but they are apparently similarly incomplete. [Genus Andrew esia Liebke, 1938] [apical is (Chaudoir, 1872)]. . . . Genus Archicolliuris Liebke, 1931 ? papua (Darlington. 1 968) PNG par (Darlington, 1968) Genus Basistichus Sloane, 1917 PNG, New Britain micans (Macleay, 1864) Genus Clarencia Sloane, 1917 EPNG papua Darlington. 1 968 NG quadridens Darlington, 1968 . . . . Genus Crassacantha Baehr, 1995 NG bidens Baehr, 1995 Genus Dicraspeda Chaudoir, 1862 Macrocentra Chaudoir, 1 869 Loxocara Sloane, 1907 Philemonia Liebke, 1938 IJ bispinosa Darlington, 1968 . . . . PNG [brunnea Chaudoir, 1 862] .... ? denticulata Baehr, 1997 PNG dubia (Gestro. 1879) IJ hebridarum Baehr. 1 998 . . New Hebrides inermis Louwerens, 1970 . . . . Solomon Is intermedia Baehr, 1 997 IJ laticollis Baehr, 1997 IJ loebli Baehr, 1 996 PNG longiloba (Liebke, 1938) PNG, New Britain minuta Baehr, 1 998 PNG nigripes Baehr, 2003 PNG obsoleta Baehr, 1996 IJ: Biak I. papuensis Baehr, 2003 PNG quadrispinosa (Chaudoir, 1 869) NG, New Britain, Solomon Is ullrichi Baehr, 1 996 PNG v/o/am? (Sloane, 1907) NG, New Britain Genus Dobodura Darlington, 1968 armata Darlington, 1968 PNG Genus Eucolliuris Liebke, 1931 fuscipennis (Chaudoir, 1 850) PNG rossi (Darlington, 1968) PNG Genus Eudalia Castelnau, 1 867 anomala Darlington, 1 968 . . NG Genus Lachnothorax Motschulsky, 1862 Lasiocolliuris Liebke, 1931 tokkia Gestro, 1875 PNG Genus Ophionea Klug, 1 82 1 Casnoidea Castelnau, 1 834 brandtH Baehr, 1996) Solomon Is gestroi Maindron, 1910 PNG, New Britain [indica (Thunberg, 1 784)] ? [nigrofasciata Schmidt-Gobel, 1 846] ? puncticollis Sloane, 1 923 PNG thouzeti Castelnau, 1 867 PNG ACKNOWLEDGEMENTS I am greatly indebted especially to G.B Monteith (QM), E. Matthews (SAM). R. Storey (MDPI), K. Walker (MV) and T.A. Weir(ANIC) for the kind loan of ample and very important material and many types. Tom Weir is also thanked for his patient re-examination of certain types. I also thank the following colleagues for kindly providing me with types or material: S. Bily (Prague), R. Davidson (Pittsburgh), T Deuve (Paris), P.M. Giachino (Turin), S. Grove (Cairns), S. Hine (London), O. Merkl (Budapest), R. Poggi (Genoa), L. Toledano (Verona), M. Uhlig (Berlin) and L. Zerche (Eberswalde). LITERATURE CITED BAEHR, M. 1986. Revision of the Australian ground-beetle genus Porocara Sloane (Coleoptera: Carabidae: Odacanthinae). Australian Journal of Zoology 34: 717-731. 1995. A new genus of Odacanthinae from New Guinea (Insecta, Coleoptera, Carabidae). Spixiana 18: 45-48. 1996a. Three new species of the genus Dicraspeda Chaudoir from New Guinea (Insecta, Coleoptera. Carabidae, Odacanthinae). Spixiana 19: 137-146. 1996b. The ground beetle genus Casnoidea Castelnau. Taxonomy, phylogeny, HAUSTRALIAN ODACANTHINE GROUND BEETLES 193 zoogeography (Insecta, Coleoptera, Carabidae, Odacanthinae). Invertebrate Taxonomy 10: 1041-1084. 1996c. The Australian ground beetle genus Porocara Sloane. Second revision (Insecta, Coleoptera, Carabidae, Odacanthinae). Spixiana 19: 253-265. 1 996d. Two new species of the genus Lachnothorax Motschoulsky from the Philippines (Insecta: Coleoptera: Carabidae: Odacanthinae). Stuttgarter Beitrage zur Naturkunde. Serie A (Biologie): 539: 1-8. 1997a. A new species of the genus Casnoidea Castelnau from Java (Coleoptera, Carabidae, Odacanthinae). Entomofauna 18: 385-389. 1997b. Three further new species of the genus Dicraspeda Chaudoir from New Guinea (Coleoptera, Carabidae, Odacanthinae). Mitteilungen der Miinchner Entomologischen Gescllschaft 87: 29-37. 1998. Two further new species of the genus Dicraspeda Chaudoir from New Guinea and the New Hebrides (Insecta, Coleoptera, Carabidae, Odacanthinae). Entomofauna 19: 173-184. 1999. A new genus of Odacanthinae from northern central Australia (Insecta, Coleoptera, Carabidae). Coleoptera 2: 115-119. 2000. Some genera and species of ground beetles new to Australia (Coleoptera: Carabidae). Memoirs of the Queensland Museum 46: 9-14. 2003a. New taxa and new records of Odacanthinae from Sulawesi (Insecta, Coleoptera, Carabidae). Spixiana 26: 57-63. 2003b. A peculiar new genus of Odacanthinae from northern Australia (Insecta, Coleoptera, Carabidae). Monographic di Museo di Scienze naturale de Torino 35: 99-1 10. 2003c. A revision of the brunnea - group of the genus Dicraspeda Chaudoir (Coleoptera, Carabidae, Odacanthinae). Spixiana 26: 249-267 BATES, II. W. 1871. Notes on Carabidae, and descriptions of new species. (No. 5). Entomologists Monthly Magazine 8: 29-34. CASTELNAU, EL. de 1834. Etudes Entomologiques, ou Description d’Insectes Nouveaux et Observations sur leur Synonymie. Mequinon- Marvis, Paris. 94 pp. 1867. Notes on Australian Coleoptera. Royal Society of Victoria: 139 pp. 1 868. Notes on Australian Coleoptera. Transactions of the Royal Society of Victoria 8: 95-225. CHAUDOIR, M. de 1862. Materiaux pour servir a Fctude des Carabiqucs. 3e partie. Bulletin de la Soci£te Imperiale des Naturalistes de Moscou 35: 275-320. 1872. Observations sur quelques genres de Carabiqucs, avec la description d’especes nouvelles. Bulletin de la Societe Imperiale des Naturalistes de Moscou 45: 382-420. CSIKI, E. 1932. Coleopterorum Catalogus. Vol. III. Carabidae III: Pars 124, Harpalinae VII: 1279-1598, (W. Junk, Berlin). DARLINGTON, P.J. Jr 1968. The Carabid beetles of New Guinea. Part III. Harpalinae continued. Perigonini to Pseudomorphini. Bulletin of the Museum of Comparative Zoology 137: 1-253. 1971. The Carabid beetles of New- Guinea. Part IV. General considerations: analysis and history of the fauna. Taxonomic supplement. Bulletin of the Museum of Comparative Zoology 142: 129-337. ESCHSCHOLTZ, J.E 1829. Zoologischer Atlas, enthaltend Abbildungen und Beschreibungen neuer Thierarten wahrend des Flottencapitains von Kotzebue zweiter Reise um die Welt auf der Russisch-Kaiserlichen Kriegsschalupp Predpriaetie in den Jahren 1823-1826, II: 13 pp. Berlin: Reimer. FRAMENAU, V.W., MANDERBACH, R. & BAEHR, M. 2002. Riparian gravel banks of upland and lowland rivers in Victoria (south east Australia): Arthropod community structure and life history patterns in a longitudinal gradient. Australian Journal of Zoology 50: 103-123. GESTRO, R. 1875. Nota sopra alcuni Carabici. Annali di Museo Civico di Storia Naturale de Genova 7: 840-894. 1879. Descrizione di nuove specie di Coleotteri raccolte nella regione Austro-Malese dal signor L. M. D'Albertis. Annali di Museo Civico di Storia Naturale Giacomo Doria 14: 552-565. KLUG, J.C.F. 1821. Entomologiae Brasilianae specimen. Nova Acta Academiae Leopoldo Carlinae 10: 277-324. LIEBKE, M. 1931. Die afrikanischen Arten der Gattung Colliuris Degeer (Col. Car.). Revue de Zoologie et de Botanique Africaines, 20: 280-301. 1938. Denkschrift iiber die Carabiden-Tribus Colliurini. Festschrift fur Prof. Dr Embrik Strand 4:37-141. LOBL, 1. & SMETANA, A. (eds) 2003. Catalogue of Palaearctic Coleoptera 1. Archostemata - Myxophaga - Adephaga. (Apollo Books: Stenstrup). LORENZ, W. 1998. Systematic List of extant Ground Beetles of the World (Insecta Coleoptera “Geadephaga”: Trachypachidae and Carabidae inch Paussinae, Cicindelinae. Rhysodidae). , (Printed by the author: Tutzing). M ACLEAY, W.J. 1 864. Descriptions of new genera and species from Port Denison. Transactions of the Entomological Society of New South Wales 1 : 106-130. 1888. The insects of King’s Sound and its vicinity. Proceedings of the Linnean Society of New South Wales 3: 443-480. MOORE, B. P. 1965. Australian larval Carabidae of the subfamilies Harpalinae, Licininae, Odacanthinae 194 MEMOIRS OF THE QUEENSLAND MUSEUM and Pentagonicinae (Coleoptera). Proceedings of the Linnean Society of New South Wales 90: 157-163. MOORE, B.P., WEIR, T.A. & PYKE, J.E. 1987. Rhysodidae and Carabidae. Pp. 17-320. In Zoological Catalogue of Australia, 4. (Australian Government Publishing Service: Canberra). PASCOE, F.P. 1860. Notices of new or little-known genera and species of Coleoptera. Journal of Entomology 1 : 36-64. SLOANE, T.G. 1890. Studies in Australian Entomology. No. IV. New genera and species of Carabidae. Proceedings of the Linnean Society of New South Wales 5: 641-653. 1900. Studies in Australian Entomology. No. IX. New species of Carabidae (with notes on some previously described species, and synoptic lists of species). Proceedings of the Linnean Society of New South Wales 24: 535-584. 1910. Studies in Australian Entomology. No. XVI. New species of Carabidae. Proceedings of the Linnean Society of New South Wales 35: 378-406. 1917. Carabidae from tropical Australia (New genera and species, notes and synonymy, and synoptic tables. Tribes Scaritini, Harpalini, Odacanthini, Lebiini, and Helluonini). Proceedings of the Linnean Society of New South Wales 42: 406-438. 1923. Studies in Australian Entomology. No. XVI II. New genera and species of Carabidae (Scaritini, Pterostichini, Merizodini, Bembidiini, Trechini, Odacanthini, Panagaeini, Licinini, and Lebiini). Proceedings of the Linnean Society of New South Wales 48: 1 7-39. TEEATTA , A NEW SPIDER GENUS FROM TASMANIA, AUSTRALIA (AMAUROBIOIDEA: AMPHINECTIDAE: TASMARUBRIINAE) VALERIE TODD DAVIES Davies, V.T. 2005 01 10: Teeatta , a new spider genus from Tasmania, Australia (Amaurobioidea: Amphinectidac: Tasmarubriinae). Memoirs of the Queensland Museum 50(2): 195-199. Brisbane. ISSN 0079-8835. Three species of Teeatta gen. nov. from Tasmania are described. They are I driesseni (type species), T. magna and T. platnicki . They are placed with Tasmarubrius, Tasmabrochus and Tanganoides gen. nov. in the subfamily Tasmarubriinae. Tanganoides nov. nom., replacement name is provided here for Tangana Davies, 2003 preoccupied in the Orthoptera by Ramme, 1929:309. □ Amphinectidae, Tasmarubriinae, Teeatta, gen. nov., Tanganoides, nov nom. Valerie Todd Davies, Queensland Museum, PO Box 3300, South Brisbane 41 01, Australia; 31 March 2004. Teatta is the fourth genus of the Tasmarubriinae Davies (2002) to be described from Tasmania. There is sexual dimorphism shown in the colour and shape of the carapace and chelicerae. In males the carapace is longer than wide (1:07), the highest part is the foveal region; the chelicerae are geniculate (Fig. 1 A). In females the carapace is darker in colour and noticeably longer than wide (1:0.6), the highest part is mid-carapace; chelicerae are strongly geniculate (Fig. IB). MATERIAL AND METHODS Almost all the material for the type species was collected by Michael Driessen from pitfall traps (PF). The material for the second species, T. magna was hand-collected and contains no males. Further collecting may yield males and at the same time confirm that the paratype from Scotts Peak Dam Rd, south of the type locality, belongs here. Notation of spines follows Platnick & Shadab ( 1 975). The left male palp is described and illustrated. Measurements are in millimetres. ABBREVIATIONS. Museums : AMNH, American Museum of Natural History, New York; QM, Queensland Museum, Brisbane; TM, Tasmanian Museum and Art Gallery, Hobart. Morphology". The usual abbreviations are used for body measurements and eyes. Others in the text: MA, median apophysis; RTA, retrolateral tibial apophysis. SYSTEMATICS KEY TO GENERA OF TASMARUBRIINAE 1. 6 palp with long fixed prolateral tegular apophysis pointing forwards 2 6 palp with tegular apophysis arising in the membraneous area just prolateral toMA or occasionally absent. ... 3 2. 6 tegular apophysis about same length as MA. Large retrobasal paracymbium. 9 lateral epigynal protuberances Tasmarubrius 3 tegular apophysis very long, extending beyond tegulum, about 1.5 x MA. Small paracymbium. 9 without lateral epigynal protuberances . Tasmabrochus 3. 6 MA short, stout; terminally bifurcate or trifurcate. Tegular apophysis small or absent. 9 spcrmathecae anterior to gonopores. Spines on tarsi III, IV . . . Tanganoides 3 MA slender, unbranched. Tegular apophysis curving across tegulum to embolic base. 9 spermathecae dorsal to gonopores. Without spines on tarsi III, IV . . Teeatta Teeatta gen. nov. TYPE SPECIES. T .driesseni sp. nov. ETYMOLOGY. Tasmanian Aboriginal teeatta , forest. Gender is masculine. DIAGNOSIS. Large (6. 0-9.0) to very large ( 1 7.0- 1 8.0) ground living spiders. Male palp with a very broad grooved embolus and long slender median apophysis. A long sclerotised tegular apophysis, arising in the membranous area prolateral to the median apophysis, is crook-shaped distally and curves across the tegulum to the embolic region (Fig. 1C). This differs in origin and direction from the fixed prolateral tegular apophysis in Tasmarubrius and Tasmabrochus, which arises on the prolateral tegulum and runs forwards. MA is unbranched di ffering from that of Tanganoides . Like the other genera it also lacks the small tarsal spines found in Tanganoides. A prolateral protrusion on the subtegulum locks with the embolic base (Fig. IE). RTA with low dorsoretrolateral apophysis and large pointed dorsal apophysis. Spermathecae small, situated behind (dorsal to) 196 MEMOIRS OF THE QUEENSLAND MUSEUM gonopores; differing from Tanganoides where they are anterior to the gonopores. DESCRIPTION. Three-clawed ecribellates with an almost glabrous carapace, geniculate chelicerae, 2 retromarginal and 2 promarginal cheliceral teeth with an intermediate row of tiny denticles, see Davies (2003, fig. 2A,B). Preening combs on metatarsi II-IV. Short thick embolus, small membranous conductor, long movable M A and long sclerotised tegular apophysis. Small rounded paracymbium. Palpal tibia with distal retroventral apophysis and excavated RTA with 2 apophyses. Epigynum with ‘lateral’ teeth, emerging posteriorly rather than laterally (Fig. 1H). Gonopores often blocked with acellular material; small spermathecae. Teeatta driesseni sp. nov. (Fig. 1). ETYMOLOGY. For Michael Driessen who collected many spiders from the moorlands of central and western Tasmania. MATERIAL. HOLOTYPE: <3, Lake St. Clair, 42°07’S, 146° 1 1 E, PF, 25 Aug. 1999, M. Driessen (QM S55299). PARATYPES: 9 . same locality and collector as holutype, PF, 26 Mar. 1 997 (QM S55305); 25d,149 , 26 June 2000 (QM S55293); 6 , 1 7 9 , 26 Apr. 2000 (QM S55296); $ , 6 Mar. 1997 (QM S55297); d, 25 Aug. 1999 (QM S55298); 9. 29 Mar. 2000 (QM S55300); d, 30 Sept. 1999 (QM 55301): 3d, Aug. 1999 (QM S55302); 29, 25 Aug. 1999 (QM S55303); 9, 26 Mar. 1997 (QM S55304); 2d, 9, Aug. 1999 (QM S53306); <3, Aug. 1997(QMS55307); 6, Aug. 1999 (QM S55308); 2d, Aug. 1999 (QM S55309); d , Aug. 1999. (QM S55315); 9, penult, d. King William Creek, 42°14’S, 146°15’E, 25 Feb. 1999 (QM S55310); 9, 2 penult d, same locality, 25 Feb. 1999 (QM S55311); 29, 25 Feb. 1999 (QM S55312); 9, Mar. 1999 (QM S55313); 29, d, July 1999 (QM S55314). DIAGNOSIS. Tegular apophysis long, sclero- tised (Fig. 1C), arising from membraneous area prolateral to the median apophysis, curving across the tcgulum to the embolus. DESCRIPTION. Male. CL 3.9, CW 2.8, AL 3.6, AW 2.3. Light brown carapace, highest at fovea. Dorsal abdomen dark brown with pattern of 6 pairs of pale spots diminishing in size posteriorly. From above PRE slightly procurved, ARE straight; from the front both rows procurved. AME smallest. All eyes ringed in black. Ratio of AME: ALE: PME: PLE is 8: 1 1 : 1 0: 1 1 . Chelicerae geniculate: 2 retromarginal and 2 promarginal teeth with an intermediate row of small thin denticles between them. Labium slightly wider than long; sternum slightly longer than wide 1:0.9. Legs 4123 (Table 1). TABLE 1. T. driesseni d(9) Leg lengths. Leg I Leg II Leg III Leg IV Femur 3.0 (3.2) 2.5 (2.8) 2.3 (2.7) 3.0 (3.3) Patella &Tibia 3.8 (4.2) 3.0 (3.3) 2.7 (2.9) 3.7 (4.2) Metatarsus 2.5 (2.5) 2.1 (2.1) 2.3 (2.2) 3.3 (3.3) Tarsus 1.7 (1.6) 1.3 (1.3) 10(1.1) L3 (1.4) Total 11.0(11.5) 8.9 (9.5) 8.3 (8.9) 11.3(12.2) Notation of spines. Femora: I, D110, P001; II, duo, pool in, duo, pool, rool iv, duo, POOL ROOL Tibiae: I, PI0I, V222: II, Pill VI 12; III, D010, Pill. V212, R001; IV. D001, Pill, V212, Rill. Metatarsi: 1. P012.V221 R002; II, P012, V221, R012; 111, D010, P101, V221, R1 12; IV, D010. PI 12, V221, R 1 1 2. Spines on metatarsi III and IV very long and robust. No tarsal spines. Preening combs on metatarsi II-IV. Male palp (Fig. 1C-F). Cymbium with slight bulge on retrolateral edge, small rounded posterior protrusion and flanged paracymbium; trichobothria in dorsal row. Thick grooved embolus, small membranous conductor, long tapered MA and very long sclerotised tegular apophysis that curves across the tegulum to the embolic region. A prolateral protrusion on the subtegulum locks the embolic base in place. Tibia with ridge-like retroventral apophysis; RTA with low dorso-retrolateral apophysis and large pointed dorsal apophysis (Fig. 1D,E). Males range in length from 6.7-8.4. Female (QM S55305). CL 4.7, CW 3.0, AL 4.4, AW 2.8. Carapace dark brown, much longer than wide, 1:0.6, highest at midcarapace. From above eye rows slightly procurved, from front both rows procurved. AME: ALE: PME: ALE 7:1 1:10:1 1 . helicerae strongly geniculate. Labium and sternum both slightly longer than wide. Legs 4 1 23 (Table 1 ). Notation ol spines. Differs from male in the following leg segments. Femora: IV, D 1 1 0 ROOL Tibiae: I, V222; II, PI 1 1, V122; III, DOlo' Pill, V212, R 1 0 1 . Metatarsi: I, P001 V22l’ R001; III, D010, PI 12, V221, R112. Two preening combs on metatarsi II-IV on either side ot distal spine. Epigynum (Fig. 1G-J) about 1/6 length ol the venter. Lateral gonopores with irregular shaped plugs. Lateral teeth. Small spermathecae dorsal to gonopores. Females 8. 1-9.4 long. DISTRIBUTION. Central Plateau of Tasmania. NEW SPIDERS FROM TASMANIA 197 FIG. 1 . A-J, Teeatta driesseni sp. nov. A,B, lateral carapace (8 9 ); C-F, 8 palp (ventral, retrolateral, prolateral, dorsal); G-J, epigynum (ventral, lateral, dorsal, posterior), da = dorsal apophysis of RTA; dra = dorso-retrolateral apophysis of RTA; fd = fertilisation duct; It = lateral tooth; ma = median apophysis; pc = paracymbium; rva = retroventral tibial apophysis; sp = spermatheca; ta = tegular apophysis. 198 MEMOIRS OF THE QUEENSLAND MUSEUM Teeatta inagna sp. nov. (Fig. 2A-C) ETYMOLOGY. Latin magmis , large. MATERIAL. HOLOTYPE: $ , Fem Tree, Mt Wellington, SE Tasmania, 42°55’S, 147 0 16’E, 22 Apr. 1952, R. Cunningham (TM J151). PARATYPES: 9, Mt Wellington near Springs, 42°54’S, 147°14’E, unknown collector (TM .1152); 9. Scotts Peak Dam Rd. SW Tasmania, 43°00S, 146°22E, 26 Apr. 1987, R.J.Raven (QM S55295). DIAGNOSIS. Much larger (16.0-18.0) than T. driesseni. Epigynum about 1/10 length of venter. Ventral spines on tibia III and IV V202, differing from T. driesseni which has V212. DESCRIPTION. Female. CL 8.1, CW 5.7, AL 10.1, AW 6.5. Carapace dark brown, highest at midlength. Abdominal pattern similar to T. driesseni. From above and in front eye-rows procurved. AME:ALE:PME:PLE 9:14:11:12. Chelicerae strongly geniculate; 2 retromarginal and 2 promarginal teeth with intermediate row of small denticles between them. Labium longer than wide 1 :0.9. Sternum longer than wide 1 :0.8. Legs 4123. 1, 17.5; II, 14.3; III, 13.9; IV, 17.8. Notation of spines. Femora: I, D110, P001; II, D110, POOL III, D100. POOL R011; IV, D210, POO 1, R00 1 . Tibiae: I, V222; II, V222; III, D00 1 , Pill, V202, ROIL IV, Pill, V202. ROIL Metatarsi: I, POOL V222, R00L II, P01 1, V221, R001; III, D100, PI 12, V221, R112; IV, D110, PI 12, V221, R112. Two preening combs on ventral metatarsi II-1V on either side of distal spine. Epigynum (Fig. 2A-C) about 1/10 length of venter. The male is unknown. DISTRIBUTION. Southern Tasmania. Teeatta platnicki sp. nov. (Fig. 2D-I) ETYMOLOGY. For Norman Platnick , co-collector of the paratype and author of many Australian spiders. MATERIAL. HOLOTYPE: 6. W of Strath cordon, just N of Mt Sprent, SW Tasmania, 42°65’S, 146°04 , E, 290m, 26Apr.l987, N. Platnick, R.J.Raven, T.Churchill (QM S64277). PARATYPES: 9, juv, Scotts Peak Dam Rd, 43°00 , S, 146°22'E, 26Apr.l987, R.J.Raven (QM S55294); d, Olga Valley, SW Tasmania, 42°43’S, 145°47 , E, litter. 31 Jan. 1977, L.Hill et al. (TM J1454). DIAGNOSIS. Much smaller than T. magna. Tegular apophysis with much longer distal branch (Fig. 2E) than T. driesseni. Continuous sclerotisation of RTA from retrolateral to pointed dorsal branch unlike T. driesseni where these are separate entities. DESCRIPTION. Mate. CL 3.6, CW 2.8, AL 3. 1 , AW 1.8. Carapace light brown, abdominal pattern similar to other species. Eyes and chelicerae similar to T. driesseni. Legs 4123. I, 9.8; II, 8.3; III, 7.6; IV, 10.3. Notation of spines. Femora: I, D110, P002; II, Dill, P01L III, Dill, POOL IV, D110, P001, R001. Tibiae: I, PI 1 L V222; II, PI 1 1, V122; III, D010, P20L V212, R101; IV, D001, PI 1 1 , V 2 1 2 , Rill. Metatarsi: I. P01 1, V221, R012; II, D010, P012, V22LR012; III.DOIO, PI 12, V221, R1 12; I V, DO 1 0, P 1 1 2, V22 1, R 1 1 2. Preening combs on metatarsi II-IV. No tarsal spines on legs. Male palp (Fig. 2D-F). Short widely grooved embolus; membraneous conductor; long sinuous MA. Tegular apophysis crook-shaped, with digitiform branch arising midway. Tibia with retroventral ridge; RTA with continuous sclerotisation between low dorsoretrolateral and long pointed dorsal apophysis. Paratype male from Olga Valley is 5.8 long, colour and pattern faded. Female. CL 3.5 CW 2.6 AL 3.8 AW 2.5. Coloration, pattern and eyes similar to that of males. Chelicerae strongly geniculate. Legs 4123.1, 8.8; II, 7.3; III, 6.8; IV, 9.4. Notation of spines. Fewer spines than in male. Femora: II, D1 10, POOL Tibiae: I, P000; II, P001, V222; III, P201; IV, R001. Metatarsi: I, P001, R001; II, D000, R001. Preening combs as in male. No tarsal spines. Epigynum (Fig. 2G-I) similar to T. driesseni. Lateral teeth not obvious. DISTRIBUTION. SW Tasmania. ACKNOWLEDGEMENTS I thank Elizabeth Turner, Curator of the Tasmanian Museum and the Curator of Arachnids, American Museum of Natural History for loan of material for this study. I am grateful to Michael Driessen for his donation of spiders to the Queensland Museum. Thanks also to the Council of the Australian Biological Resources Study for its financial support of illustrator, Catherine Harvey. Special thanks to John K. Page, Zoolological Record for indicating the homonymy of Tangana with an orthopteran. LITERATURE CITED DAVIES, V.TODD 2002. Tasmabmchus, a new spider genus from Tasmania, Australia (Araneae, Amphinectidae, Tasmarubriinae). Journal of Arachnology 30: 219-226. NEW SPIDERS FROM TASMANIA 199 FIG. 2. A-I, Teeatta spp. nov. A-C\ T magna epigynum (ventral, dorsal lateral); D-I, T. platnicki ; D-F, — Australosuchus clarkae Baru wickeni 20 FIG 6. Graph showing the log transformed width of alveoli in closely related early crocodyloids. Specimens based on: Kcimbara murgonensis (QMF29666); K. implexidens (QMF21116); K. molnan ; (QMF 12364); Runcorn crocodylomorph ( UQF73 1 95); Baru wickeni (NTMP86 1-14); Australosuchus clarkae (QMF 1 8 1 52). Reproduced from Salisbury (1994). Willis, 1996) following recognition of K. implexidens. In relation to QMF 12364, these diagnostic features include: medium sized, moderately broad-snouted; mandibular symphysis extending caudally to between the 6th and 7th dentary alveoli; splenial extending anteriorly to between the 6th and 7th dentary alveoli but not entering the mandibular synthesis; 1 7- 1 8th dentary alveoli; teeth slightly compressed transversely with distinct anterior and posterior carinae. Salisbury & Willis (1996) listed 3 other features of the mandible of K. implexidens that they thought diagnostic of the genus pending collection of additional material of K. murgonensis : articular fossa deeply notched laterally; short retroarticular process; medial wall of retroarticular fossa concave. In dorsal view, the preserved tooth row of QMF 12364 is in the same plane as the caudal half of the mandibular ramus. This indicates a generalised brevirostrine skull, with proportions similar to the holotype skull of K. murgonensis (as opposed to a shorter rostrum, where the medial angle between the tooth row and the caudal half of the ramus would be considerably <180°). Similar to K. murgonensis and K. implexidens , QMF 12364 has 17 dentary alveoli. The morphology of the teeth and the pattern in the width of the dentary alveoli up to the 9th alveolus (Fig. 6) are also similar. In K. implexidens , the 10th and 11th alveoli are approximately the same size, with the remainder being considerably smaller. In K. mutgonensis and QMF 12364, the 10th alveolus is smaller than the 11th and 12th. The 1 1th and FIG. 7. Kambara molnari sp. nov., holotype. Dorsal view of the angular and the caudal portion of the surangular, showing the two surangular pits and the rostrocaudally aligned ridge. Rostral is to the left. 214 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 8. Kambara molnari sp. nov., holotype. Medial view of the splenial. Broken line indicates probable splenial continuation. Rostral is to the right. 12th are also similar in size, and there is a corresponding decrease in size in the caudal most portion of the tooth row. The major difference between K. murgonensis and QMF 1 2364 is in the middle of the dentary (the 10th- 12th alveoli). In K. murgonensis , the 10th alveolus is closer in size to the 9th, whereas in QMF 12364, the 1 1th is closer in size to the 10th. The mandibular rostrum is not preserved in QMF 12364. Nevertheless, the splenial extended rostrally to a point that is at least level with the caudal margin of the 6th dentary alveolus. Such a condition is consistent for Kambara. In light of these similarities and its close geographical and temporal occurrence, we feel confident in assigning QMF 12364 to Kambara. DIAGNOSTIC OSTEOLOGICAL FEATURES. Differences between K. molnari and other material assigned to Kambara that warrant specific designation include the following. Occlusal Pattern. A major distinguishing features between K. murgonensis and A. implexidens relates to the way in which the jaws occlude (Salisbury & Willis, 1996). In K. murgonensis, the dentary teeth occlude medial to the rostral teeth, whereas in K. implexidens they interlock (Salisbury & Willis, 1996). Kambara molnari shows what could be considered an intermediate condition. The margins ot the 7th and 8th alveoli in K. molnari are raised slightly (Fig. 4), with occlusal grooves for maxillary teeth on the lateral surface of the dentary, at positions midway between adjacent alveoli. This indicates that at least in this portion of the rostrum, there was interlocking dentition with the rostral-most maxillary teeth. Reception pits between some of the caudal-most alveoli ( 1 2th- 1 7th) also suggest that the teeth interlocked in this part of the rostrum. On the other hand, the 10th- 12th alveoli are confluent (Fig. 5), indicating that in this part of the rostrum, the maxillary teeth occluded lateral to those of the dentary. The resulting inferred occlusal pattern is therefore different to those present in either K. murgonensis and K. implexidens , and is more reminiscent of that seen in several types of Palaeogene crocodyloids and some species of Borealosuchus ( Leidyosuchus ) (Brochu, 1 997) and Diplocynodon (Brochu, 1 999). Osteology > of the Retroarticular Process and the Articular. The articular and associated retroarticular process is not well known for Kambara. The only other specimen besides QMF 1 2364 to have part of the articular preserved is the paratype mandible of K. implexidens (QMF30077) (Salisbury & Willis, 1996, figs 8, 9). Based on this specimen, Salisbury & Willis (1996) considered a short retroarticular process as possibly diagnostic of the genus. In K. molnari , the retroarticular process is approximately 3 X as long in a rostrocaudal direction as it is wide in dorsal view; a condition that could not be described as ‘short 1 . Salisbury & Willis (1996) also noted that unlike K. implexidens , the retroarticular fossa of A. molnari was divided into 2 smaller fossae by a sagittally aligned ridge (Fig. 8), similar to the condition seen in Cwcodylus. It has also been suggested that the medial wall of the retroarticular fossa being concave may be diagnostic of Kambara (Salisbury & Willis, 1996). This hypothesis is again based on the paratype mandible of K. implexidens. K. molnari also shows this feature (Fig. 7). Therefore while still diagnostic, this feature is no longer restricted to K. implexidens. Surangular Pits. The presence of 2 small pits on the surangular of K. murgonensis was used as a diagnostic feature of that species by Salisbury & Willis (1996). Similar pits occur in K. molnari (Fig. 7). There was no mention in the description of Australosuchus clarkae of these pits (Willis & PALAEOGENE CROCODILIAN FROM QUEENSLAND 215 Kambara murgonensis Kambara implexidens Kambara molnari — — Runcorn crocodylomorph - Australosuchus clarkae Baru wickem FIG 9. Graph showing the log transformed interalveolar distances of the holotype of Kambara molnari (QM FI 2364) and other closely aligned crocodyloids. Specimens based on: Kambara murgonensis (QMF29666); K implexidens (QM F2 1 1 16); K. molnari (Q MF 12364); Runcorn crocodylomorph (UQF73 195); Baru wickeni (NTMP861-14); Australosuchus clarkae (QMF18152). Reproduced from Salisbury (1994). Molnar, 1991b). These pits in a closely related genus may indicate that K. implexidens was more plesiomorphic than the other 2 species (as it lacks these pits). We have revised the diagnostic value of these pits, so as to not be confused with the heavy sculpture pitting also present on the surangular. TAXONOMIC CONCLUSIONS. Based on features discussed above, we erect Kambara. molnari sp. nov. and distinguish it by: 1) 17 dentary alveoli; 2) 2 shallow, rostrocaudally elongate pits lateral to the articular fossa on the dorsolateral surface of the surangular; 3) retroarticular process approximately 3 x longer than it is wide in dorsal view; 4) retroarticular fossa divided into 2 fossae by a shallow, sagittally aligned ridge; 5) occlusal grooves for the reception of maxillary pseudocanines lateral to a point midway between the 7th and 8th, and the 8th and 9th alveoli; 6) reception pits for maxillary teeth between the 1 2th- 1 7th alveoli;7) 10th- 12th alveoli confluent. DISCUSSION The recognition of a third species of Kambara at Rundle, near Gladstone on the central Queensland coast, extends the range of this genus approximately 250km farther north. Previously, Boat Mountain, near Murgon was the only recorded location for Kambara. Although it is unlikely that K. molnari was contemporaneous with K. murgonensis and K. implexidens , its discovery points to high species level diversity for Kambara during the Eocene. Similarities between the depositional environments in which all three species were found additionally indicate that like other mekosuchines, Kambara was probably restricted to freshwater habitats. During the early Tertiary, it seems likely that species of Kambara were found throughout the inland waterways of Queensland, if not all of eastern Australia if temperature conditions permitted. This distribution is similar to that of Crocodylus johnstoni in northern Australian today. Kambara molnari shows features of both K. implexidens and K. murgonensis. Salisbury & Willis (1996) considered the interlocking dentition seen in K. implexidens to be convergent on the condition found in crocodylids and closely allied taxa such as Brachyuranochampsa and Harpacochampsa camfieldensis . In this light, the overbite seen in K. murgonensis and other mekosuchines was considered the ancestral condition for the group. However, the partially interlocking occlusal pattern in K. molnari suggests that the condition in K. implexidens is not as unusual as initially thought. UQF73195, a rostral dentary fragment from the early Tertiary (late Palaeocene or early Eocene) of Runcorn, SE Queensland also shows features indicative of an 216 MEMOIRS OF THE QUEENSLAND MUSEUM interlocking occlusal pattern (Salisbury & Willis, 1996; Willis & Molnar, 1991a). As was suggested by Salisbury & Willis (1996) and later supported by the phylogenetic analysis of Brochu (2001) (based on Australosuchus clarkae )* the interlocking occlusal pattern (or at least partial) in 3 taxa of early Tertiary Australasian croc- odilians may indicate a close taxonomic affinity between Mekosuchinae and Crocodylidae. The sagittally divided retroarticular fossa in K. molnari is a feature only known for Crocodylus (Norell & Clark, 1990) and possibly Crocodilus ’ clavis (Norell & Storrs, 1998). In extant crocodilians, the retroarticular fossa serves as the point of insertion for the m. depressor mandibulae, the primary muscle responsible for jaw opening (Busbey, 1989; Schumacher, 1973). The low, sagittally aligned ridge on the retroarticular fossa, as in Crocodylus and K. molnari , suggests that the insertion of this muscle may have been partially divided. As to whether such a division influenced the action ot the muscle during contraction is unclear, and it may or may not be related to an interlocking occlusal pattern. The fact that a rostrocaudally divided retroarticular fossa is absent in several other taxa that possess an interlocking occlusal pattern (e.g., Tomistoma schlegelii and Gavialis gangeticus) suggests that this is not the case. It is hoped that the collection of further crocodilian material from the Rundle Formation will help elucidate any similarities between Kambara and crocodylids, shedding much needed light on the origins of mekosuchines. ACKNOWLEDGEMENTS We are grateful to Alex Cook and other staff in the Geosciences Program, Queensland Museum for access to material in their care. Further thanks must be given to the two anonymous reviewers of this report. Financial support for SWS at the University of Queensland was provided by the Australian Research Council Linkage-Project scheme (LP0347332), in association with the Queensland Museum, Land Rover Australia, Winton Shire Council and Isisford Shire Council. LITERATURE CITED BALL, L.C. 1914. Tertiary oil shales of The Narrows, Port Curtis District. Queensland Government Mining Journal 15: 73-76. 1946. Oil shales of The Narrows, central Queensland. Queensland Government Mining Journal 47: 176-179. BALOUET, J.C. & BUFFETAUT, E. 1987. 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Memoirs of the Queensland Museum 39: 711-717. RIEK, E.F. 1952. The fossil insects of the Tertiary Redbank Plains Series. Part 1 : an outline of the fossil assemblages with descriptions of the fossil insects of the orders Mecoptera and Neuroptera. University of Queensland Papers: Department of Geology. 4: 3-14. ROSSMAN, T. 2000. Skelerranatomomische Beschreibung von Pristichampus rollinatii (Gray) (Crocodilia, Eusuchia) aus dem Palaogen von Europa, Nordamerika und Ostasien. Courier, Forschungsinstitut Senckenberg 221: 1-107. SALISBURY, S.W. 1994. Interspecific variation, palaeoecology and phylogenetic relationships of early Eocene crocodylids from Murgon, southeastern Queensland. Unpubl. Honours thesis. University of New South Wales, Sydney. 2001. A biomechanical transformation model for the evolution of the eusuchian-type bracing system. Unpubl. PhD thesis. University of New South Wales, Sydney. SALISBURY, S.W. & FREY, E. 2001. A biomedical transformation model for the evolution of semi-spheroidal articulations between adjoining vertebral bodies in crocodilians. Pp. 85-134. In Grigg, GC., Seebacher, F. & Franklin, C.E.(eds) Crocodilian biology and evolution. (Chipping Norton: Sydney). SALISBURY, S.W. & WILLIS, P.M.A. 1996. A new Crocodylian from the early Eocene of south-eastern Queensland and a preliminary investigation of the phylogenetic relationships of crocodyloids. Alchcringa 20(3-4): 179-226. SALISBURY, S.W., FREY, E., MARTILL, D.M. & BUCHY, M.C. 2003. A new crocodilian from the Lower Cretaceous Crato Formation of north-eastern Brazil. Palaeontographica 270: 3-47. SALISBURY, S.W., WILLIS, P.M.A., PEITZ, S. & SANDER, P.M. 1999. The crocodilian Goniopholis sinnis from the Lower Cretaceous of north-western Germany Pp. 121-148. In Unwin, D.M.(ed.) Cretaceous Fossil Vertebrates - Special papers in Palaeontology. (The Palaeontological Association: London). SCHUMACHER, GH. 1973. The head muscles and hylaryngeal skeleton of turtles and crocodilians. Pp. 101-199. In Gans, C. & Parsons, T.S. (eds) Biology of the Reptilia. (Academic press: New York). SMITH, J.B. & DODSON, P. 2003. A proposal for a standard terminology of anatomical notation and orientation in fossil vertebrate dentitions. Journal of Vertebrate Paleontology 23: 1-12. VICKERS-RICH, P.V. 1996. Early Cretaceous polar tetrapods from the Great Southern Rift Valley, southeastern Australia. Memoirs of the Queensland Museum 39: 719-723. WILLIS, P.M.A. 1993. Trilophosuchus rackhami gen. et sp. nov. A new crocodilian from the Early Miocene limestones of Riversleigh, northwestern 218 MEMOIRS OF THE QUEENSLAND MUSEUM Queensland. Journal of Vertebrate Paleontology 13: 90-98. 1995. The phylogenetic systematics of Australian crocodilians. Unpubl. PhD thesis, University of New South Wales, Sydney. 1997. New Crocodilians from the Late Oligocene White Hunter Site, Riversleigh, northwestern Queensland. Memoirs of the Queensland Museum 41(2): 423-238. 2001. New crocodilian material from the Miocene of Riversleigh. Pp. 64-74. In Grigg, G.C., Seebacher, F. & Franklin, C.E. (eds) Crocodilian biology and evolution. (Surrey Beatty & Sons: Chipping Norton). WILLIS, P.M.A. & MACKNESS, B.S. 1996. Quinkana babarra , a new species of ziphodont mekosuchine crocodile from the Early Pliocene Bluff Downs Local Fauna, northern Australia with a review of the genus. Proceedings of the Linnean society of New South Wales 116: 143-151. WILLIS, P.M.A. & MOLNAR, R.E. 1991a. A longi- rostrine crocodile from the Early Tertiary of southeastern Queensland. Alcheringa 15: 229-233. 1 99 1 b. A new Middle Tertiary Crocodile from Lake Palankarinna, South Australia. Records of the South Australian Museum 25: 39-55. 1997. A review of Plio-Pleistocene Crocodilian genus Pallimnarchus. Proceedings of the Linnean Society of New South Wales 177: 223-242. WILLIS, P.M.A., MOLNAR, R.E. & SCANLON, J.D. 1993. An early Eocene crocodilian from Murgon, southeastern Queensland. Kaupia 3: 27-33. WILLIS. P.M.A., MURRAY, P. & MEGIRIAN, D. 1990. Bam darrowi gen. et. sp. nov., A large, broad-snouted crocody line (Eusuchia: Crocodylidae) from the mid-Tertiary freshwater limestones in Northern Australia. Memoirs of the Queensland Museum 29(2): 521-540. WITMER, L.M. 1995. Homology of facial structures in extant Archosaurs (Birsds and Crocodilians), with special reference to paranasal pneumaticity and nasal conchae. Journal of Morphology 225: 269-327. NEW CHAROPID LAND SNAILS CHIEFLY FROM LIMESTONE OUTCROPS IN NSW (EUPULMONATA: CHAROPIDAE) ISABEL T. HYMAN AND JOHN STANISIC Hyman, I.T. & Stanisic, J. 2005 01 10: New charopid land snails chiefly from limestone outcrops in eastern New South Wales (Eupulmonata: Charopidae). Memoirs of the Queensland Museum 50(2): 219-302. Brisbane. ISSN 0079-8835. Nineteen new species of Charopidae are described from mainly limestone habitats in eastern New South Wales. Eleven new genera are introduced and some existing genera and species revised. Rhophodon Hedley, 1924 is more clearly defined and R. palethorpei sp. nov., R. silvaticus sp. nov., R. mcgradyorum sp. nov. and R. duplicostatus sp. nov. are described from limestone and rainforest habitats in NE NSW; Egilodonta Iredale, 1937 (type species: Charopa bairnsdalensis Gabriel, 1930) is removed from the synonmy of Rhophodon and E. wyanbenensis sp. nov., E. bendethera sp. nov. and E. paucidentata sp. nov. arc described from restricted limestone localities in SE NSW: Letomola lanalittleae sp. nov. is described from restricted limestone localities in the Macleay Valley. NE NSW; Macrophallikoropa gen. nov. is introduced for the widespread, moist forest dweller Helix belli Cox, 1 864 and M. stenoumbilicata sp. nov. and M. depressispira sp. nov. are described from restricted localities in the Sydney Basin Bioregion; Allocharopa Iredale, 1937 (type: Helix brazieri Cox, 1 868 from the Sydney Basin Bioregion) is shown to be a synonym ot Elsothera Iredale, 1933; Egilomen Iredale, 1937 is redefined and a new diagnosis is presented for the type, E. cochlidium (Cox, 1868), which is restricted in distribution; Whiteheadia gen. nov. is introduced for Egilomen globosa Stanisic, 1990; Coricudgia wollemiana gen. et sp. nov. is described from Mt Coricudgy, Wollcmi NP; Decoriropa gen. nov. is introduced for the widespread Helix lirata Cox, 1 868; Marilyniropa jenolanensis gen. et sp. nov. is described from limestone localities near Jenolan, SE Oberon; Cralopa Iredale, 1941 is redefined and revised diagnoses and new distribution details are presented tor the widespread C. stwudensis (Cox, 1 864) and C. kaputarensis Stanisic, 1 990 from the Nandewar Range; Gouldiropa gen. nov. is introduced for C. carlessi Stanisic, 1990 which is redefined and restricted in distribution to the northern New England Tableland; Sharniropa gen. nov. is introduced for S. wollondillvana sp. nov. and 5. borenorensis sp. nov. from limestones in the Abercrombie and Orange areas of central NSW respectively, and S. xanana sp. nov. from limestone outcrops in the southern New' England Tableland; Acheronopa attunga gen. et sp. nov. is described from limestones in the Manilla-Attunga region of the New' England Tableland; Hedleyropa yarrangobillyensis gen. et sp. nov. is described trom the Yarrangobilly limestones of SE NSW:* Scelidoropa gen. nov. is introduced for the widespread S. sarahjaneae sp. nov. and the narrowly retricted S. nandewar sp. nov. trom the Nandewar Range; Diphyowpa gen. nov. is introduced for Helix saturni Cox, 1868 from the Sydney Basin Bioregion and D. macleayana sp. nov. trom the Macleay Valley, NE NSW. Patterns of morphological variation are analysed and discussed and a phylogenetic hypothesis for the species is presented based on 32 shell characters and 12 anatomical characters. Biogeographic discussions focus on the effects of climate-induced fragmentation and isolation of mesic communities since the Miocene on species distributions and the role of limestone outcrops as important secondary refugia (next to rainforest) for this biota. A conservation priority for these species is outlined. □ Charopidae, systematics, new species, limestone, rainforest, biogeography, New South Wales. Isabel Hyman (e-mail: ihyman@bio.usyd.edu.au). Australian Museum, 6 College Street, Sydney, 2010; School of Biological Sciences (AOS). University of Sydney, New South Wales 2006; John Stanisic, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia; 1 June 2004. The small to minute Charopidae form a significant part of the land snail fauna in Australia. The family occurs in many parts of Australia (Smith, 1984, 1992; Solem, 1983; 1 984) but is particularly diverse along the eastern edge of the continent (here defined as the area within approximately 200km of the coastline and stretching from the NSW — Vic Border to approximately Cooktown, NE Old). In this region charopids are generally found in the litter zone of rainforests (moist and dry) with comparatively few species occurring in adjacent 220 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 1. Limestone localities. 1, Manilla-Attunga; 2, Macleay Valley; 3, Timor; 4. Glenrock; 5, Borenore; 6, Jenolan; 7, Abercrombie; 8, Church Creek-Colong; 9, Wombeyan; 10, Bungonia; 11, Coolcman; 12, Yarrangobilly; 13; Marble Arch; 14, Wyanbene; 15, Bendethera; 16, Buchan. eucalypt forests (Stanisic, 1990, 1994a). However, within the drier zone of eucalpypt forest and woodland, an archipelago of limestone outcrops forms an important secondary node of diversity for the family. Stanisic (1990) reviewed 50 species (27 new) belonging to 18 genera from subtropical rainforests in eastern Australia and provided a summary of previous Australian and extralimital taxonomic studies on the group. Only minor additions to the Australian species inventory have occurred subsequently (Stanisic, 1993a,b; 1996). One of the major obstacles preventing more comprehensive reviews of the group in the past has been the dearth of material available for study. These snails are very small (mostly <5mm, and often <3mm in diameter), extremely cryptic in their preferred microhabitat and, as experience has shown, usually very difficult to find alive in great numbers. Since 1980, the Queensland Museum (QM) has been particularly focussed on collecting Charopidae in eastern Australia and through a program of leaf litter collecting and sorting (mainly to recover dead shells), it has been possible to gain some insight into the true diversity of the family in this region. The QM collections now contain more than 25,000 specimens of Charopidae from eastern Australia belonging to approximately 440 species of which more than 370 require description. [In contrast Smith (1992) listed 130 described species for the entire Australian continent]. The Australian Museum, Sydney (AMS) has important historical holdings of the family from eastern Australia as well as a significant amount of contemporary material but these collections are limited in their geographic scope. However, in spite of the QM ’s intense collecting efforts (covering more than 2,000 sites) it is highly probable that there are major gaps in the charopid collection base. A significant number of species in the collections are still represented only by single lots and in some cases single specimens from single localities, flagging the possibility that many more species have yet to be discovered. A particularly critical problem for potential reviewers of material currently held in collections is the relative lack of spirit material for anatomical studies. Preserved animal material is available for less than 50% of species so that recollection of many sites is another necessary prerequisite to monographing the group. NEW CHAROPID LAND SNAILS 221 A specialised component of the QM’s wide-ranging collecting effort directed at the Charopidae was an investigation of the land snail fauna of limestone outcrops in eastern NSW. This was conducted mainly from 1992-1995 with casual follow-up collecting in the ensuing years. The main aims of that study were to obtain very detailed data on charopid diversity and distribution both within and between the many limestone outcrops that stretch chiefly along the Great Dividing Range from the Ashford Caves, NE NSW to the NSW-Vic Border (Lismund et al., 1986) (Fig. 1). These limestone formations vary in size from large tower-karst to small, barely discernible outcrops of very low relief, but all act as ‘moisture sinks' in a sea of dry countryside. Many support rainforest in the form of well-developed dry vine thicket that contrasts markedly with the surrounding sclerophyll vegetation. In numerous cases the surrounding landscape has been greatly altered by European settlers for farming activities whereas the outcrops have been left largely intact, sometimes even being fenced to prevent entry of cattle. In some examples however, such as those in the Cooma area, scattered Kurrajong trees are the only reminder of what vegetation may have been present on the outcrop in the past. But, even in these cases, charopids continue to eke out a living among the little microhabitat that remains. Consequently, it has been possible to obtain fairly robust data on both species presence and distribution across a wide range of these specialised habitats in spite of the fact that much of the surrounding countryside has been severely altered. An important contributing factor to the continued survival of these snail communities has been the very nature of the limestone outcrops. Nestled amongst a vegetation community that these days is subject to almost perennial fires, the rock outcrop causes a fire shadow effect which contributes to the survival of both the fire sensitive rainforest vegetation and the associated invertebrate biota. Many of the charopids occurring in these ‘island-like' environments are morphologically, and possibly phylogcnctically distant from those living in the adjacent countryside. Others have sister species in non-limestone habitats. Some are extremely localised ‘limestone' endemics whereas others have broader and more widespread distributions among the archipelago of limestone outcrops. Other species occur on the outcrops but have their main distribution in non-limestone habitats. This study examines a small subset of this diverse and somewhat enigmatic charopid fauna. SCOPE OF STUDY The charopids examined in this study were chosen on the basis of: 1) being well-differentiated on shell characters but covering a range of shell features (sculpture, rib spacing, umbilical width); 2) being well represented in collections so that the shell measurement datasets would be amenable to statistical analysis: and 3) being well represented by preserved material for dissection. Beyond these basic criteria every attempt was made to include all available material that appeared to belong to the various grades of conchological organisation chosen for study (putative clades). Hence, in spite of trying to only deal with large and comprehensive datasets, some species are represented by shells only, sometimes in extremely limited numbers. The geographic limitation of the study (restricted to NSW) also means that some extralimital relatives (Qld in particular) have been excluded. Some chiefly non-limestone species have also been included in this study but only in as far as they contribute to a more comprehensive under- standing of the systematics. Due to the difficulty of assessing the relatively brief and comparatively imprecise descriptions of early authors such as Cox (1868) and Iredale (1933, 1937, 1 94 1 a,b), many of the detailed morphological comparisons in this study are made only with other taxa dealt with either in this study or with those in Stanisic (1990, 1993a,b, 1996). Where comparisons to charopid taxa outside these studies are made much of the data presented may include previously unpublished information and relies heavily on the observ- ations of one of the authors (JS). In attempting identification it should be kept in mind that the subset of species dealt with herein represents a small geographic and phylogenetic component of the total family diversity in eastern Australia and that taxonomic concepts may only have limited application. However, many of the species have restricted distributions and in most cases these are considered to be accurate reflections of the species' true range. Hence, given accurate collection data, the likelihood of mis-identification is considered to be low. 222 MEMOIRS OF THE QUEENSLAND MUSEUM MATERIALS AND METHODS All material used in this study came from the collections of the Australian Museum, Sydney or the Queensland Museum, Brisbane. Species descriptions were generally based on SEM data from 3 shells, measurements from these 3 shells plus 20more empty shells, and anatomical and radular details from at least three spirit specimens. In addition, all the shells of a species were examined for major variation. However, in spite of careful selection ‘wet’ material was not available for all of the species, and for some species very few shells were available for SEM and quantitative analysis. Shells were cleaned in warm dilute bleach, then mounted using silver tape and sputter-coated with gold. Shells were examined and photo- graphed with a Leo 435VP Scanning Electron Microscope. Shells were measured using XTree Gold 2.51 (Johnson et al., 1989-1991). Variables measured were shell height (H) and diameter ( D), spire height (SH), aperture height (AH) and width (AW) and umbilical width (U) (see Solem, 1976 for definitions). Snails were dissected using a Leica MZ8 microscope. Drawings were made using a camera lucida attachment. Radulae were cleaned in warm potassium hydroxide, then mounted on glass slides using carbon tape, then gold-coated and photographed as for the shells. Anatomical and shell terminology largely follows Solem (1984). Shell and anatomical data were scored using DELTA 1.03T (Description Language lor Taxonomy) (Dallwitz et al., 2000). The DELTA files were analysed in PAUP* 4.08b (Phylogenetic Analysis Using Parsimony) (Swofford, 2001) using a heuristic search (100 replicates) with random selective addition and tree-bisection-reconnection. Bootstrap values were calculated using the 4 fast stepwise-addition function in PAUP* with 100 replicates. Trees were viewed in MacClade 3.08a (Maddison & Maddison, 1999). Only taxa for which both anatomical and shell data were available were included in the analysis. Outgroup taxa (Aaadonta constricta constricta and Minidonta hendersoni , Endodontidae) were scored from Solem (1976). Unless otherwise stated classification follows Smith (1992). ACRONYMS AND ABBREVIATIONS USED. General : AMS, Australian Museum, Sydney; QM, Queensland Museum; NSW, New South Wales; Qld, Queensland; Vic, Victoria; SF, State Forest; NP, National Park; NR, Nature Reserve; Ra, Range; R, River; SC, spirit or wet collection; RC, dr\T or reference collection; N, north; S, south; E, east; W, west; NE, northeastern; SE, southeastern. Collectors : DP, Darryl Potter; Gl, Glen Ingram; IL, Ian Loch; JC, Jan Chaseling; JS, John Stanisic; MS, Michael Shea; OG, Owen Griffiths; PHC, Phil Colman; WFP, Winston Ponder. Habitat Data : cnvf, complex notophyll vine forest; drf, dry rainforest; ew, eucalypt woodland; JCt, junction; lm, limestone outcrop; Irf, littoral rainforest; mvf, microphyll vine forest; nvf, notophyll vine forest; rf, rainforest; ro, rocky outcrop; snvf, simple notophyll vine forest; strf, subtropical rainforest; trib.. tributary; vt, vine thicket; wsf, wet sclerophyll forest; wtrf, warm temperate rainforest. Anatomical Data : dg, prostate; e, epiphallus; g, ovotestis; gd, hermaphrodite duct; gg, albumen gland; gt, talon; h, heart; hv, principal pulmonary vein; i, intestine; kdl, primary ureter; kd2, secondary ureter; kp, pericardial lobe of kidney; kr, rectaflobe of kidney; me, mantle collar; mg, mantle gland; p, penis; pp, penial pilaster; ppt, preputial tube; pv; penial verge; prm, penis retractor muscle; ss, spermathecal stalk; ut, uterus; v, vagina; vd, vas deferens. SYSTEMATICS Subclass EUPULMONATA Family CHAROPIDAE Hutton, 1884 Subfamily CHAROPINAE Hutton, 1884 Riiophodon Hedley, 1924 Rhophodon Hedley, 1924: 219; Iredale, 1937: 329; Iredale, 1941b: 2, Kershaw, 1955: 30; Burch, 1976b: 133; Smith & Kershaw, 1979:175; Stanisic, 1990: 114; Smith, 1992: 202 . TYPE SPECIES. Rhophodon peregrinns Hedley, 1 924; by original designation. DIAGNOSIS. Shell very small to minute, discoidal, flammulated to monochrome,with few to many apertural barriers; spire depressed to Hat or very slightly elevated. Protoconch with prominent, slightly curved radial ribs that become more crowded toward the protoconch/teleoconch boundary and very fine, wrinkle-like, discontinuous spiral cordlets. Teleoconch sculpture of very crowded to widely spaced, straight to weakly protractively sinuated radial ribs; secondary sculpture of intersecting microradials and microspirals that form strong beads at their intersection. Umbilicus very wide NEW CHAROPID LAND SNAILS 223 cup-shaped to saucer-shaped. Kidney weakly bilobed with pericardial lobe elongate, triangular, slightly rcflexed at the apex. Penis internally with longitudinal pilasters, occasionaly with additive circular pads. DISTRIBUTION AND HABITAT. From Mt Booroon Booroon, SE Qld, to the Macleay Valley, NE NSW. Species are obligate closed-forest dwellers living among litter and rocks in warm temperate to subtropical (wet to dry) rainforest. REMARKS. Rhophodon Hedley, 1924 was introduced for 3 Australian charopids with apertural barriers. Letomola Iredale, 1941 was subsequently introduced for Rhophodon conforms Hedley, 1924 but the author gave no convincing reasons for the separation from Rhophodon . Stanisic ( 1 990) redefined Rhophodon as a genus of very small to minute charopids with apertural barriers, very wide umbilicus, primarily radial protoconch and prominent radial ribs on the teleoconch. Stanisic (1990) also included three new species and placed the monotypic Egilodonta Iredale, 1937 (type species: Champa bairnsdalensis Gabriel, 1930) into its synonymy but maintained Letomola as a separate genus. A malleate protoconch, the presence of a supraperipheral sulcus and unusual radular features were considered to be sufficient reasons for its separation from Rhophodon. Smith & Kershaw (1979) and Smith (1992) included the Victorian 4 Charopa ’ problematic 'a Gabriel, 1947 in Rhophodon but this species was not examined by Stanisic (1990). Following the examination of additional new material of 4 C\ bairnsdalensis and the discovery of several closely related species in the NSW/Vic Border area, Egilodonta is herein reinstated for a group of southern charopids with dentate apertures. These species are conchologically and anatomically cohesive. In particular the position and less exuberant development of the apertural barriers are features that combine to separate these species from Rhophodon. The position of 4 C\ pmblematica still needs to be determined. Smith & Kershaw (1979) mention the presence of apertural lamellae in the shell of this species, presumably a major reason for its generic placement by those authors, but these lamellae were not mentioned by Gabriel (1947). Hence, until the type specimen of 4 C\ pmblematica is re-examined this species should be regarded as a doubtful member of Rhophodon. Rhophodon Hedley, 1924 is most similar to Egilodonta Iredale, 1937 in shell shape, in the protoconch sculpture (the radial ribs on the protoconch become increasingly crowded at the protoconch/teleoconch margin) and in the possession of apertural barriers. However, Egilodonta differs from Rhophodon in having consistently more widely spaced and thicker teleoconch ribs and consistently fewer apertural barriers that differ in their relative positions on both the palatal and parietal edges of the aperture. In Egilodonta there is only a single parietal barrier (positioned almost half way down the parietal wall), compared with at least two barriers (situated approximately one-half and two thirds down the parietal wall) and occasionally an infraparietal trace in Rhophodon. Rhophodon also usually has at least two palatal barriers (one in R. silvaticus sp. nov.) in contrast to the single palatal barrier in all Egilodonta. Anatomically these two genera show little difference in general features of the pallial and reproductive systems (strong elongation) but this could be largely related to convergent shell shape. Shell design in both groups is character- ised by many tightly coiled whorls and a very much reduced whorl cross-section. Elongation of structures such as the pericardial kidney lobe and epiphallus have may have developed as a spatial adjustment in response to this shared evolutionary trend in fundamental shell shape. Rhophodon and Egilodonta are not only separated by considerable geographic distance (c.600km) but they are also ecologically distinct. Rhophodon is an obligate closed forest group inhabiting warm temperate to subtropical rainforest, albeit sometimes on limestone, whilst Egilodonta appears to be chiefly a calciphile group with a tendency to also occur in eucalypt woodland. Letomola Iredale, 1941 from the Macleay Valley, NSW (which is sympatric with R. kempseyensis) also has apertural barriers. It also has an exsert protoconch and wide umbilicus (similar to Rhophodon) but has a more depressed spire, supraperipheral sulcus and malleate protoconch sometimes with fine, widely spaced, thin, curved, radial ridgelets. The teleoconch sculpture is degenerate consisting of broad, shingle-like overlapping sheets. Letomola also has fewer barriers than Rhophodon. Decor iropa gen. nov. has the same general shell form as Rhophodon (discoidal shell with radial protoconch, wide umbilicus and prominent 224 MEMOIRS OF THE QUEENSLAND MUSEUM radial ribs on the teleoconch) but lacks apertural barriers and has relatively evenly spaced radial ribs on the protoconch. Solem (1983) showed that the shape of microprojectons on the apertural barriers could be a useful means of elucidating phylogenetic relationships within those Pacific Island charopids with apertural barriers and Stanisic (1990) showed that this may also apply to Australian species. Specifically, Letomola was shown to have quite different apertural micro- projections to those of Rhophodon spp., which correlated with fundamental differences in conchological and anatomical features between the two groups (Stanisic, 1990). However, the microsculpture of the apertural barriers was not investigated in the current study. Rhophodon species can be distinguished from each other by a combination of shell size and rib spacing on the teleoconch, but most definitively, on the basis of apertural barrier conformation - number, form and position. Rhophodon kempseyensis Stanisic, 1990 (Figs 2-11; Table 1) Rhophodon kempseyensis Stanisic, 1990: 125, figs 77-79; Smith 1992: 202. TYPE LOCALITY. Natural Arch, Carrai SF, NSW. MATERIAL. All NSW: AMSC168611, C308079, QMMO37076, MO37096, M037151, M056006. M049295, M052724. DIAGNOSIS. Shell very small, chocolate brown with scattered, radially disposed lighter blotches. Teleoconch with extremely crowded, weakly protectively sinuated, narrow radial ribs. Mean ribs/mm 33. Aperture with three parietal barriers (infraparietal present only as a low trace); palatal barriers four, one at the baso-palatal margin. Penis internally with three to four longitudinal pilasters. DISTRIBUTION AND HABITAT. W of Kempsey, from the Yessabah limestone outcrop to The Castles limestone outcrop in the Carrai SF; often found in great numbers in leaf and soil litter, or under logs and rocks, on limestone outcrops covered in rainforest. REMARKS. Rhophopon kempseyensis Stanisic, 1990 has very fine and extremely crowded ribs (mean 34.56 ribs/mm on the last whorl). R. consobrinus (Hedley, 1924) and R. peregnnus (Hedley, 1924) also have crowded ribs, but only about half as many on the last whorl as R. kempseyensis (see Stanisic, 1990). R. duplicostatus sp. nov. has more crowded ribs than kempseyensis (mean 142.3 ribs/mm) but these are very narrow, more like thickened micro- radials, and are grouped in pairs. R. silvaticus sp. nov (mean 38.70 ribs/mm) also has a higher rib count but is smaller with a lower whorl count. R. pcdethorpei sp. nov (mean 1 3.06 ribs/mm) and R. mcgradyorum sp. nov. (mean 8.04 ribs/mm) have considerably lower rib counts. The barrier arrangement in R. kempseyensis is very similar to that of R. palethorpei (three parietal barriers and four palatal barriers) but differs in having the infraparietal barrier present as a low trace rather than a high crescent shaped lamellae. Additional fieldwork since the original description of this species has managed to extend the range of R, kempseyensis further westward (The Castles) in the Macleay Valley but still always in association with limestone covered in rainforest. Rhophodon spp. in general show a strong affinity with rainforest, suggesting that the present day restriction of R. kempseyensis to rainforest-covered limestone may be linked to the possible past restriction of this vegetation type to these rocky refugia. Although rainforest now flourishes elsewhere in the local country- side, R. kempseyensis has yet to disperse into these non-limestone associated habitats. Stanisic ( 1 990) showed that the radula of/?, kempseyensis has enlarged and almost spade-like mesocones on the lateral teeth rather than the more typical lanceolate, possibly an adaptation for scraping food from rock surfaces. This may be evidence that R. kempseyensis has become specialised to living on limestone. Rhophodon palethorpei sp. nov. (Figs 2-1 1,34; Table 1) ETYMOLOGY For Hugh Palethorpe. MATERIAL. HOLOTYPE: QMM070390, Werrikimbe National Park, at Youdales Hut on Kunderang Brook, NSW (3 1°04’45"S, 1 52°1 5’40"E), under fern roots, drflm, 4. i. 1997, JS, JC. Height of s h e 1 1 = 1 .30mm, diameter=2.76mm, H/D=0.47, D/U=2.16, number of whorls=52. ribs on last whorl=89. PARATYPES: All same locality as holotype. QMMO60151. 3SC/73RC, under fem roots, drflm, 4.i.l997, JS, JC; QMM055962, 10SC. on limestone, drf/lm, 7. i. 1995, JS, JC; QMM055961, 4RC, on limestone, drf/lm, 7.i.l995, JS, JC; QMMO59704, 110RC, AMSC205155, 10RC, on limestone, drflm, 7.i. 1 995, JS, JC; QMMO59706, 1 RC, in litter, drflm, 7.U995, JS, JC. DIAGNOSIS. Shell very small, dark golden brown with a wide umbilicus. Teleoconch with slightly sinuated, moderately crowded, relatively wide ribs. Mean ribs/mm 13.06. Aperture with NEW CHAROPID LAND SNAILS 225 three parietal barriers; four palatal barriers, one at the baso-palatal margin. Penis internally with one round basal pilaster. DESCRIPTION. Shell very small, brown to golden brown, with 4.3-5. 1 (mean 4.65) tightly coiled whorls, the last descending more rapidly. Shell diameter 2.4 1-3. 10mm (mean 2.67mm). Apex and early spire flat. Height of shell 1.05- 1.40mm (mean 1.24mm), H/D 0.42-0.50 (mean 0.46). Protoconch exsert of 1.4- 1.7 whorls. Apical sculpture of slightly curved, moderately spaced radial ribs, becoming more crowded toward the protoconch/teleoconch border, width of interstices (in multiples of rib width) 7 at apex, 3 at post-nuclear boundary; secondary sculpture of moderately spaced, weak spiral wrinkles. Teleoconch sculpture of numerous slightly protractively sinuated radial ribs, 68-152 (mean 1 1 0) on last whorl. Ribs/mm 8.76-13.27 (mean 13.06). Ribs wide, height equal to width; straight in section, rounded on top. Rib inter- stices on first post-nuclear whorl equal to width of three major ribs; interstices on penultimate whorl equal to width of 2.5 major ribs. Interstitial sculpture of microradial riblets and fine microspiral cords. Microradial riblets low, 4-6 between major ribs on first post-nuclear whorl, 7-8 on penultimate whorls; microspiral cords low, crossing major radials, forming rounded beads at their intersection with microradials. Umbilicus very wide, saucer-shaped, diameter 1.00- 1.28mm (mean 1.14mm), D/U 2.16-2.57 (mean 2.36). Sutures im- pressed, whorls evenly rounded above and below periphery. Aperture ovate; parietal callus present. Three parietal barriers and four palatal barriers present, all crescent-shaped lamellae. Based on 23 measured adults (QMM070390, MO60151). Ovotestis containing two clumps of alveoli; with one alveolar lobe per clump. Talon stalk 1 -3 times diameter of talon; talon circular. Penis retractor muscle half the length of penis or greater; inserting at junction of penis and epiphallus. Epiphallus longer than penis; wider than vas deferens; pilasters longituduinal; retractor muscle not entwined with epiphallus. Penis sheath present; penis tubular (with a large bulge on one side at base); internally with one round pilaster basally. Vagina shorter than penis. Pallial cavity with unilobed kidney. 226 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 3. Dorsal view of shells. A, Rhophodon kempseyensis, AMSC 153720. B, Rhophodon palethorpei QMM070390. C, Rhophodon mcgradyonnn , QMMO70392. D, Rhophodon silvaticus, QMMO70391 E Rhophodon duplicostatus , QMMO70393. F, Egilodonia w yanbenensis, QMMO70394. G Egilodonia bende'hera, QMMO70395. H, Egilodonia paucidentata, QMM049447. 1 , Egilodonia paucidentata QMMO70396. J, Egilodonia baimsdalensis (Yarrangobilly Caves, Kosciusko National Park) AMSC 142959’ K. Egilodonia baimsdalensis (Lakes Entrance), AMSC 153706. L, Egilodonia baimsdalensis (Alpine Wav’ Kosciusko National Park), AMSC 154808. Scale bars=0.5mm in C, D, E; 0.8mm in F-L- 1mm in A B NEW CHAROPID LAND SNAILS 227 Radula with central tooth strongly tricuspid, considerably smaller than laterals; lateral teeth tricuspid; mesocone of first lateral tooth slender, slightly diamond-shaped; mesocone of marginal teeth rounded, spade-like; ectoconc of outer marginal teeth split into three teeth; endocone occasionally split into two or more teeth. Number of lateral teeth: 7; marginal teeth: 7; radular rows: 123. Based on 3 dissected specimens (QMM055962). DISTRIBUTION AND HABITAT. Known only from the type locality; living on limestone outcrops covered in dry rainforest (vine thicket). REMARKS. Rhophodon palethorpei sp. nov. appears to be very closely related to R. kempseyensis. The barrier configuration of the two species is basically the same except that in R. palethorpei the infraparietal barrier is much more developed. R. palethorpei is also similar in to R. kempseyensis in most aspects of shell measure- ment, but differs most dramatically in having a lower rib count (13.06 ribs/mm compared with 34.56 ribs/mm in R. kempseyensis). R. palethorpei is readily distinguished from R. silvaticus sp. nov. by its larger size, lower rib count and in having a greater number of palatal barriers (four as opposed to one). The many similarities between the shells of R. kempseyensis and R. palethorpei , in particular the configuration of the apertural barriers, suggest common ancestry. The two species are now geographically isolated in two different drainage systems: R. kempseyensis in the Macleay River valley and R. palethorpei in the Kundcrang Brook watershed. This may be related to past restriction of rainforest habitats to refugia such as rocky outcrops and the consequent long-term isolation of once conjunct populations. Although R. palethorpei is currently known only from one locality, there is a possibility that it also exists on other, yet to be sampled, limestone outcrops along Kunderang Brook. Rhophodon silvaticus sp. nov. (Figs 2-7; Table 1) ETYMOLOGY. Latin silvaticus, pertaining to forests; referring to the forest habitat. MATERIAL. HOLOTYPE: QMMO70391, Thumb Ck SF, slopes of Blue Knob, c. 11.5km NW Burrapine, W Macks vi lie, NSW(30°4r45"S, 152°33T5"E), under logs, wtrf, 29.ix.1993, JS, JC. Height of shell=0.67mm, diameter=1.39mm, H/IX).48, D/U=ratio 2.36, number of whorls=4.1, ribs on last whorl=169. PARATYPES: QMM049318, 2RC, same data as holotype; QMM0373 1 8, 4SC, same locality data as holotype, under logs on rocky hillside, wtrf. 15.i.l992, JS, JC. DIAGNOSIS. Minute, golden brown shell with wide umbilicus. Teleoconch with narrow, slightly sinuated, very closely spaced ribs. Mean ribs/mm 38.70. Aperture with two parietal barriers and one palatal barrier. Anatomy unknown. DESCRIPTION. Shell minute golden brown, with c.4.1 tightly coiled whorls, the last descending more rapidly; shell diameter c. 1 .39mm. Apex and early spire fiat. Height of shell c.0.67mm, H/D c.0.48. Protoconch exsert of c. 1 .6 whorls. Apical sculpture of slightly curved, moderately spaced radial ribs, becoming more crowded toward the protoconch/telcoconch border, width of interstices (in multiples of rib width) 4 at apex, 3 at post-nuclear boundary; secondary sculpture of moderately spaced, weak spiral wrinkles. Teleoconch sculpture of numerous weakly protectively sinuated radial ribs, c. 169 on last whorl. Ribs/mm c.38.70. Ribs narrow, height less than width; straight in section, rounded on top. Rib interstices on first post-nuclear whorl equal to width of 2 major ribs; interstices on penultimate whorl equal to width of 1.5 major ribs. Interstitial sculpture of fine microradial riblets and microspiral cords. Microradial riblets low, 2 between ribs on first post-nuclear whorl; 3 between ribs on penultimate whorl; microspiral cords low, crossing major radials, forming elongated beads at their intersection with the microradials. Umbilicus very wide saucer-shaped, diameter c. 0.59mm, D/U c.2.36. Sutures impressed, whorls evenly rounded above and below periphery. Aperture ovately lunate; parietal callus present. Two parietal barriers and one palatal barriers present, all crescent shaped lamellae. Based on the measured holotype (QMMO70391). Anatomy unknown. DISTRIBUTION AND HABITAT. Known from the type locality; living under logs among volcanic rocks in temperate rainforest. REMARKS. Rhophodon silvaticus sp. nov. is most similar to R. kempseyensis in rib spacing and general satin-like appearance of the shell surface, however it is much smaller, has the teleoconch ribs slightly more widely spaced (ribs/mm =38.70) and possesses only one palatal barrier (four in R. kempseyensis). R. palethorpei has the teleoconch ribs more widely spaced than those of R. silvaticus and like R. kempseyensis 228 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Protoconch of shells from above. A, Rhophodon kemosevemii AMSriS'mn r palethorpei, QMM070390. C. Rhophodon tncgradyorum^QMMOlOi 92^D ^ Rhophodon ^sdvaticus QMMO70391. E Rhophodon duplicostatus, QMMO70393. F, Egilodonia wyanbenensis QMMO70394 G EgUodonta bendethera QMMO70395. H. Egilodonta paucidentata, b MM 04944 7^ I '^EsilodoiUa paucidentata , QMMO70396. J, Egilodonta bairnsdalensis , AMSC142959 K Eeilodonta hairncdnlen • AMSC1 53706. L, Egilodonta bairnsdalensis , AMSC154808. Scale bars=0. 1mm ^ batrnsdalensts , NEW CHAROPID LAND SNAILS 229 also has four palatal barriers. This is the smallest of the Rhophodon group of species, and additional material is needed to adequately define the morphometries of the shell and to detail the anatomy. R. silvaticus is currently only known from the one locality in the Thumb Creek SF and more fieldwork is needed to confirm its range. There would appear to be no obvious reason why this species is not more widespread in the upper Taylors Ann watershed (Nulla Five Day SF) where similar habitats exist. The microhabitat data (on rocky hillside) suggests that it may have a patchy localised distribution confined to areas with rocky substrate. Rhophodon megradyorum sp. nov. (Figs 2-7, 33; Table 1) ETYMOLOGY. For Jim and Audrey McGrady. MATERIAL. HOLOTYPE: QMMO70392, Bcllbird Gully, Gibraltar Ra., NSW, 4.U988, G Annabell. Height of shell=0.99mm, diameter=l .67mm, H/D=0.59, D/U=2.61, number of whorls=4.7, ribs on last whorl=47. PARATYPES: QMMO66108, 3RC, same data as holotype. DIAGNOSIS. Shell minute, golden brown, with a wide umbilicus. Teleoconch with wide, straight (not sinuated) widely spaced ribs. Mean ribs/mm 8.04. Aperture with 3 parietal and 3 palatal barriers and a baso-columellar barrier; barriers thickened at the apex. Anatomy unknown. DESCRIPTION. Shell very small, golden brown to yellow, with 4. 5-4. 9 (mean 4.7) tightly coiled whorls, the last descending more rapidly in front. Shell diameter 1.64- 1.69mm (mean 1.67mm). Apex and early spire slightly concave to flat. Height of shell 0.82-0. 99mm (mean 0.88mm), H/D 0.50-0.59 (mean 0.53). Protoconch exsert of 1.7- 1.8 whorls. Apical sculpture of curved, moderately spaced, radial ribs, becoming more crowded toward protoconch/teleoconch boundary; width of interstices (in multiples of rib width) 6 at apex, 3 at post-nuclear boundary; secondary sculpture of weak, discontinuous spiral cords. Teleoconch sculpture of numerous, protractively sinuated widely spaced radial ribs; 35-47 (mean 42) ribs on last whorl. Ribs/mm 6.79-8.96 (mean 8.04). Ribs relatively broad, height less than width; straight in section, rounded on top. Rib interstices on first post-nuclear whorl equal to width of 4-6 ribs; interstices on penultimate whorl equal to width of 7 ribs. Interstitial sculpture of microradioal riblets and fine microsopiral cords. Microradial riblets low, 12-13 between ribs on first post-nuclear whorl and 20 between ribs on penultimate whorl; microspiral cords low, crossing major radials, forming strong, round to square beads at their intersection with microradial riblets. Umbilicus widely open, saucer-shaped, diameter 0.61 -0.64mm (mean 0.63mm). D/U 2.61-2.67 (mean 2.68). Sutures impressed, whorls evenly rounded above and below periphery. Aperture ovately lunate, parietal callus present. Three parietal barriers, one baso-columellar barrier and four palatal barriers present; barriers thickened. Based on 4 measured adults (QMMO70392, MO66108). Anatomy unknown. DISTRIBUTION AND HABITAT. Known only from the type locality in the Gibraltar Range NP, east of Glen Innes, NSW; habitat and microhabitat unknown. REMARKS. The teleoconch ribs of/?, megradyorum sp. nov. are relatively widely spaced compared with other NE NSW Rhophodon taxa and the shell bears a greater overall similarity to /?. minutissimus Stanisic, 1990 from SE Qld. It is, however, distinguished from that species by having only two parietal barriers (three in R. minutissimus) and four palatal barriers (three in R. minutissimus). The barriers of R. megradyorum are also comparatively more thickened and lack the blade-like appearance of those in other found in other NE NSW Rhophodon . In this respect the barriers resemble those of /?. colmani Stanisic, 1990 from the Kenil worth SF, SE Qld. However, the latter species differs dramatically from /?. megradyorum in having many more apertural barriers (six palatal) and teleoconch ribs that are broader and much more widely spaced (mean ribs/mm 5.81). Rhophodon duplicostatus sp. nov. (Figs 2-7, 34; Table 1) ETYMOLOGY. Latin duplico , double and costatus , ribbed; referring to the major rib doublets on the teleoconch. MATERIAL. HOLOTYPE: QMMO70393, Glenugie SF, slopes of Glenugie Peak (= Mt Elaine), SE Grafton, NSW (29°50 , 0rS, 153°04’47"E), under rocks on rocky hillside, remnant warm temperate rainforest, 1 6.i. 1 992, JS. JC. Height of shell=0.78mm, diameter- 1 .7mm, H/D =0.46, D/U =2.54, number of whorls=4.7, rib pairs on body 3 1 3. PARATYPES: QMM037344, 7RC, same data as holotype. DIAGNOSIS. Shell very small, golden brown, with a wide umbilicus. Teleoconch ribs slightly 230 MEMOIRS OF THE QUEENSLAND MUSEUM Lf'eral view of shells. A. Rhophodon kempseyensis , AMSC 153720. B, Rhophodon palethorpei , QMM070390 C Rhophodon mcgradyorum , QMMO70392. D, Rhophodon silvaticus, QMMO70391 E Rhophodon duplicostatus, QMMO70393. F, Egilodonta wyanbenensis, QMMO70394. G, Egilodonta Egilodonta paucidentata, QMM049447. 1 , Egilodonta paucidentata , QMMO70396. J, Egilodonta bairnsdalensis, AMSC \ 42959. K. Egilodonta bairnsdalensis, AMSC 153706. L, tgi/oaonta bairnsdalensis , AMSC 154808. Scale bars=lmm in A, B; 0.5mm in C-L. NEW CHAROPID LAND SNAILS 231 TABLE 1 . Shell measurements for species of Rhophodon. (AH/AW, ratio of aperture height to aperture width; D, shell diameter; D/U, ratio of shell diameter to umbilicus width; H, shell height; H/D, ratio of shell height to diameter; N, number of specimens; NW, total number of whorls; PW, number of protoconch whorls; RIB, number of ribs on last whorl; RIBS/M, number of ribs per mm on the last whorl; SP, spire height; TW, number of telcoconch whorls; UMB, umbilicus width).* indicate actual number of measurements taken. Measurement Rhophodon kempseyensis Rhophodon palethorpei Rhophodon silvaticus Rhophodon duplicostatus Rhophodon megradvorum N 23 (*22, **13) 23 1 3 (*1) 4 D 2.45-2.83 (2.628) ±0.111 2.41-3.01 (2.674) ± 0.141 1.39 1.49-1.7(1.627) ± 0.118) 1.64-1.69(1.672) ± 0.024 PW 1.3-1.6(1.514) ± 0.064 * 1.5-1.7(1.543) ± 0.059 1.6 1.5 * 1.7-1.8(1.725) ± 0.05 TW 3. 3-3. 8 (3.582) ± 0.113 * 3.2-4. 1 (3.704) ± 0.203 2.5 3.2* 2.8-3.2(2.95) ± 0.173 NW 4.8-5.3 (5.095) ± 0.143 * 4.9-5. 6 (5.248) ± 0.175 4.1 4.7* 4. 5-4.9 (4.675) ± 0.171 H 1.03-1.24(1.118) ± 0.059 1.05-1.4(1.239) ± 0.095 0.67 0.72-0.83 (0.777) ± 0.055 0.82-0.99 (0.88) ± 0.075 SP 0 0-0.16(0.083) ± 0.004 0.01 0-0.1 (0.003) ± 0.006 0-0.04 (0.018) ± 0.021 AH/AW 1 .5-1.4 (1.4) ± 0.08 1.06-1.7(1.32) ± 0.14 1.54 1.29-1.57(1.42) ± 0.14 1.21-1.41 (1.3) ± 0.11 UMB 1-1.31 (1.169) ± 0.08 1-1.28(1.136) ± 0.077 0.59 0.58-0.67 (0.633) ± 0.047 0.61-0.64(0.625) ± 0.017 RIB 246-358 (288) ± 32.738 ** 68-152(110) ± 20.967 169 - 35-47 (42.25) ± 5.123 RIBS/MM 28.17-40.27 (34.56) ± 3.678 ** 8.763-13.27(13.06) ± 2.084 38.7 - 6.79-8.96 (8.035) ± 0.906 H/D 0.38-0.47 (0.43) ± 0.03 0.42-0.5 (0.46) ± 0.02 0.48 0.46-0.49 (0.48) ± 0.02 0.5-0.59 (0.53) ± 0.04 D/U 2.08-2.46 (2.25) ± 0.1 2.16-2.57(2.36) ± 0.11 2.36 2.54-2.6 (2.57) ± 0.03 2.61-2.77 (2.68) ± 0.07 sinuated, very narrow and extremely crowded, complex, each consisting of a pair of ribs. Ribs/mm c. 142.3. Aperture with three parietal and six blade-like palatal barriers present. Anatomy unknown. DESCRIPTION. Shell minute, golden brown, with c.4.7 tightly coiled whorls, the last descending more rapidly in front. Shell diameter 1.49-1. 70mm (mean 1.63mm). Apex and early spire slightly concave. Height of shell 0.72-0. 83mm (mean 0.88mm), H/D 0.46-0.49 (mean 0.48). Protoconch exsert of c.1.5 whorls. Apical sculpture of moderately spaced, slightly curved radial ribs, becoming more crowded at protoconch/teleoconch boundary; width of interstices (in multiples of rib width) 5 at apex, 2.5 at post-nuclear boundary; secondary sculpture of weak, discontinuous spiral wrinkles. Teleoconch sculpture of very numerous and extremely crowded weakly protractively sinuated radial ribs that usually occur as doublets. Rib pairs on last whorl 3 1 3, ribs pairs/mm 142.3, giving the shell an appearance of being smooth. Height of ribs less than width; ribs straight in section, rounded on top. Rib interstices on the first post-nuclear whorl about the size of the rib doublet (i.e. the size of two ribs). Interstitial sculpture of fine microradial riblets and thin microspiral cords. Microradial riblets low, 3-4 between ribs on first post-nuclear whorl and 2 between ribs on penultimate whorl; microspiral cords forming prominent elongate spiral beads at their intersection with the microradials. Umbilicus wide saucer-shaped, diameter 0.58-0. 67mm (mean 0.63mm), D/U 2.54-2.60 (mean 2.57). Sutures impressed, whorls evenly rounded above and below the periphery. Aperture ovately lunate, parietal callus present. Three slender parietal barriers present; six very fine palatal barriers present, three in the baso-columellar region; barriers blade-like. Based on 3 measured adults (QMMO70393, M037334). Anatomy unknown. DISTRIBUTION AND HABITAT. Rhophodon duplicostatus sp. nov. is known only from the type locality in Glenugie SF, NE NSW; found in litter among volcanic talus in a small patch of remnant warm temperate rainforest. 232 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 6 Aperture of shells, showing apcrtural barriers. A, Rhophodon kempseyensis, AM SC 153720 B Rh °Pl‘ odon QMM070390. C Rhophodon mcgradyorum , QMMO70392. D, Rhophodon nmSw r? ' B, Rhophodon dupljcos talus, QMMO70393. F, Egilodonla n yanbenensis, QMMO70394. G, Egilodonla bendelhera, QMMO70395. 11, Egilodonla pa, .edemata, QMM049447. I, Egilodonla paucidentata, QMMO70396. J, Egilodonla bairnsdalensis, AMSC14' ) 959 K Eeilodonla bairnsdalensis, AMSC 153706. L, Egilodonla bairnsdalensis, AMSC 154808. Scale bars=0.2mm. ‘ NEW CHAROPID LAND SNAILS 233 REMARKS. Rhophodon duplicostatus sp. nov. has very fine and extremely crowded radial ribs that are almost too dense to count using light microsopy. Under SEM the ribs appear as doublets with each doublet separated by an interstice sculptured with fine, low microradials. In this respect the species cannot be confused with any of its congeners. The apertural dentition of R. duplicostatus is also diagnostic in having a proliferation of lamellae in the baso-columellar region. On the basis of having three parietal teeth the species is most readily associated with the R. kemps evens is- R. palethorpei-R. peregrinus group of species. The isolation of R. duplicostatus , a member of a closed forest group of land snails, on the unusual Glenugie Peak, suggests derivation from an ancestral population inhabiting once more widespread closed forests in NE NSW. Glenugie Peak (= Mt Elaine on early maps) is a pile of volcanic talus supporting a small patch of rainforest amidst an otherwise eucalypt dominated countryside. As a shift to more xeric climates restricted rainforest in the Tertiary (Kemp, 1981; Webb & Tracey, 1981; Adam 1 992), mesic communities in the tow coastal area around Grafton would probably have been cornered on such local topographic refiigia. The refugial status of Glenugie Peak is reinforced by the fact that it is also home to 1 5 other species of land snails that include several possible endemic species (Stanisic, unpubl. data). This level of overall diversity is high in the context of the local area where site diversity is generally less than 1 0 species (Stanisic, 1994). According to Floyd (1987), Glenugie Peak is also floristically unusual, thus further emphasising the complex evolutionary history of this refugium. Geological data suggests that other similar refugia (yet to be investigated for land snails) exist closer to the coast in the vicinity of Glenugie SF (e.g. Brown's Knob) and it would not be surprising to find that R. duplicostatus has a broader local distribution among these. Egilodonta Iredale, 1937 Egilodonta Iredale, 1937: 328; Gabriel, 1947: 120; Gabriel & Macpherson, 1947:162; Kershaw, 1956a: 142; Burch, 1976b: 133; Smith & Kershaw, 1979: 174; Stanisic, 1990: 1 14 (in synonymy); Smith, 1992: 202 (in synonymy). TYPE SPECIES. Champa baimsdalensis Gabriel, 1930; by original designation. DIAGNOSIS. Shell minute, discoidal, monochrome golden brown, with relatively few apertural barriers; umbilicus very wide saucer- shaped. Protoconch with prominent, slightly curved radial ribs that become more crowded toward the protoconch/teleoconch boundary and very fine, wrinkle-like, discontinuous spiral cords. Teleoconch sculpture consisting of bold, widely spaced radial ribs whose interstices are sculptured with fine microradial riblets; secondary' sculpture of low microradial riblets and microspiral cords that form strong beads at their intersection. Kidney weakly bilobed with pericardial lobe long, triangular. Epiphallus thick and muscular and well differentiated from the vas deferens. Penis internally sometimes with vergic structures. DISTRIBUTION AND HABITAT. Egilodonta ranges from the Baimsdale region, SE Vic to just south of Braidwood, SE NSW. Species live in the litter and appear associated with limestone habitats, although E. baimsdalensis (Gabriel, 1930) also shows some preference for surrounding eucalypt forest. REMARKS. On the basis of limited material, Stanisic ( 1 990) synonymised Egilodonta Iredale, 1937 with Rhophodon Hedley, 1924. The discovery of additional species of Charopidae with dentate apertures from the NSW- Vic Border area has enabled a reassessment of the status of Egilodonta. This group of charopids displays a number of consistent differences from the NE NSW species grouped in Rhophodon. Bolder and more widely spaced ribs in combination with fewer apertural barriers serve to circumscribe the SE NSW/NE Vic species. Consequently Egilodonta is restored to full generic status to include the type and three new species from this region. Inexplicably Solem (1972) placed ‘C\ bairndalensis in Dentherona Iredale, 1933 (type species: Helix dispar Brazier, 1871) without formally documenting the implied generic synonymy. Distinguishing characters that separate Egilodonta Iredale, 1937 and Rhophodon Hedley, 1924 have been discussed under the latter genus. Conchological differences among Egilodonta spp. arc less dramatic than those among Rhophodon spp., which may indicate a relative recent separation of Egilodonta populations. Apertural dentition, rib spacing, and shell size vary marginally; however, genitalia show considerable variation compared with Rhophodon spp. These differences in penial chamber structure (verge and pilasters) most likely relate to species level interactions due to relatively recent microsympatry (see Solem, 1983). Significantly the one species that is truly 234 MEMOIRS OF THE QUEENSLAND MUSEUM F ' G K . : View of shells. A, Rhophodon kempseyensis , AMSC153720. B, Rhophodon palethorpei, QMM070390 C Rhophodon mcgracfyonim QMMO70392. D. Rhophodon silvaticus . QMMO70391. E, Rhophodon duphcos tutus QMMO70393 . F. Egilodonta wyanbenensis, QMMO70394. G, Egilodonta nutmlimff 0 , 7 95 '/-' Egilodonta paucidentata, QMM049447. I, Egilodonta paucidentata , QMMO70396. j, Egilodonta bairmdalemts ,AMSC 142959. K, Egilodonta bairnsdaknsis , AMSC 1 53706. L, Egilodonta batrnsdalensis. AMSC 154808. Scale bars=lmm in A, B; 0.5mm in C-L. NEW CHAROPID LAND SNAILS 235 allopatric, E. bairnsdalensis (Gabriel, 1930), lacks a penial verge. It is probable that Egilodonta is still in a stage of comparatively exuberant speciation (in an historical context) compared with the environmentally restricted Rhophodon. The most widely collected species is E. bairnsdalensis which appears to be a eucalypt woodland species that also takes advantage of limestone habitats. E. wyanbenensis sp. nov., E. bendethera sp. nov. and E. paucidentata sp. nov. appear to be specifically associated with lime- stone environments based on current evidence, but more fieldwork in surrounding habitat needs to be completed before this association is firmly established. Egilodonta bairnsdalensis (Gabriel, 1930) (Figs 2-7, 10-12, 34; Table 2) Charopa bairnsdalensis Gabriel. 1930: 78, pi. 2, figs 11-12. Endodonta bairnsdalensis (Gabriel). Kershaw, 1956a: 137. Egilodonta bairnsdalensis (Gabriel). Iredale, 1937: 329; Gabriel, 1947: 120; Gabriel & Macphcrson, 1947:162; Smith & Kershaw, 1979: 174. Dentherona bairnsdalensis (Gabriel). Solem, 1972: 85, figs 17, 18. Rhophodon bairnsdalensis (Gabriel). Stanisic, 1990: 138, figs 89-91; Smith, 1992:202. TYPE LOCALITY. Baimsdale, Vic. MATERIAL. NSW: AMSC355056, C355058, QMM065004, MO65017. DIAGNOSIS. Shell minute, straw-coloured, with a wide saucer-shaped umbilicus. Teleoconch with wide, almost straight, relatively broad, very widely spaced ribs. Mean ribs/mm 5.32. Aperture with one strong parietal barrier, one basal barrier and one palatal barrier. Penis internally with 2-4 longitudinal pilasters; verge absent. DISTRIBUTION AND HABITAT. The Great Dividing Range from the Baimsdale area in SE Vic north to the Yarrangobilly Caves, Kosciusko NP, SE NSW; found living among litter in eucalypt woodland. Specimens cited in Stanisic (1990) from Marble Arch are now included in E. paucidentata sp. nov. REMARKS. Egilodonta bairnsdalensis (Gabriel, 1930) can be distinguished from E. bendethera sp. nov. and E. paucidentata sp. nov. by its stronger apertural barriers, slightly wider ribs, the presence of two to four penial pilasters and the absence of a penial verge. E. wyanben- ensis sp. nov. is readily distinguished by having more crowded ribs on the last whorl. Some individuals of E. bairnsdalensis (mostly from around Lakes Entrance, Vic) have no palatal barrier, while others (mostly from around Kosciusko NP) have both basal and palatal barriers with the palatal weakly developed. However, in the absence of other shell (including shell size and shape) or anatomical differences between these populations, they are all included in E. bairnsdalensis. Egilodonta wyanbenensis sp. nov. (Figs 2-11; Table 2) ETYMOLOGY. From the type locality. MATERIAL. HOLOTYPE: QMMO70394, Wyanbene Caves, S Braidwood, NSW (35°48 , 25"S, 149°4r20 M E), lm, in litter, 6.U990, JS, JC. Height of shell=0.96mm, diameter= 1.99mm, H/D=0.48, D/U=2.16, number of whorls=4.6, ribs on last whorl=66. PARATYPES: QMM037793, 44RC, AMSC205156. 5RC, same data as holotvpe; QMMO29270, 34SC73RC, Wyanbene Caves, S Braidwood. (35°48 , 25' , S, 149°4r20 M E), among litter on lm, 611990, JS, JC. DIAGNOSIS. Minute straw-coloured shell with wide umbilicus. Teleoconch with moderately broad, slightly sinuated, moderately crowded ribs. Mean ribs/mm 10.32. Aperture with one parietal barrier, one basal barrier and one palatal barrier; barriers very weakly developed. Penis internally with a short verge and no pilasters. DESCRIPTION. Shell very small, straw coloured, with 4.3-5. 1 (mean 4.65) tightly coiled whorls, the last descending more radpidly in front. Shell diameter 1.78-2. 39mm (mean 2.14mm). Apex and early spire flat to slightly concave. Height of shell 0.9 1-1. 15mm (mean 1.03mm), H/D 0.43-0.52 (mean 0.48). Protoconch exsert of 1.4- 1.7 whorls. Apical sculpture of curved radial ribs, becoming more crowded at protoconch/teleoconch boundary; width of interstices (in multiples of rib width) 6 at apex, 3 at post-nuclear boundary; secondary sculpture of discontinuous, very narrow, spiral wrinkles. Teleoconch scuplpture of modcraretly crowded, slightly protectively sinuated radial ribs; 56-85 (mean 69) ribs on last whorl. Ribs/mm 8.49-13.05 (mean 10.32). Ribs moderately broad, height less than width; straight in section, rounded on top. Rib interstices on first post-nuclear whorl equal to width of 2.5 ribs; interstices on penultimate whorl equal to width of 3 ribs. Interstitial sculpture of fine radial riblets and crowded microspiral cords, about equal in strength. Microradials low, 3-6 between ribs on first post-nuclear whorl, 8 between ribs on penultimate whorl; microspiral cords crossing 236 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 8. Radula. A-C, Rhophodon kempseyensis, QMMO37096. D-F, Rhophodon palelhorpei, QMM055962 G-I, Egilodonia wyanbenensis , QMMO29270. J-L. Egilodonta bendethera , QMMO70395 Scale bars = 0.01mm. major ribs and forming strong elongate beads at their intersection with microradials. Umbilicus very wide saucer-shaped, diameter 0.80- 1 . 1 4mm (mean 0.48mm), D/U2.08-2.31 (mean 2.18). Sutures impressed, whorls evenly rounded above and below the periphery. Aperture ovately lunate, parietal callus present. One slender parietal barrier, a weak basal barrier and one palatal barrier present; all cresent shaped lamellae. Based on 16 measured adults (QMMO70394, M037793, M037796). Genitalia with ovotestis containing two clumps of alveoli; with more than two alveolar lobes per clump. Talon stalk more than 3 times diameter of talon; talon circular. Penial retractor muscle entwined with epiphallus, inserting at junction of the penis and epiphallus. Epiphallus longer than penis, entering penis through a verge; length of NEW CHAROPID LAND SNAILS 237 verge less than half the length of penis. Epiphallus muscular, reflexed, wider than the vas deferens, internally with longitudinal pilasters. Penis tubular without internal pilasters; sheath present. Vagina tubular, longer than penis. Pallial cavity with almost unilobed kidney; pericardial lobe elongate, triangular with apex slightly reflexed. Radula with central tooth strongly tricuspid, considerably smaller than laterals; lateral teeth tricuspid; mesocone of first lateral tooth slender, slightly diamond-shaped; marginal teeth rounded; ectocone of outer marginal teeth split into four or more teeth; endocone occasionally split into two or more teeth. Number of lateral teeth 5; marginal teeth 6; radular rows 1 10. Based on 3 dissected specimens (QMMO29270). DISTRIBUTION AND HABITAT. Known only from the type locality; in litter on weakly vegetated limestone among open eucalypt woodland. REMARKS. Egilodonta uyanbenensis sp. nov. is readily distinguished from E. baimsdalensis , E. bendethera sp. nov. and E. paucidentata sp. nov. by the more crowded ribs on the teleoconch. Anatomically, E. wyanbenensis can be dis- tinguished from its congeners by a combination of the presence of a short penial verge and absence of any penis pilasters. E. baimsdalensis has two to four longitudinal pilasters and lacks a verge; E. bendethera has a long penial verge and a single longitudinal pilaster; and E. paucidentata has a short penial verge and several longitudinal pilasters. The Wyanbene Caves locality has been relatively poorly surveyed, and the presence of£. paucidentata sp. nov. in the geographically proximate Bendethera and Marble Arch limestone outcrops might suggest that further fieldwork at this site could yet add this species to the inventory. Egilodonta bendethera sp. nov. (Figs 2-11; Table 2) ETYMOLOGY. For the type locality. MATERIAL. HOLOTYPE: QMMO70395, Moruya, c.60km W at Bendethera Cave, NSW (35°55'54"S, 149°42T2 H E), lm/Ficus, Acacia and eucalypt scrub, in litter on limestone rocks, 6.i.l994, JS, JC. Height of shell=0.91mm, diameter= 1.64mm, H/D^O.55, D/U=2.34, number of whor!s=4.3, ribs on last whorl=27. PARATYPES: QMM068759, I4RC/14SC, same data as holotype; QMMO68760, 160RC, AMSC205157, 10RC, same data as holotype. DIAGNOSIS. Shell very small, light golden brown with a wide umbilicus. Teleoconch with broad, widely spaced, protractively sinuated ribs. Mean ribs/mm 4.67. Aperture with one parietal barrier, and one weakly developed basal barrier and one weakly developed palatal barriers. Penis internally with long penial verge and one longitudinal pilaster. DESCRIPTION. Shell very small, light golden brown, with 4.3-5. 8 (mean 5.3) tightly coiled whorls, last descending more rapidly in front. Shell diameter 1.64-2. 63mm (mean 2.34mm). Apex and early spire flat. Height of shell 0.9 1-1. 38mm (mean 1.78mm), H/D 0.48-0.55 (mean 0.50). Protoconch exsert of 1.5- 1.7 whorls. Apical sculpture of slightly curved, moderately spaced radial ribs, becoming more crowded toward protoconch/teleoconch boundary; width of interstices (in multiples of rib width) 7 at apex, 3 at post-nuclear boundary; secondary sculpture of moderately spaced, discontinuous radial wrinkles. Teleoconch sculpture of bold, widely spaced, almost straight radial ribs, 27-47 (mean 34) on last whorl. Ribs/mm 3.92-6.08 (mean 4.67). Rib height greater than width; straight in section, rounded on top. Rib interstices on first post-nuclear whorl equal to width of 5-6 ribs; interstices on penultimate whorl equal to width of 6 ribs. Interstitial sculpture of fine radial riblets and crowded microspiral cords, about equal in strength. Microradials low, 12 between ribs on first post-nuclear whorl, 25 between ribs on penultimate whorl; microspiral cords crossing major ribs and forming strong round to elongate beads at their intersection with microradials. Umbilicus very wide saucer-shaped, diameter 0.7- 1.5mm (mean 0.96mm), D/U 2.19-2.63 (mean 2.44). Sutures impressed, whorls rounded above and below periphery. Aperture ovately lunate; lip sinuous; parietal callus present. One very fine parietal barrier, one basal and one palatal barrier present; barriers low blade-like. Based on 10 measured adults (QMMO70395, M068759). Genitalia with ovotestis containing two clumps of alveoli; more than two alveolar lobes per clump. Talon stalk more than 3 times diameter of talon; talon circular. Penial retractor muscle inserting at the junction of the penis and epiphallus. Epiphallus longer than penis; wider than vas deferens, muscular, reflexed, entering penis through a verge; internally with longitud- inal pilasters. Penis tubular, internally with one 238 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 9. Reproductive system. A, Rhophodon kempseyensis, QMMO37096. B, Rhophodon paleihorpei, QMM055962. C, Egilodonia bainisdalensis, AMSC 154770. D, Egilodonta wyanbenensis, QMMO29270. E, Egilodonta bendethera, QMMO70395. F, Egilodonia pmcidentata , QMM049447. G Letomola contortus, QMM056119. H, Letomola lanalittleae, QMM056007. I, Macrophallikoropa belli, AMSC162176. J, Macmphall ikompa stenoumbilicata, QMM028661. Scale bar=2mm in A and D, and 1.6mm in all others. Abbreviations: dg, prostate; e, epiphallus; g, ovotestis; gd, hermaphrodite duct; gg, albumen gland; gt, talon; p, penis; ppt, preputial tube; prm, penis retractor muscle; ss, spermathecal stalk; ut, uterus; v, vagina; vd, vas deferens. NEW CHAROPID LAND SNAILS 239 longitudinal pilaster and long verge; length of verge greater than half the length of penis; sheath present. Vagina tubular, shorter than penis. Pallial cavity with complete secondary ureter; kidney almost unilobed with rectal lobe vestigial and pericardial lobe elongate, triangular and with apex reflexed. Radula with central tooth strongly tricuspid, considerably smaller than laterals; lateral teeth tricuspid; mesocone of first lateral tooth slender, slightly diamond-shaped; marginal teeth rounded; ectocone of outer marginal teeth split into four or more teeth; cndocone not split. Number of lateral teeth 6; marginal teeth 7; radular rows 1 15. Based on 3 dissected adults (QMM068759). DISTRIBUTION AND HABITAT. Known only from the type locality; living in litter on strongly vegetated limestone outcrop (including Ficus sp.) among open eucalypt woodland. REMARKS. Egilodonta bendethera sp. nov. is microsympatric with E. paucidentata sp nov., to which it bears a strong resemblance. The two species can be distinguished by a difference in shell colour (E. bendethera is lighter), and a difference in aperture shape. The whorls of E. bendethera are rounded above and below a flattened periphery, giving the aperture a less rounded appearance than E. paucidentata. E. bendethera is also larger, has wider rib spacing and has a narrower (wide cup-shaped instead of shallow saucer-shaped) umbilicus than E. paucidentata. Anatomically E. bendethera differs from E. paucidentata in that the epiphallus enters the penis through a long verge (more than half the length of the penis), and in having only one penial pilaster. In E. paucidentata the verge is considerably shorter (less than half the length of the penis), and there are six penial pilasters. Both species are similar to E. baimsdalensis , but differ in having weaker apertural barriers and less widely spaced teleoconch ribs. E. bendethera is readily distinguished from E. wyanbenesis by having less crowded ribs on the teleoconch. Egilodonta paucidentata sp. nov. (Figs 2-1 1,33; Table 2) ETYMOLOGY. Latin paucus, few, and dentatus, toothed; referring to the poorapertural dentition. MATERIAL. 1 10L0TYPE: QMMO70396, Marble Arch, S of Braidwood, NSW (35°43’19"S, 149°42T2”E), limestone outcrop/remnant rainforest, under rocks and in litter, 11.x. 1992, JS, GI. Height of shell=0.95mm, diameter= 1 .92mm, H/D=0.49, D/U=2.06, number of whorls=4.8, ribs on last whorl = 37. PARATYPES: All NSW. QMMO42081, 5SC/49RC, same data as holotype; AMSC 126761, 20+RC, W of Batemans Bay, Deua NP, Bendoura Ra, Reedy Ck, Marble Arch (35°43.6’S, 149°41.4’E), WFP, W.F. (Jr) Ponder; AMSC346063, 20+RC, W of Batemans Bay, Deua NP, Bendoura Ra, Reedy Ck, 5km E of Bcrlang Camping Ground, Marble Arch (35°43.6*S, 149°41 .4'E), MS. OTHER MATERIAL. QMM049447, M068004. DIAGNOSIS. Minute golden brown shell with a wide saucer-shaped umbilicus. Teleoconch with widely spaced, broad, protractively sinuated ribs. Mean ribs/mm 6.5 1 . Aperture with one parietal, one basal and one palatal barrier; barriers weakly developed, blade-like. Penis internally with a short apical verge and six longitudinal pilasters. DESCRIPTION. Shell discoidal, golden brown, with 4.5-5. 5 (mean 5.0) tightly coiled whorls, the last descending more rapidly in front. Shell diameter 1.72-2. 26mm (mean 1.99mm). Apex and early spire flat. Height of shell 0.80-1 . 1 1 mm (mean 0.93), H/D 0.41-0.54 (mean 0.47). Protoconch exsert of 1.2- 1.7 whorls. Apical sculpture with prominent, strongly curved radial ribs, becoming increasingly crowded toward prortoconch/teleoconch boundary; width of interstices (in multiples of rib width) 7 at apex, 2 at post-nuclear boundary; secondary sculpture of moderately spaced, discontinuous radial wrinkles. Teleoconch sculpture of bold, widely spaced, protractively sinuated radial ribs; 28-57 (mean 41 ) ribs on last whorl. Ribs/mm 4.87-8.91 (mean 6.5 1 ). Rib height equal to width; straight in section, rounded on top. Rib interstices on first post-nuclear whorl equal to width of 4-5 ribs; interstices on penultimate whorl equal to width of 3-5 ribs. Interstitial sculpture of fine radial riblets and crowded microspiral cords, about equal in strength. Microradials low, 10-13 between ribs on first post-nuclear whorl, 15-21 between ribs on penultimate whorl; microspiral cords crossing major ribs and forming strong elongate beads at their intersection with microradials. Umbilicus very wide, saucer-shaped, diameter 0.85- 1.07mm (mean 0.94mm), D/U 1.95-2.32 (mean 2.13). Sutures impressed, whorls rounded above and below a flattened periphery. Aperture ovately lunate; lip sinuous; shiny parietal callus present. One very fine parietal barrier, one basal and one palatal barrier present; barriers low blade-like. Based on 43 measured adults (QMMO70396, M049447, M068004, MO42081). 240 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 10. Penis, dissected. A. Rhophodon kempseyensis, QMMO37096. B, Rhaphodon palethorpei, QMM055962. C. Egilodonla baimsdalensis, AMSC 154770. D, Egilodonta wyanbenensis , QMMO29270. E, Egilodonta bendethera , QMMO70395. F, Egilodonta paucidentata, QMM049447. G, Letomola lanalittleae, QMM056007. H, Macrophallikoropa belli, QMMOI6867. 1, Maerophallikoropa stenoumbilicata , QMM028661. J. Elsothera brazieri, QMMO44810. K. Decoriropa lirata, QMMO42109. L, Coricudgia wollemianti, QMM028659. M, Marilyniropa jenolanensis , QMM037465. N, Gould iropa kaputarensis, QMM049175. O, Cralopa stmudensis, QMM029722. P, Gouldiropa carlessi , M032080. Q, Sharniropa wollondillyana , QMM029241. R, Acheronopa attunga, QMM049218. S, Hedleyropa yarrangobillyensis, QMMO39840. T, Scleridoropa sarahjaneae, QMM039974. U, Scleridoropa nandewar, QMM049158. V, Diphyoropa macleayana , AMSC 162 184. Scale bar= 1.25mm in L t and 1mm in all others. Abbreviations: e, epiphallus; pp, penial pilaster; ppt, preputial tube; pv, penial verge, vd, vas deferens. NEW CHAROPID LAND SNAILS 241 Genitalia with ovotestis containing two clumps of alveoli; more than two alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penis retractor muscle half the length of penis or greater; inserts at junction of penis and epiphallus. Epiphallus present; enters penis through a verge; length of verge less than half the length of penis. Epiphallus longer than penis; wider than vas deferens; pilasters longituduinal; retractor muscle not entwined with epiphallus. Penis sheath present; penis tubular; pilasters longitudinal; 6 pilasters present. Vagina shorter than penis; atrium shorter than penis; preputial tube absent. Pallial cavity with complete secondary ureter; primary ureter normal; kidney almost unilobed; apex reflexed. Mantle gland absent. Radular morphology unknown. Based on 1 dissected adult (QMM049447). DISTRIBUTION AND HABITAT. Known only from two limestone environments in the Bendoura/Minuma Ranges, S of Braidwood, SE NSW; living in litter on strongly vegetated limestone outcrops that support some remnant rainforest plant species; the surrounding countryside comprises open eucalypt woodland. REMARKS. Features that differentiate Egilodonta paucidentata sp. nov. from other members of the genus are given above. The species is most similar to E. bairsnsdalensis in general shell features and was mis-identified as that species by Stanisic (1990). E. paucidentata has com- paratively weaker apertural barriers and slightly more crowded ribs than E. bairnsdalenesis. Anatomically, the presence of a verge in E. paucidentata is a major difference from the vergeless condition in E. bairnsdalensis. The two species are allopatric and accurate locality data should eliminate the chance of misidentification. Additional fieldwork needs to be conducted in the vicinity of the limestone localities in order to determine whether this species is an obligate calciphile. Letomola Iredale, 1941 Letomola Iredale, 1941a: 267; Stanisic, 1990: 109; Smith 1992: 110. Letomala (error) Kershaw, 1956b: 9. TYPE SPECIES. Rhophodon contortus Hedley, 1924; by original designation. DIAGNOSIS. Small to minute charopids, with discoidal shells; colour brown with white radial streaks. Shell with or without apertural barriers; spire flat to depressed. Protoconch strongly exsert with sculpture of irregular pits; sometimes with narrow, very widely spaced, curved radial ribs; spiral cords absent but discontinuous spiral grooves sometimes present. Teleoconch sculpture degenerate, without distinct radial ribs, instead with broad, shingle-like, overlapping radial thickenings; secondary sculpture of low microradial ridges and low spiral cords that form weak beads at their intersection. Umbilicus very wide saucer-shaped to broad V-shaped. Kidney moderately bilobed with apex of pericardial lobe slightly reflexed. Penis internally with longi- tudinal pilasters and with or without vergic structures. DISTRIBUTION AND HABITAT. Letomola Iredale, 1941 is known only from the Macleay Valley, NE NSW. Species are obligate limestone dwellers usually living in litter or on the rock surface of outcrops among dry rainforest. REMARKS. Iredale (1941a) introduced Letomola for Rhophodon contortus Hedley, 1924 without giving clear reasons for the separation from Rhophodon Hedley, 1924. Kershaw (1956b) briefly discussed the genus. Subsequently Smith & Kershaw (1979) added Helix barrenensis Petterd, 1879 (type locality: Furneaux Group of islands, Bass Strait) without detailed justification for this placement. These authors probably based their decision on the presence of apertural lamellae since other details of the shell do not agree with those of Letomola. Smith (1992) maintained this classification. //. barrenensis is not discussed here but the fact that it has well developed radial sculpture on the teleoconch (as opposed to the degenerate sculpture of Letomola) indicates strongly that it does not belong to Letomola. The generic status of this species most probably needs to be determined in a context of the Tasmanian charopids. In spite of the absence of apertural barriers, the addition of L. lanalittleae sp. nov. does not alter the basic concept of Letomola as presented by Stanisic (1990). Barriers have developed independently in a number of Australian charopid genera and numbers of barriers vary from species to species. In the case of L. lanalittleae barrier loss merely represents an extreme example of reduction. Protoconch sculpture, teleoconch sculpture, and to a lesser degree, general shell form all agree with Stanisic’s (1990) view of the genus. A slightly amended diagnosis is presented to accommodate the idiosyncrasies of the new species. 242 MEMOIRS OF THE QUEENSLAND MUSEUM C FIG 1 1 . Pallial cavity. A, Rhophodon kempseyensis , QMMO37096. B, Rhophodon palethorpei, QMM055962. C, Egilodonta haimsdalemis , AMSCI54770. D, Egilodonta wyanbenensis , QMMO29270. E, Egilodonta bendethera , QMMO70395. F, Egilodonta paucidentata, QMM049447. G, Letomola contortus , QMM056 119. 1 1, Macrophallikoropa belli , AMSC162 1 76. 1, Macrophallikoropa stenoumbilicata, QMMD28661 . Scale bar = 1.5mm in B, and 1 .2mm in all others. Abbreviations: h, heart; hv, principal pulmonary vein; i, intestine; kdl, primary ureter; kd2, secondary ureter; kp, pericardial lobe of kidney; kr, rectal lobe of kidney; me, mantle collar; mg, mantle gland. NEW CHAROPID LAND SNAILS 243 Letomola Iredale, 1941 possesses a number of features that distinguish it from other charopid genera examined in this study. Having a shell with a pitted protoconch, supra-peripheral sulcus and degenerate teleoconch sculpture and a radula with greatly reduced central tooth are a unique combination of features that readily identify Letomola among all hitherto described eastern Australian Charopidae. Ecologically Letomola is distinguished by living on the limestone karst surface as well as in the interstitial litter. No other charopid is currently known to live on limestone karst in eastern Australia although this is common among groups such as the Hydrocenidae ( Georissa spp.), Helicinidae ( Pleuropoma spp.) and Pupillidae ( Gyliotrachela spp.) (Stanisic, pers. obs.). Solem (1974) suggested that ribs have a universal functional role among land snails of maintaining a dirt free shell surface for the snail as it crawls through moist, adherent litter. In which case the modified (reduced?) shell sculpture of Letomola may be related to its tendency to a non-litter lifestyle. A similar pattern is seen in the semi-arboreal charopids, Lenwebbia protoscrobiculata Stanisic, 1990 and Lenwebbia paluma Stanisic, 1993. Both have a shell with pitted apical sculpture and reduced teleoconch sculpture. Letomola contortus (Hedley, 1924) (Figs 9, 11-12, 14-18; Table 2) Rhophodon contortus Hedley. 1924: 220, pi. 32, figs 35-37; Kershaw, 1955: 30. Letomola contortus (Hedley). Iredale, 1941a: fig. 6; Iredale, 1941b: 2; Stanisic, 1990: 110. figs 67-69; Smith, 1992: 193. Letomala (sic) contortus (Hedley). Kershaw, 1956b: 9. MATERIAL. All NSW: QMMO37150, M019796, M0561 1 9, M059998, M052762, AMSC 1 57303, C168612. DIAGNOSIS. Shell minute, brown with white radial streaks. Whorls tightly coiled with a strong supraperiphcral sulcus. Protoconch pitted, without any radial ribs or spiral cords, spire depressed. Teleoconch lacking distinct radial ribs, instead with broad, shingle-like, radially ridged, overlapping thickenings. Umbilicus wide and shallow saucer-shaped. Two apcrtural barriers (one parietal, one palatal) present. Ovotestis con- sisting of one clump of alveoli with two alveolar lobes. Penis internally with an apical round pilaster and several longitudinal pilasters basally. DISTRIBUTION AND HABITAT. Eastern end of the lower Macleay Valley, NE NSW on limestone outcrops among dry rainforest. Found living on roof, walls and in floor litter of limestone caves, or sheltered vertical limestone rock faces. REMARKS. Letomola contortus (Hedley, 1924) is readily identified by its degenerate adult sculpture, very wide umbilicus and the presence of a supraperipheral sulcus. Differences from L. lanalittleae sp. nov. are presented under that species (see below). Anatomically L. contortus is not very different from other ‘dentate' charopids such as those grouped under Rhophodon Hedley, 1924 and Egilodonta Gabriel, 1930. However, the shell patterns are a dramatic departure from general charopid patterns seen in this study and probably relate to a slightly altered lifestyle not requiring the functional advantages of a ribbed shell. Interestingly this species lives on the limestone karst rather than the litter. Letomola lanalittleae sp. nov. (Figs 9-10, 12, 14-18, 33-34; Table 2) ETYMOLOGY. For Lana Little. MATERIAL. All NSW. HOLOTYPE: QMMO70397, Kempsey, WNW, c. 1.5km E Mt Sebastopol, Kempsey-Carrai Rd, NSW(30‘ , 57'(K) , 'S, 152°28’40"E),on rocks, drtTlm, 28.xi. 1989, JS, DP. Height of shell= 1 .38mm, diameter=2.86mm, H/D=0.48, D/U=3.62, number of whorls=4.7. PARATYPES: QMM032292, 19SC/44RC, same data as holotype; QMM037761, 50RC, Kempsey, WNW, c. 1.5km E Mt Sebastopol, Kempsev-Carrai Rd (30°57WS, 152°28'40"E), in litter, drflm,'28.xi.l989, JS, DP; QMM056007, 20SC/42RC, Kempsey. WNW, c. 1.5km E Mt Sebastopol, Kempsey-Carrai Rd (30 l, 57 , 00 , 'S. 152°28’40"E), on limestone and in litter, drf/lm, 8. i. 1995, JS, JC. OTHER MATERIAL. QMM049294. DIAGNOSIS. Shell moderately small, chocolate brown with cream blotches. Whorls tightly coiled with a strong supraperipheral sulcus; Umbilicus wide V-shaped. Spire flat with exsert protoconch. Apertural barriers absent. Pitted protoconch with faint, curved, irregular radial ribs and dis- continuous spiral grooves. Teleoconch without pronounced radial ribs, instead with broad, radially ridged, shingle-like overlapping thickenings. Ovotestis containing two clumps of alveoli, with more than two alveolar lobes per clump. Penis internally with round to ovate pilasters. DESCRIPTION. Shell small, chocolate brown with cream radial streaks, with 4. 7-5.6 (mean 5.3) tightly coiled whorls, the last descending very rapidly in front; supraperipheral sulcus present. Shell diameter 2.81 -3.74mm (mean 3.40mm). 244 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 12. Distribution of Letomola and Macrophallikoropa. Key: o Letomola contortus , ★ Letomola lanalittleae , □ Macrophallikoropa belli , • Macrophallikoropa stenoumbilicata, ▲ Macrophallikoropa depress ispira. Spire flat. Height of shell 1.30- 1.74mm (mean 1.48mm), H/D 0.40-0.49 (mean 0.44). Protoconch strongly exsert of 1.5- 1.7 whorls. Apical sculpture of irregular pits, with irregularly spaced, curved radial ribs; rib spacing variable, decreasing towards post-nuclear boundary (but not evenly); width of interstices (in multiples of rib width) 1 5 at apex; 2 at post-nuclear boundary. Scattered, discontinuous spiral grooves also present. Teleoconch sculpture consisting of broad, radially ridged, shingle-like overlapping thickenings. Microsculpture of low microradial radial ridgelets and microspiral cords, about equal in strength; forming weak round beads at their intersection. Umbilicus wide V-shaped, diameter 0.79- 1.1 8mm (mean 1.02mm), D/U 3.08-3.74 (mean 3.35). Base flattened with microradial ridgelets becoming more bladelike. Aperture ovately lunate; parietal callus present; apertural barriers absent. Based on 2 1 measured adults (QMM037761 , M032292, MO70397). Genitalia with ovotestis containing two clumps of alveoli; with more than two alveolar lobes per clump. Talon stalk more than 3 times diameter of talon; talon circular. Penial retractor muscle less than half the length of penis; inserting onto the penis apex. Epiphallus present, longer than penis; wider than vas deferens, entering penis through a simple pore. Penis tubular, pilasters modified (round); penial sheath present. Vagina shorter than penis. Pallial cavity with kidney moderately bilobed; apex very slightly reflexed. Radula with central tooth weakly tricuspid; lateral teeth tricuspid; first and second laterals equal in width; mesocone of first lateral tooth rounded at tip; marginal teeth flattened and rectangular; ectocone of outer marginal teeth split into three teeth; endocone not split. Number of lateral teeth 7; marginal teeth 7; radular rows 1 1 9. Based on 4 dissected specimens (QMM056007). DISTRIBUTION AND HABITAT. Letomola lanalittleae sp. nov is known from limestone outcrops among dry rainforest at the western end of the lower Macleay Valley; living on vertical limestone surfaces. REMARKS. Letomola lanalittleae sp. nov. is distinguished from L. contortus by its larger size, fiat spire (depressed in L. contortus ), narrower umbilicus and the absence of apertural lamellae (two present in L.contortus). Anatomically, L. lanalittleae differs by having 2 clumps of alveoli in the ovotestis compared with only 1 in L.contortus. The radula of L. lanalittleae also has NEW CHAROPID LAND SNAILS 245 TABLE 2. Shell measurements for species of Egilodonta and Letomola. (Abbreviations as in Table 1). f— Measurement Egilodonta bairnsdalensis Egilodonta wvanbenensis Egilodonta bendethera Egilodonta paucidentata Letomola contortus Letomola lanalittleae N 49 (*48) 16 (*15) 10 42 15 21 (*20) D 1.51-2.19(1.86) ± 0.09 1.78-2.39(2.14) ± 0.157 1.64-2.63(2.34) ± 0.32 1.72-2.26(1.99) ±0.11 1.78-2.23 (2.063) ± 0.133) 2.81-3.74(3.4) ± 0.28 PW 1.5-1.8(1.66) ± 0.07 * 1.4-1.7(1.573) ± 0.088 * 1.5-1.7(1.55) ± 0.07 1.2-1.7(1.56) ± 0.1 1.5-1.6(1.527) ± 0.046 1.5-1.7(1.61) ± 0.07 TW 2. 9-4.9 (3.27) ± 0.3) * 2.7-3.4 (3.073) ± 0.215 * 2. 8-4.2 (3.78) ± 0.46 3-4 (3.44) ± 0.2 2.2-2. 7 (2.467) ± 0.172 3.1-4(3.66) ± 0.25 NW 4.6-6.6 (4.93) ± 0.3 * 4.3-5. 1 (4.647) ± 0.185 * 4.3-5. 8 (5.33) ± 0.47 4. 5-5.5 (5) ± 0.19 3. 7-4. 2 (3.993) ± 0.175 4.7-5.6 (5.26) ± 0.27 H 0.73-1.08 (0.89) ± 0.09 0.91-1.15(1.033) ± 0.074 0.91-1.38(1.18) ± 0.16 0.8-1.11 (0.93) ± 0.07 0.85-1.05 (0.979) ± 0.057 1.3-1.74(1,48) ± 0.11 SP 0-0.12(0.03) ± 0.04 0-0.02(0.001) ± 0.005 0.01-0.18(0.07) ± 0.06) 0-0.14(0.04) ± 0.04 0-0.01 (0.002) ± 0.004 0.08-0.42 (0.25) ± 0.1 AH/AW 1.1-1.62(1.31) ± 0.12 1.29-1.76(1.5) ± 0.13 1.13-1.57 (1.33) ± 0.14 1.14-1.76(1.34) ± 0.12 0.97-1.31 (1.18) ± 0.08 0.71-1.05 (0.87) ± 0.09 UMB 0.67-0.95 (0.79) ± 0.06 0.8-1.14(0.985) ± 0.082 0.7-1.15(0.96) ± 0.13 0.85-1.07(0.94) ± 0.05 0.64-0.88 (0.783) ± 0.067 0.79-1.18(1.02) ± 0.11 * RIB 23-50 (33.57) ± 6.53 56-85(69.125) ± 7.83 27-47 (34.3) ± 7.04 28-57 (40.67) ± 7.07 - - RIBS/MM 2.78-7.27 (5.32) ± 1.11 8.49-13.05(10.32) ± 1.271 3.92-6.08 (4.67) ± 0.71 4.87-8.91 (6.51) ± 1.02 - - H/D 0.43-0.55 (0.48) ± 0.03 0.43-0.52 (0.48) ± 0.03 0.48-0.55 (0.5) ± 0.03 0.41-0.54 (0.47) ± 0.03 0.43-0.53 (0.48) ± 0.03 0.4-0.49 (0.44) ± 0.03 D/U 2.11-2.58(2.37) ± 0.11 2.08-2.31 (2.18) ± 0.07 2.19-2.63 (2.44) ± 0.12 1.95-2.32 (2.13) ± 0.09 2.38-2.79 (2.64) ± 0.13 3.08-3.74(3.35) ± 0.21 ♦ the first lateral tooth the same size as the other lateral teeth, whereas in L. contortus the first lateral is considerably larger than the others. L. contortus and L. lanalittleae are allopatric within the lower Macleay Valley and the considerable shell differences point to either a long period of isolation or a major shift in environmental regime. The larger size of L. lanalittleae may be a reflection of its adaptation to a less insular lifestyle above the litter on the limestone rock where living conditions are harsher and requiring of a larger shell. A somewhat analgous phenomenon is seen in populations of littoral zone mollusks where larger individuals can exist higher up on the seashore because they are less prone to dessication. In the case of Letomola , the evolution of a larger shell has probably allowed L. lanalittleae more flexibility in living space than L. contortus . Macrophallikoropa gen. nov. ETYMOLOGY. Greek macro, long and phallikos, penis; referring to the unusually long penial apparatus. TYPE SPECIES. Helix belli Cox, 1 864; herein designated. DIAGNOSIS. Shell minute, discoidal, with tightly coiled whorls. Colour golden brown to very light golden brown, sometimes with darker flammulations. Protoconch exsert; spire slightly elevated to depressed. Protoconch sculptured with narrow, high, continuous spiral lirae; num- ber of lirae 13-20. Secondary apical sculpture of radial elements present as narrow, discontinuous ridgelets (= pressure folds) between the cords. Teleoconch with slightly sinuated to straight, very crowded ribs; interstitial sculpture with microradials and microspirals about equal in strength, forming strong rounded beads at their intersection. Umbilicus wide U-shaped to very wide saucer-shaped. Kidney strongly bilobed. Epiphallus moderately long to very short. Penis with long to very long preputial tube; internally with very short apical verge or apical pore; no other visible thickenings. DISTRIBUTION AND HABITAT. Macro- phallikoropa gen. nov. ranges from SE NSW to SE Qld. Species live under logs and rocks in temperate to subtropical rainforest (humid and dry), dry vine thicket and wet to dry sclerophyll forest and occur under a variety of geological regimes ranging from sandstone to basalt and limestone. REMARKS. Allocharopa Iredale, 1937 (type species: Helix brazieri Cox, 1868) was intro- duced to accommodate a number of charopids 246 MEMOIRS OF THE QUEENSLAND MUSEUM from New South Wales, Victoria and Tasmania, including Helix belli Cox, 1864. One of the key characters given in the generic diagnosis was that the protoconch should be ‘radially ribbed as remainder of shell’. Closer inspection reveals that the conglomeration of species included in this genus represents a variety of protoconch types. This is a strong indication that the group is polyphyletic. H. belli has a prominent spirally lirate protoconch which immedately separates it from the other species listed by Iredale, 1937 under Allochciropa. As will be shown below, Allochciropa is a junior synonym of Elsothera Iredale, 1933 and most of the species included in this genus by Smith (1992) will need reallocation. A major revision of all the species is beyond the scope of this study and only H. belli and H. brazier i (see below) are dealt with herein. It is highly probable that the other species (mostly Tasmanian) will need to be reviewed in the context of that local charopid fauna since many of the southern Charopidae (Tasmanian, Victorian) appear unrelated to the eastern NSW radiation (Stanisic, unpubl. data). Smith & Kershaw (1979) placed H. belli in Roblinella Iredale, 1937 (type species Helix roblini Pctterd, 1879) which was introduced to accommodate a series of species with ‘spirally striated nuclear whorls'. These authors did so without giving reasons. As introduced by Iredale (1937), Roblinella contains species with dramatically different apical spiral lirae indicating that it is polyphyletic (possibly at least three genera [Stanisic, unpubl. data]). The spirally lirate protoconch of //. belli comes closest to that of the type, R. roblini , but differs in having more regular and crowded lirae. The very fine teleoconch ribs and coiling pattern of R. roblini (loose coiling with tiny umbilicus) are also features that indicate the two species are not congeneric. On available evidence H. belli deserves generic recognition and Macrophallikoropa gen. nov. is introduced for this widespread rainforest charopid plus two additional new species from very restricted localities in the Sydney Basin Bioregion. Macmphallikompa gen. nov. can be distinguished from all other genera in this study by the presence of a spiral protoconch. The studies of Stanisic (1987, 1990, 1993a) show that spiral apical sculpture has developed independently in distantly related groups. In contrast to the apical sculpture of Setomedea Iredale, 1941 (low. narrow, moderately spaced, strongly to slightly wavy spiral cords with vague, scattered radial ridges); Oreokera Stanisic, 1987 (low, crowded, regular to wavy spiral cords with radial rugosities developing at the protoconch/ teleoconch boundary); Mussonula Iredale, 1937 (low, fine to bold, crowded to very crowded spiral cords with curved radial ridges appearing toward the protoconch/teleoconch border); Sinployea Solem, 1983 (low, widely spaced radial cords); and Omphaloropa Stanisic, 1990 (low, moderately spaced spiral cords that become indistinct toward the inner half of the embryonic whorls), that of Macrophallikoropa features spiral cords that are strongly developed, crowded, narrow and high in section. In each of the above genera the lirae and general protoconch appearance seem macroscopically similar, but when viewed by scanning electron microscopy they reveal subtle but significant differences in architectural detail. These seemingly minor differences in protoconch sculpture correspond with major differences in shell form, teleoconch sculpture and anatomy suggesting that they are phylogenetically significant. Iredale (1937, 1941a,b) and Smith & Kershaw (1979) list a number of Australian charopids with a spirally lirate protoconchs, but do not provide adequate detail on microstructural diversity for this level of interpretation. An additional feature on the protoconch of Macrophallikoropa is the presence of short, discontinuous radial elements between the cords. These may represent shrinkage wrinkles associated with a dried periostracum (^pressure folds) in a manner similar to the discontinuous spiral elements shown by Solem (1984) to be present on the apices of species with predom- inantly radial protoconchs such as Discocharopa aperta (Mollendorff, 1888). The biogeography of this group is intriguing with a widespread species, M. belli (Cox, 1864) occurring mainly in the rainforests along the Great Dividing Range from SE NSW to SE Qld, in the sandstone country around Sydney and in several southern limestone localities (Jenolan, Bungonia, Wombeyan, Marble Arch, Wyanbene). Two additional species, M. depressispira sp. nov. and M. stenoumbilicata sp. nov., occur in restricted habitats at Jenolan Caves and Mt Coricudgy and environs respectively. Much of the area that encompasses these latter distributions is sandstone dominated by sclero- phyll forests with rainforest occurring only as small refugia in damp gullies or on rocky outcrops. Jenolan Caves (limestone) and Mt NEW CHAROPID LAND SNAILS 247 30 ° 32 ° 34 " Coricudgy (a basalt capped peak) represent two unusual isolated topographies within this sandstone landscape. Both maintain remnant rainforest communities. Judging by the widespread distribution of M. belli within currently dis- connected coastal rainforests blocks in eastern NSW and SE Qld, this species must have been present in these rainforests at a time when they were more extensive and more strongly interconnected. Presumably as rainforest became climatically restricted and fragmented in distribution from approx- imately the mid-Miocene onwards, M. belli would have continued to exist as isolated populations in widely separated refugia, probably along drainage lines, on mountain tops and in select topographic niches. Presumably also, climatically induced, shorter- term expansion of traditional M. belli habitat through the many wet climatic phases in the Plio-Pleistocene would have enabled M. belli to once again disperse and also occupy the wetter, non-rainforest habitats it now in does in southern areas, including much of the Sydney Basin Bioregion. M depressispira and M. stenoumbilicata appear to represent two climatically isolated derivatives of M. belli since they maintain many of the features of that species while also showing a number of significant departures from the ancestral pattern (depressed spire, narrower umbilicus, reduced length of preputial tube, lengthening of the umbilicus). Macrophallikoropa belli (Cox, 1864) comb. nov. (Figs 9-12, 14-18; Table 3) Helix belli C ox, 1864: 22; Cox, 1868: 17, pi. 6, fig. 3. Endodonta (Champa) belli (Cox). Cox, 1 909: 1 1 . Allochampa belli (Cox). Ircdalc, 1937: 326; Iredale, 1941a: 261; Kershaw, 1956a: 141; Smith, 1992: 181. Roblinella belli (Cox). Smith & Kershaw, 1979: 180. MATERIAL. LECTOTYPE (herein designated): AMSC205161. Height of shelH).84mm, diameter= 1.67mm, H/IX).5, D/LK3.04, number of whorls=4.5, ribs on last whorl=175. PARALECTOTYPES: AMSC 136899, 26RC, same data as lectotype. KEY MATERIAL. AMSC 162 176, 36 ° 200 km 148 ° 150 ° 152 ° 154 ° 156 ° FIG 13. Distribution of Egilomen , Whiteheadia and Decoriropa. Key: ■ Egilomen cochlidium , o Whiteheadia globosa , • Decoriropa lirata. 16SG2RC, S of Quirindi. Liverpool Ra, Cedar Brush Nature Res, Cedar Brush Gap, off Warrah Ck Rd, NSW, 4WD Track (31°5034'S, 15041.4'E), under logs, if with broken canopy, 8.xi.l985, IL, JW; QMM029277, 6SC, Misty Mt, c.l9km along Bolaro Mtn Rd, offBatemans Bay-Braidvvood Rd, NSW (35°37'S, 149°57‘E), wtrf, under logs, 0611990, JS, JC; QMMOI6867, 3SC, Lamington NP, Binna Burra, Qld Mt Hobwee circuit, nvf, under logs, 01.x. 1985, JS, DP, JC; AMSC 171 336, 2(HRC, Sydney, Bellevue Hill. Cooper Park NSW (33°53.26’S, 151°14.9’E), wsf, 23.V.1982, MS; QMMO 10760, 8RC, Dundiurabin, ca.2.3km N. Dundurrabin - Grailon Rd NSW (30°1 1 'S, 15r34*E).m\f, 15vi. 1981, WFP, JS, OGDP; QMM042080, 15RC, Marble Arch. S of Braidwood NSW (35"43'S, 1 49*42 ’E), Lsocrrf, 11.x. 1992, JS, GI: QMMO10512, 12RC, Mt Warning, NSW. summit (28‘24'S, I53°16’E), nvf, 07.vii.1981. WFP, JS. OG DP. OTHER MATERIAL. Southeastern NSW: QMM029275, AMSC357398, C3574024, C357410, C357415, C318486, QMM029263, M037985. Sydney Basin: AMSC377544, C3 1 8382, C3 1 8383, C3 1 8385, C3 1 8390, C3 1 8393, C3 1 8398, 248 MEMOIRS OF THE QUEENSLAND MUSEUM J K L FIG. 14. Dorsal view of shells. A, Letomola contort us, AMSC1 19349. B, Letomola lanalittleae , QMMO70397. C, Macrophallikoropa belli (Sydney), AMSC171336. D, Macrophallikoropa belli (Mt Warning), QMMO10512. E, Macrophallikoropa stenoumbilicata , AMSC205162. F, Macrophallikoropa depressispira , AMSC205163. G, Elsothera brazieri , QMM028775. H, Decoriropa lirata (Tarce), AMSC 135842. I, Decoriropa lirata (Kempsey), QMM052755. J, Decoriropa lirata (Wombcyan Caves), QMM064764. K, Coricudgia wollemiana. AMSC205164. K, Marilyniropa jenolanensis , AMSC205165. Scale bars=0.5mm in A, C-F, H; 0.8mm in I-L; 1mm in B, NEW CHAROPID LAND SNAILS 249 C3 18403X3 18407, C318422,C318412,C3 18416X3 18417, C3 1 84 19X"3 18420X3 18425X3 18435X3 18430X3 18431, C3 1 8439, C3 1 8444, C3 1 8432, C3 1 8433, C3 1 8442, C3 1 8438, C3 18440, C3 18441, C3 18443, C3 18483, C377468, QMM037777, M034782, MO31920. Northeastern NSW. QMM03 1 938, M0375 1 7, M029739, M049246, M05268 1 , M052699, M029749, AMSC377871, C377941, C377867, C377840, Cl 62009, QMMO10781, M010804, M010820, MO 10753, MO 16773, MO10892, MO9830, MO10539. SE Qld: AMSC 136491, C129240, QMMO10992, M012863, MO 12697, MO6079, M08361. DIAGNOSIS. Minute, golden brown to very light golden brown shell, sometimes with darker flammulations. Spire flat, protoconch with crowded spiral lirae. Teleoconch with slightly sinuate to straight crowded ribs; interstitial sculpture with microradials and microspirals about equal in strength, forming strong beads. Umbilcus very wide cup-shaped, mean D/U 2.75. Penis internally with very short penial verge, very long preputial tube. Epiphallus very short. DESCRIPTION. Shell tiny, golden brown (occasionally with darker flammulations), with 3. 8-5. 5 (mean 4.46) tightly coiled whorls, the last descending more radidly in front. Sutures impressed, whorls evenly rounded above and below periphery. Shell diameter 1.43-2.22mm (mean 1.76mm). Apex and early spire flat. Height of shell 0.73- 1.1 2mm (mean 0.92mm), H/D 0.48-0.56 (mean 0.52). Protoconch exsert of 1.5- 1.8 whorls. Apical sculpture of crowded, narrow spiral lirae; number of lirae 14-20. Secondary apical sculpture of discontinuous radial ridglets that may be periostracal pressure folds. Teleoconch sculpture of very crowded, protectively sinuated radial ribs; 1 02-248 (mean (179) ribs on last whorl. Ribs/mm 14.69-42.19 (mean 32.65). Ribs medium to narrow; height equal to width; straight in section; rounded on top. Rib interstices on the first post-nuclear whorl equal to width of one to four major ribs; interstices on the penultimate whorl equal to one to three ribs. Interstitial sculpture of fine radial riblets and spiral cords, about equal in strength. Microradials low; 2-7 between ribs on first postnuclear whorl, 3-6 between ribs on penultimate whorl. Microspirals forming strong round to elongate beads at their intersection with the microradials; crossing major ribs. Umbilicus very wide, cup-shaped; diameter 0.42-0.84mm (mean 0.64mm), D/U 2.31-3.43 (mean 2.76). Aperture ovately lunate, with parietal callus. Based on 31 measured adults (AMSC171336, C3 18439, C205161, QMM010760, MO10512, M042080, M032185). Genitalia with ovotestis containing 2 clumps of alveoli with 2 alveolar lobes per clump. Talon stalk shorter than or equal to talon diameter; talon circular. Penial retractor muscle less than half the length of the penis inserting at or near the junction of the penis and epiphallus, on either the penis or epiphallus. Epiphallus short wider than epiphallus, entering penis through a verge; length of verge less than half the length of the penis. Penis tubular, internally with an apical verge but without any other noticeable thickenings. Vagina shorter than or equal in length to penis; atrium shorter than penis; long preputial tube present. Kidney bilobed; apex of pericardial lobe slightly reflexed. Radula with central tooth strongly tricuspid; lateral teeth tricuspid; first and second laterals equal in width. Mesocone of first lateral tooth slender, slightly diamond-shaped; marginal teeth skewed towards the centre; ectocone of outer marginal teeth split into two teeth; endoconc not split. Number of lateral teeth 5; marginal teeth 6; radular rows 78-80. Based on 7 dissected specimens (AMSC162176, QMM029277, M016867, M032185). DISTRIBUTION AND HABITAT. Macrophal - likoropa belli is a wide ranging species inhabiting a range of forest types from dry sclerophyll to rainforest on limestone, volcanics and sandstone; found living under logs and rocks. REMARKS. Macrophallikompa belli (Cox, 1 864) comb. nov. is sympatric with M. depressispira sp. nov. at the Jenolan Caves limestone locality but can be distinguished from that species by its smaller size, less open umbilicus and flatter spire. M. belli differs from M. stenoumbilicata sp. nov. by its smaller size and more open umbilicus. A/, belli is readily distinguished from other co-habiting charopids through a combination of very small size, spiral protoconch, strongly curved, narrow and very crowded radial ribs, wide cup-shapcd umbilicus and the lack of apertural lamellae. Some Rhophodon spp. are similar in general shell form (shape, teleoconch ribbing and umbilical width) but are distinguished by having radially ribbed protoconchs and apertural lamellae. Decoriropa lirata (Cox, 1864) comb. nov. also has a small discoidal shell with wide umbilicus but has a radial protoconch and widely spaced, bold ribs on the teleoconch. Anatomically the long penial apparatus of M. belli is unusual among the eastern Australian Charopidae and hitherto 250 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 15. Protoconch of shells from above. A, Letomola con tortus, AMSC 119349. B, Letomola lanalittleae , QMMO70397. C, Macrophallikoropa belli (Sydney), AMSC 171336. D, Macrophallikoropa belli (Mt Warning), QMMO10512. E, Macrophallikoropa stenoumbilicata , AMSC205162. F, Macrophallikoropa depressispira , AMSC205163. G, Elsothera brazieri , QMM028775. H, Decoriropa lirata (Taree), AMSC 135842. I, Decoriropa lirata (Kempsey), QMM052755. J, Decoriropa lirata (Wombeyan Caves), QMM064764. K, Coricudgia wollemiana , AMSC205164. K, Marilyniropa jenolanensis, AMSC205165. Scale bars=0. 1 5mm in A, E, F, H, I; 0. 1 mm in C, D; 0.2mm in B, G, J-L. NEW CHAROPID LAND SNAILS 251 matched only by that of Rotachciropa densilamellata Stanisic, 1990. Stanisic (1990) suggested that in the case of Rotachciropa this development was organ elongation in response to a marked increase in whorl numbers from the clade average. However, this is not the case in A/. belli where whorl numbers are fairly similar to both A/. stenoumbilicata and M. depressispira that lack this condition. It could be that this elongated condition is a retained ancestral feature. From a practical viewpoint it is difficult to undertstand how the penial apparatus could function as an eversible organ. M. belli is a relatively widespread species in the context of eastern Australian charopids, especially for a species that inhabits moist rainforest. Discocharopa aperta (M ollcndorff, 1888) has a wider distribution (Solem, 1989) than A/, belli but that species also shows a greater environmemtal adaptability in preferring drier habitats such as deciduous vine thickets. But while M. belli distribution is largely restricted to wetter environments these do cross a wide range of floristic and geological boundaries, from scelorphyll forest to rainforest and through basalt, limestone and sandstone landscapes. A/. belli has s significant presence in the Sydney Basin Bioregion where it persists in quite marginal habitats (e.g. Sydney Botanic Gardens) thereby also implying an inherent adaptability. That M. belli has been able to disperse into and adapt to this broad set of habitats across a wide geographic area indicates that the species has a long temporal history. In spite of the large amount of material in the AM and QM, few of these specimens are adult. Adult specimens are readily identifiable by the more rapid descension of the last whorl, which effectively elevates the remainder of the shell's spire. Hence, the shell datasets do not allow any rigorous analysis of geographic trends in shell variation. Of the specimens measured those from Marble Arch were distinctive in displaying a slightly more elevated spire and a more closed umbilicus. Not all available material in the AM and QM has been listed herein. The listing is meant to be a guide to significant datasets, general distribution and key localities for the species. Additional lots, in many cases containing 1-3 specimens and duplicating many of the listed localities, are held in the AM and QM. The sheer quanitiy of material is a poignant indication of the wide- spread distribution of A/. belli compared with other species discussed herein. Macrophallikoropa stenoumbilicata sp. nov. (Figs 9-12, 14-18; Table 3) ETYMOLOGY. Greek stenos, narrow and Latin umbilicus ; referring to the relatively narrow umbilicus. MATERIAL. All NSW. HOLOTYPE: AMSC205162, Mt Coricudgy, E Rylstone, southern slope (32°50.82’S, 150°2’E), in litter off road, 2.x. 1983, PHC. Height of shcll=1.03mm, diameter= 1.92mm, H/D=0.54, D/U=3.20, number of whorls=4.2, ribs on last whorl=197. PARATYPES: AMSC 162008, 12RC, same data as holotype; AMSC 1 62007, 2RC, Mt Coricudgy, E Rylstone, Hanging Swamp (32°50.2’S, 150°2!.2'E), in litter, 2.x. 1983, PHC; QMM028661. 4SC/3RC, slopes of Mt Coricudgy, W of Rylstone (32°51'S, 1 50°2TE), under logs, temperate rainforest, 15.LI990, JS, JC; QMMO44706, IRC, summit, Mt Coricudgy, N Rylstone (32°49'48 M S, 150°20 , 24”E), under rocks, remnant rainforest, IO.i.1993, JS, JC. OTHER MATERIAL. All NSW. QMM044716, M044724, M059873, MO59860. DIAGNOSIS. Shell minute, golden brown to very light golden brown, sometimes with darker flammulations; whorls tightly coiled. Spire and apex slightly elevated, protoconch with crowded spiral lirae. Teleoconch with almost straight, very crowded ribs. Umbilicus wide U-shaped, mean D/U 3.1 1. Penial verge absent; moderately long preputial tube present, epiphallus moderately long. DESCRIPTION. Shell minute, light to golden brown (occasionally with darker flammulations), with 4. 0-4. 7 (mean 4.33) tightly coiled whorls, the last descending more rapidly in front. Sutures strongly impressed, whorls evenly rounded abovcT and below the periphery. Shell diameter 1.6 1-1. 22mm (mean 1.95mm). Apex and early spire slightly elevated. Height of shell 0.77-1. 19mm (mean 1.00mm), H/D 0.48-0.54 (mean 0.51). Protoconch slightly exsert, of 1.5- 1.8 whorls. Apical sculpture of crowded, narrow spiral lirae; number of lirae 17. Secondary apical sculpture of discontinuous radial ridglets that may be periostracal pressure folds. Teleoconch sculpture of very crowded, weakly protractvely sinuated radial ribs; 147-197 (mean 171) ribs on the last whorl. Ribs/mm 26.43-32.66 (mean 28.94). Ribs narrow; height equal to width; straight in section, rounded on top. Rib interstices on the first post-nuclear whorl equal to width of three to four major ribs. Interstitial sculpture of fine radial riblets and spiral cords, about equal in strength. Microradials low, forming prominent, elongate beads at their intersection with the microradials. Umbilicus wide U-shaped, diameter 0.53- 0.73mm (mean 0.63mm), D/U 3.04-3.20 (mean 3.11). Aperture ovately lunate. Based on 6 measured adults (AMSC 162008, C205162). 252 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 16. Lateral view of shells. A. Letomola contortus, AMSC1 19349. B. Letomola lanalittleae, QMMO70397 C, Macrophall ikoropa belli (Sydney), AMSC171336. D, Macrophallikoropa belli (Mt Warning) QMMO10512. E, Macrophallikoropa stenoumbilicata, AMSC205I62. F, Macrophallikoropa depressispira AMSC205163. G Elsothera brazieri, QMM028775. H, Decoriropa lirata (Taree), AMSCI35842 1 Decoriropa lirata (Kempscy), QMM052755. J, Decoriropa lirata (Wombeyan Caves), QMM064764 K.’ Coricudgia wollemiana, AMSC205 164. K. Marilyniropa jenolanensis, AMSC205 1 65. Scale bars=0.5mm in A, C, D, H-J; 0.8mm in E, F; 1mm in B, G, K, L. NEW CHAROPID LAND SNAILS 253 Genitalia with ovotestis containing two clumps of alveoli; with two alveolar lobes per clump. Talon stalk shorter than talon; talon circular. Epiphallus moderately long, shorter than penis; wider than the vas deferens, entering penis through a simple pore. Penial retractor muscle inserted on head of penis. Penis tubular, internally without any visible thickenings; moderately long preputial tube present. Vagina shorter than penis, without unusual features. Kidney bilobed with apex of pericardial lobe weakly reflexed. Radular morphology unknown. Based on 3 dissected adults (QMMO42150, M028661). DISTRIBUTION AND HABITAT. Macro - phallikoropa stenoumbilicata sp. nov. is known from rocky refugia on and in the vicinity of Mt Coricudgy at the northern end of the Wollemi NP; found living under logs and rocks in remnant rainforest and eucalypt forest. REMARKS. The shell of Macrophallikoropa stenoumbilicata sp. nov. is larger in size than that of M. belli , and has a smaller umbilicus (wide U-shaped), slighly more elevated spire and radial ribs that arc only weakly protractively sinuated. Anatomically M. stenoumbilicata differs from M. belli by lacking a penial verge, having a longer epiphallus and shorter preputial tube. M. stenoumbilicata differs from M. depress ispira sp. nov. by having a smaller shell with more elevated spire (rather than flat to slightly elevated) and an umbilicus that is less excavate (wide saucer-shaped in A/, depressispira). M. stenoumbilicata is confined to the northern end of the Wollemi NP which is characterised by basalt capped peaks that are scattered among a landscape of Narrabeen sandstone. These peaks support rainforest and wet sclerophyll forest near their summits and southeastern aspects and were probably critical refugia for the survival of this species during the drier climatic phases of the Plio-Pleistocene. Much of this area still needs to be investigated in order to determine the full extent of the species range. Macrophallikoropa depressispira sp. nov. (Figs 12, 14-17, 33-34; Table 3) ETYMOLOGY. Latin depressus, low and spira, spire; referring to the depressed spire. MATERIAL. All NSW. HOLOTYPE: AMSC205163, Jenolan Caves, E side Grand Arch near Blue Lagoon, near Devils Coach house (33°49.3’S, 150°1.6’E), in litter. among rocks and grass, ll.xii.1979, WFP, JS . Height of shell=1.14mm, diameter=2 .2 1 mm, H/D=0.52, D/U=ratio 2.60, number of whorls=5.0, ribs on last whor 1=293. PARATYPES. AMSC 124275, many RC, same data as holotype; AMSC6377 1 , 1 RC, Jenolan Caves, same data as holotype; QMM09699, many RC, Jenolan Caves, entrance to the Devils Coachhouse, litter, 13.xii. 1 979, JS. DIAGNOSIS. Shell minute, golden brown to very light golden brown, with tightly coiled whorls. Spire concave, protoconch with crowded spiral lirae. Teleoconch with very crowded radial ribs. Umbilicus very wide saucer-shaped, mean D/U 2.57. Anatomy unknown. DESCRIPTION. Shell minute, golden brown with 4.6-5.2 (mean 4.96) tightly coiled whorls, the last descending more radiply in front. Sutures weakly impressed, whorls evenly rounded above and below periphery. Shell diameter 1.97-2.37mm (mean 2.16mm) Spire depressed (concave). Height of shell 0.90- 1.1 4mm (mean 1.06mm), H/D 0.46-0.52 (mean 0.49). Protoconch slightly exsert of 1.6- 1.8 whorls. Apical sculpture of crowded, narrow spiral lirae; number of lirae 13. Secondary apical sculpture of discontinuous radial ridglets that may be periostracal pressure folds. Teleoconch scuplture of crowded, weakly protractively sinuated radial ribs; 165-293 (mean 222) ribs on the last whorl. Ribs/mm 37.97-26.39 (mean 28.94). Ribs narrow, height equal to width; straight in section; rounded on top. Rib interstices on the first post-nuclear whorl equal to width of two to three ribs. Rib interstices sculptured with low radial ridges and low microspiral cords frorming elongate to round beads where they intersect. Microradials between ribs on first postnuclear whorl 3; between ribs on penultimate whorl 2-4. Umbilicus very wide saucer shaped, diameter 0.76-0.9 1mm (mean 0.84mm), D/U 2.34-2.72 (mean 2.57). Aperture ovately lunate; parietal callus present. Based on 19 measured adults (AMSC124275, C205163, C124275). Anatomy unknown. DISTRIBUTION AND HABITAT. Macro- phallikoropa depressispira sp. nov. is known only from the type locality at Jenolan Caves; found in litter collected from among limestone rocks. REMARKS. Macrophallikoropa depressispira sp. nov. differs from both M. belli and M. stenoumbilicata in having a larger shell with depressed spire (rather than flat to slightly elevated) and an umbilicus that is more excavate 254 MEMOIRS OF THE QUEENSLAND MUSEUM J K L FIG 1 7. Ventral view of shells. A, Letomola con tortus, AMSC 1 19349. B, Letomola lanalittleae, QMMO70397. C, Macrophallikoropa belli (Sydney), AMSC171336. D, Macrophallikoropa belli (Mt Warning), QMMO10512. E, Macrophallikoropa stenoumbilicata , AMSC205162. F, Macrophallikoropa depressispira , AMSC205163. G, Elsothera brazieri , QMM028775. H, Decoriropa lirata (Sydney), AMSC427984. 1, Decoriropa lirata (Kempsey), QMM052755. J, Decoriropa lirata (Wombeyan Caves), QMM064764. K, Coricudgia wollemiana , AMSC205164. K, Marilyniropa jenolanensis , AMSC205165. Scale bars=0.5mm in A, C-F, I; 0.8mm in H, J; 1mm in B, G, K, L. NEW CHAROPID LAND SNAILS 255 (wide saucer-shaped). It can be readily distinguished from other sympatric charopids by its protoconch features. There has been considerable land snail collecting undertaken in the Blue Mountains area and Jenolan Caves area in particular, suggesting that the distribution is accurate and that this species is a true, narrow limestone endemic. No live material of A/, depress ispira, which could be critical to a better understanding of character shifts within the genus, has yet been collected. Elsothera Iredale, 1933 Elsothera Iredale, 1933: 53; Iredale, 1937: 324; Iredale, 1941a: 267; Kershaw. 1956a: 140; Burch, 1976b: 132; Stanisic, 1990: 160; Smith, 1992: 187; Stanisic, 1996: 345. Allocharopa Iredale, 1937: 326; Iredale 1941a: 269; Kershaw, 1956a: 141; Burch, 1976b: 132; Smith, 1992: 181. TYPE SPECIES. Helix sericatula Pfeiffer, 1850; by original designation. DIAGNOSIS. Shell very small with evenly to loosely coiled whorls. Colour greyish brown with darker streaks or monochrome brown. Protoconch sculpture of thin, regularly spaced, curved radial ribs (sometimes absent) and with prominent to very weak, sometimes discon- tinuous, microspiral cords. Teleoconch with crowded to moderately widely spaced, protractively sinuated radial ribs. Microsculpture of bladed microradial ribs crossed by low microspiral cords; prominent beads formed at their intersection. Umbilicus ranging from widely open to closed. Epiphallus and vas deferens long and strongly coiled about penial retractor muscale and/or penis. Penis with numerous longitudinal pilasters. DISTRIBUTION AND HABITAT. Elsothera Iredale, 1933 is widespread through central western Qld to eastern and central western NSW, Victoria and eastern South Australia. Records of the genus from Tasmania (Smith & Kershaw, 1979; 1981) need to be verified; living under rocks and logs in wet and dry sclerophyll forest, vine thickets and open eucalypt woodland. REMARKS. In dealing with Macrophallikoropa belli (Cox, 1 864) comb. nov. it became apparent that Allocharopa Iredale, 1937 waspolyphyletic. The mixture of species placed here by Iredale (1937) comprised some with primarily radially ribbed protoconchs (sometimes accompanied by a secondary spiral sculpture) and some, such as M. belli, which had strong spiral lirae. While a review of all the ‘radially ribbed' species is beyond the scope of this study a cursory study of the type of Allocharopa (= Helix brazieri Cox, 1868) revealed that it has significant characters (shell and anatomy) in common with Elsothera . In particular the protoconch sculpture and unusual epiphallic coiling were similar to E. sericatula which is sympatric with the former in the Sydney Basin. These characters are also shared with E. hemttorum Stanisic, 1996 from the Qld Brigalow Lands. On these shared characters H. brazieri is reassigned to Elsothera and Allocharopa is relegated to synonymy with Elsothera, In as much as Helix brazieri (Cox 1868) is placed in Elsothera Iredale 1933 it should be noted that as currently defined (Smith, 1 992) this genus probably is polyphyletic and the generic diagnosis herein should be regarded as tentative. The protoconch sculpture within the genus comprises several distinct forms including crowded curved radials with weak, sometimes discontinuous microspiral wrinkles (pressure folds) as seen in in E. brazieri (Cox, 1 868) comb, nov. and E, sericatula (Pfeiffer, 1850); curved radials with distinct continuous, crowded microspiral cords (E. nautilodea); and crowded microspiral cords with weak to obsolete microradials (E. heuittorum Stanisic, 1996, E. funerea (Cox, 1 868). In the case of E. genithecata Stanisic, 1990 the radial ribs are stronger and more widely spaced and this species may yet be shown to belong to another clade (Stanisic, 1 990; 1996: unpubl. data). There is also a considerable variation in shell colour within the genus ranging from grey to greyish brown with darker radial flecks to all brown. Elsothera requires revision but this is beyond the scope of this study. A key character that appears to bind the group is the extensive coiling of the epiphallus which in the case of E. brazieri is coiled around the penis. This feature has not been seen in any other eastern Australian charopid genus but because of the comparative dearth of anatomical studies within the group, its phylogenetic significance has yet to be fully determined. Elsothera brazieri (Cox, 1868) comb. nov. (Figs 10, 14-18, 20-22, 33-34; Table 3) Helix brazieri Cox, 1868: 14. Endodonta (Charopa) brazieri (Cox). Cox, 1909: 13. Allocharopa brazieri (Cox). Iredale, 1937: 326; Iredale 1941a: 269, fig. 6 (part); Kershaw, 1956a: 141; Smith & Kershaw, 1979: 161; Smith, 1992: 181. 256 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 18. Radula. A-C, Letomola contortus , QMM056119. D-F, Letomola lanalittleae, QMM056007. G-I belIi ’ QMMO 16867. J-L. Elsotliera brazieri, QMMO44810. M-O. Decoriropa lirata QMM029722. Scale bars=0.0 1mm in A-I.M-O; 0.02mm in J-L. NEW CHAROPID LAND SNAILS 257 MATERIAL. HOLOTYPE. AMSC63498, Cooks River, Sydney S (33°.55’S, 151°.70’E), under stones in dry places, 1868, J. Brazier. Height of shells 1.69mm, diameter=3.43mm, H/D-0.49, D/U=3.46, whorls=4.4, ribs on last whorl- 138. KEY MATERIAL. All NSW. QMMO44810, 5SC, Blue Mountains NP. Federal Pass. Echo Pt-Scenic Rwy, Katoomba, 33°44’06"S, 1 50°38'42 M E, wt6rt7m, under logs and rocks, 1 6.i. 1 993, JS. JC; QMM028775, 2SC/7RC, Jenolan Caves (33°49’S, 150°0rE), lm, under rocks, ()9.i.l990; JS, JC. OTHER MATERIAL. All NSW. Sydnev Basin. AMSC 159053, Cl 746 14, C346733, C348657, C368692, Cl 09350, Cl 549 14, Cl 549 17, Cl 54926, C3 18526, C3 19927, C3 19931, C3 19932, C3 19933, C3 19934. C3 19935, C3 19936, C3 19937, C3 19938, C3 19939, C346810, C 148 1 1 1 , C3468 1 9, C367089, QMMO 1 6940, M028668. M042139, M042143, M044782, M07I653, M034741, M071657. DIAGNOSIS. Small very small with evenly coiled whorls. Protoconch with curved, crowded radial ribs that become more crowded at the protoconch/teleoconch boundary and weaker, sometimes discontinuous spiral cords. Teleoconch sculpture of numerous, crowded and prominent curved radial ribs, mean ribs/mm 15.32. Microsculpture of strongly bladed microradial ribs and low, crowded microspiral cords with beads formed at their intersection. Umbilicus wide V-shaped, mean D/U 3.58. Epiphallus long and coiled around penial retractor muscle and basal part of penis. Penis internally with five longitudinal pilasters. DESCRIPTION. Shell very small, greyish brown in colour with darker radial streaks, 3 8 . 4.4 (mean 4.17) evenly coiled whorls. Shell diameter 2. 72-3. 99mm (mean 3.37mm). Spire flat. Height of shell 1.64-1. 94mm (mean 1.80mm), H/D 0.49-0.58 (mean 0.52). Protoconch of 1.5-1 . 8 whorls with primarily radial sculpture consisting of 80-90 regularly spaced, weakly curved radial ribs that become more crowded at the protoconch/teleoconch boundary and very weak, low regularly spaced spiral lirae that cross the radial ribs. Ribs wide, height less than width; spacing uniform; width of interstices (in multiples of rib width ) 4-5 at apex. Teleoconch sculpture of bold, protractively sinuated, crowded radial ribs; 136-206 (mean 1 62) ribs on the last whorl, ribs/mm 1 3.70-1 7.96 (mean 1 5.30). Ribs medium in size height greater than width, straight in section, rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 6-8 ribs; interstices on the penultimate whorl equal to width of 6-7 ribs. Microsculpture of low microradial ribs and stronger microspiral cords that cross the microradials and form strong elongate to round beads at their intersection. Microradials low; 6-8 between ribs on first postnuclear whorl; 10-12 between ribs on penultimate. Umbilicus wide V-shaped, diameter 0.80- 1.3 1mm (mean 0.97mm), D/U 3.05-4.04 (mean 3.57). Sutures weakly impressed, whorls rounded above and rounded below a rounded periphery. Aperture roundly lunate, parietal callus present. Based on 10 measured specimens (QMM028775, AMSC63498). Genitalia with ovotestis containing 2 clumps of alveoli; with more than 2 alveolar lobes per clump. Talon stalk narrower than talon diameter; talon circular. Penial retractor inserting onto the penis head. Epiphallus longer than penis, coiled around basal part of penial retractor muscle and basal part pof the penis; wider than the vas deferens. Penis tubular internally with numerous longitudinal pilasters. Pallial cavity with kidney very weakly bilobed; apex slightly reflexed. Radula with central tooth strongly tricuspid and smaller than the first lateral; lateral teeth tricuspid.; Ectocone of outer marginal teeth split into four or more teeth; endocone not split. Number of lateral teeth: 4, marginal teeth: 8 ; radular rows: 85. Based on 2 dissected specimens (QMMO44810). DISTRIBUTION AND HABITAT. Elsothera hrazieri is known only from the Sydney Basin and the northern part of the SE Highland Bioregions; living under logs and rocks in temperate rainforest and wet and dry sclerophyll forest. REMARKS. Elsothera hrazieri is micro- sympatric with E. sericatula (Pfeiffer, 1850) but is distinguished by having a widely open rather than closed umbilicus, having the radial ribs more widely spaced (coarser sculpture) and in having the epiphallus more strongly coiled and twisted around the penis. Diphyoropa saturni (Cox, 1864) comb, nov., which is also widely sympatric with E. hrazieri in the Sydney Basin, may be confused with the latter judging by identifications on existing museum material. However, D. saturni is readily distinguished by the bolder, more widely separate ribs on the teleoconch and in having a protoconch with a bimodal sculpture wherein initial spirals give way to radial ribs and ridges on the latter part of the embryonic whorls. Anatomically, D. saturni lacks the coiled epiphallus of Elsothera spp. 258 MEMOIRS OF THE QUEENSLAND MUSEUM mm n rtm**. f tf iff? f . r. rr a -fr FIG 19. Radula. A-C, Coricudgia wollemiana , QMM028659. D-F, Marilyniropa jenolanensis , QMM037465. Scale bars=0.01 in A-C, F; 0.02mm in D,E. Most of the El sot hera group have an open umbilicus. E. nautilodea (Cox, 1866) and E. genithecata Stanisic, 1 990 from NE NSW and E. sericatula from the Sydney Basin are exceptions. In the present case the major shift in umbilical width from closed in E. sericatula to widely open in the E. brazieri may be linked to underlying species recognition changes that have occurred under conditions of congeneric sympatry. Also E. brazieri appears to be less abundant in the field than the sympatric E. sericatula based on the quantity of material in museum collections. Reasons for this are not immediately obvious. Egilomen Iredale, 1937 Egilomen Iredale. 1937: 328; Iredale, 1941a: 267, 269; Kershaw, 1955: 29; Burch, 1976b: 132; Stanisic, 1990: 193; Smith, 1992: 187. TYPE SPECIES. Helix cochlidium Cox 1868; by original designation. DIAGNOSIS. Shell very small, colour white. Protoconch with relatively widely spaced, very bold, slightly curved radial ribs (approx. 30) that do not become more crowded, even at the protoconch/teleoconch boundary; spiral elements vague, consisting of discontinuous periostracal wrinkles (?=pressure folds). Teleoconch with broad, widely spaced, weakly protractively sinuated radial ribs; secondary sculpture of prominent crowded, low microspiral cords and relatively less prominent microradials ribs that form strong beads at their intersection. Umbilicus moderately wide U-shaped. Anatomy unknown. DISTRIBUTION AND HABITAT. Egilomen is restricted to the Clarence and Richmond River drainages in NE NSW. However, it would not be surprising if the range of the genus is extended beyond this area. Material in the collections of the QM from the Macleay Valley, NE NSW, appears to fall within Egilomen based on protoconch characters (Stanisic, unpubl. data). Egilomen prefers dry subtropical rainforest among litter and rocks; historical records suggest that it can also be found in riparian environments. REMARKS. Egilomen was introduced by Iredale (1937) for Charopidae that had a wide umbilicus, smooth protoconch and radial ribs on the teleoconch. Helix cochlidium Cox, 1 868, as type: H. lirata Cox. 1 864, H. barrenense Pettard, 1879, H. pexum Cox, 1868 and H. saturni Cox, 1864 were included. However, species included here by Iredale do not have smooth protoconchs and in fact cover an assortment of protoconch sculptures ranging from bold radial ribs, to radial ribs crossed by spiral cords and in one case a uniquely bimodal pattern of spiral and radial elements (//. saturni). Stanisic (1990), on the basis of limited comparative material, redefined Egilomen to include taxa with a protoconch sculpture of prominent radial ribs with spiral NEW CHAROPID LAND SNAILS 259 elements reduced or absent and a teleoconch sculpture of bold radial ribs. Anatomically, the defining features were a strongly bilobed kidney and one longitudinal penis pilaster with an accessory circular pad. Stanisic (1990) included only the type and a new species, E. globosa. Although not reviewed, Stanisic (1990) suggested that most other species included in Egilomen by I redale (1937) belonged elsewhere. A redefinition of Egilomen was considered necessary in order that other taxa dealt with herein could be placed into appropriate taxonomic context. During the course of this study it became apparent that Stanisic's (1990) interpretation of Egilomen , and in particular E. cochlidium, was somewhat expansive. In addition, closer inspection of the protoconchs of the types of both E. cochlidium and E. globosa revealed differences indicating that these two species are generically distinct on the basis of discriminatory criteria used in the current study. Hence, Egilomen is herein once again redefined and restricted to include only the type. Revised diagnoses of both Egilomen and E. cochlidium are presented together with a revised list of ‘material studied' for the latter species. Consequently, the distribution of E. cochlidium is herein restricted to include only the region surrounding the Richmond and Clarence River drainages. E. globosa is not able to be placed in any existing genus and hence is herein reassigned to Whiteheadia gen.nov. Two additional species, H. lirata and H. saturni , previously placed here by Iredale (1941a), are also reviewed and generically re-assigned. H. pexa and H . barrenense are not considered to belong to any of these groupings but their re-allocation must await further study. The protoconch of Egilomen Iredale, 1937 has bold, widely spaced ribs and vague, dis- continuous spiral creases (^pressure folds?). Most significantly however, the ribs on the protoconch show no tendency to become more crowded toward the protoconch/teleoconch boundary'. This is a characteristic of many local charopid genera that have radial elements on the protoconch and usually provides a convenient means for locating the nuclear/post-nuclear transition point. This boundary is less easily distinguished in the case of Egilomen wherein the apical rib spacing is confluent with the teleoconch rib spacing. In both E. cochlidium and Whiteheadia globosa (Stanisic, 1990) comb. nov. the spiral elements on the protoconch that are visible at low magnification consist of irregularly spaced, periostracal wrinkles that do not cross the radial elements. However, and most significantly, the latter species has much more crowded radials on the protoconch, and the protoconch- teleoconch boundary is more apparent because the denser apical ribbing contrasts clearly with the more widely spaced teleoconch. e teleoconch. This is considered to represent a generic level difference between the two species. W globosa also differs significantly from E. cochlidium in having a partially thickened lip and closed umbilicus. The shell of Egilomen (bold teleoconch ribs and open umbilicus) is most similar to that of Decoriropa gen. nov. but the latter has distinct, continuous apical spiral cords and finer, much more crowded apical ribs. Decoriropa has shell whorls of a similar profile to that of Egilomen but has the umbilicus more widely open and a weak sutural sinus. The absence of apertural barriers readily separates Egilomen from both Rhophodon and Egilodonta , which also have similarly shaped, though much smaller and more widely umbiiicate shells (cup to saucer shaped). The difficulty in dealing with this genus highlights the general problems posed by the availability of very limited material of a series of very tiny species with radially scuptured white shells (Stanisic, unpubl. data). Many are represented by only a few dry specimens, a direct consequence of relying onMitter-sorting’ for recovery of material without having located the species' true microhabitat. Such constraints will need to be overcome before a comprehensive revision of this group can be completed. Egilomen cochlidium (Cox, 1868) (Fig. 13; Table 3) Helix cochlidium Cox, 1868: 13. pi. 8, fig. 1. Egilomen cochlidium (Cox). Iredale, 1937: 138; Iredale, 1941a: 269; Smith 1992: 187; Stanisic, 1990: 103 (in part), fig. 128a-c. e-g (only ). MATERIAL. LECTOTYPE. AMSC63505, Clarence River, NSW. Height of shell= 1.25mm, diameter=2.26mm, H/D=0.55, D/U=3.14, whorl$=4 5. ribs on last whorl=30. PARALECTOTYPES. AMSC153714, 2RC, same collection data as lcctotype. OTHER MATERIAL. All NSW. AMSC57247, C8737, C152253, C153715, Cl 54732, QMM056176. DIAGNOSIS. Shell white, very small, with regularly coiled whorls, mean whorl count 4.5. Apex and early spire fiat to weakly elevated, mean H/D 0.55. Teleoconch with broad, widely spaced, weakly protractively sinuated radial ribs, 260 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 20. Reproductive system. A, Elsothera brazieri , QMMO44810. B, Decoriropa lirata , QMMO42109. C, Coricudgia wollemiana , QMM028659. D, Marilynirvpa jenolanensis , QMM037465. E, Cralopa stroudensis, QMM029722. F, Gouldiropa carlessi , QMM032080. G, Gouldiropa kaputarensis , QMM049175. Scale bai=2.5mm in A, C and G and 1 .6mm in all others. Abbreviations: dg, prostate; e, epiphallus; g, ovotestis; gd, hennaphrodite duct; gg, albumen gland; gt, talon; p, penis; prm, penis retractor muscle; ss, spcrmathecal stalk; ut, uterus; uv, free oviduct; v, vagina: vd. vas deferens. mean ribs/mm 5.57. Umbilicus moderately wide U-shaped, mean D/U 2.85. Anatomy unknown. DESCRIPTION. (Amended from Stanisic, 1990). Shell very small, white to light straw yellow, with 4.0-5. 1 (mean 4.5) evenly coiled whorls, the last descending in front. Shell diameter 1.91-3. 07mm (mean 2.20mm). Apex and early spire flat to slightly elevated. Height of shell 0.95- 1.83mm (mean 1.26mm). H/D 0.49-0.61 (mean 0.55). Protoconch flat of 1.5- 1.6 whorls. Apical sculpture of slightly curved, relatively widely spaced, broad radial ribs (approx 30), that do not become more crowded at the protoconch/teleoconch boundary; apical spiral elements vague, consisting of discontinuous NEW CHAROPID LAND SNAILS 261 periostracal wrinkles (? = pressure folds). Teleoconch with bold, broad, widely spaced, protractively sinuated radial ribs, 30-47 (mean 37) ribs on the last whorl. Ribs/mm 4.22-5.99 (mean 5.07). Interstitial sculpture of prominent crowded, low microspirals cords and relatively less prominent microradial ribs that form strong beads at their intersection. Microradial riblets low, 8-12 between each pair of major ribs. Umbilicus moderately wide U-shaped, diameter 0.60-1. 03mm (mean 0.82mm). D/U 2.59-3.47 (mean 2.85). Sutures strongly impressed . Whorls Rattened below and rounded above a laterally compressed periphery. Aperture ovately lunate, lip simple. Based on 14 measured specimens (AMSC63505, Cl 53714, C8737, C57247, C153715). REMARKS. Stanisic’s (1990) description of E. cochlidium included specimens from the Dorrigo area, NE NSW and the Border Ranges of NE NSW and SE Qld that had white shells with radially ribbed protoconchs, bold radials on the teleoconch and widely open umbilici. Although minor differences in coiling pattern, whorl profile and shell appearance were noted, these were considered variations in widespread populations. Specimen lots were small and spirit material was available in only one case. Little material has been added to this data set but in light of the emphasis on subtle differences in shell features, particularly the use of protoconch sculpture as a generic character, these specimens were subjected to closer scrutiny. Re-examination of this material (AMSC57247, Cl 28524, C 128637, C 1 286 1 8, C 1 283 1 9, C154735, QM MO 10794, M017291, MO16905, MO17290) has revealed several unrelated species (and possibly genera) on the basis of protoconch sculptural detail alone. These protoconch types are superficially similar to that of E. cochlidium in being macroscopically radial but differ in both rib architecture and spacing. In contrast to E. cochlidium these species have continuous spiral cords on the protoconch and more closely spaced and more curved apical ribs than that seen in E. cochlidium. An important consequence of the above taxonomic rearrangment is that the anatomy of£. cochlidium remains unknown. The anatomical detail presented by Stanisic (1990) refers to an unnamed species from the Dorrigo area, W of Coffs Harbour, NE NSW that will be dealt with elsewhere. Whiteheadia gen. nov. ETYMOLOGY. For Thora Whitehead. TYPE SPECIES. Egilomen globosa Stanisic, 1990; herein designated. DIAGNOSIS. Shell minute, white. Protoconch with numerous, bold, relatively closely spaced, slightly curved radial ribs (approx. 50) that do not become more crowded toward the protoconch/teleoconch boundary; secondary apical sculpture of vague, discontinuous periostracal spiral wrinkles. Teleoconch with prominent, slightly protractively sinuated, moderately widely spaced radial ribs; secondary sculpture of prominent crowded, low microspirals cords and relatively less prominent microradial ribs that form strong beads at their intersection. Umbilicus closed or reduced to a tiny chink. Kidney evenly bilobed; pericardial lobe not reflexed. Penial retractor muscle inserted on the epiphallus some distance from the penis/epiphallus junction. Penis internally with a single longitudinal pilaster and accessory circular pad. DISTRIBUTION AND HABITAT. From NE NSW to SE Qld; under logs and among rocks in closed forest. REMARKS. Characters distinguishing Whiteheadia and Egilomen have been discussed under the latter genus but two unusual characters in W globosa (Stanisic, 1 990) comb, nov require special mention. The baso-columellar thickening in W. globosa from Mt Guyra is interpreted as intraspecific variation in a disjunct population since all other characters fall within the species concept. Perhaps of greater significance though, is the epiphallic insertion of the penial retractor muscle. Most east Australian charopids hitherto dissected have the penial retractor muscle inserted on or very near to the penis/epiphallus junction. Relatively few exceptions have been noted (Stanisic, 1990) and no coherent phylogenetic pattern for this altered character state has yet emerged. Whiteheadia globosa (Stanisic, 1990). comb. nov. (Fig. 13) Egilomen globosa Stanisic, 1 990: 1 97, figs 131, 132; Smith, 1992: 187. MATERIAL. All SE Qld: QMMO 19460, M023300, M025968. TYPE LOCALITY. Sidetrack off Mt Archer Rd, Mt Mee SF, SE Qld. 262 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 21. Pallial cavity. Reproductive system. A, Elsothera brazieri , QMMO44810. B, Decoriropa lirata, QMMO42109. C, Coricuagia wollemiana, QMM028659. D. Marilyniropa jenolanensis , QMM037465. E, Cralopa stroudensis, QMM029722. F, Gouldiropa earless i, QMM032080. G, Gouldiropa kaputarensis , QMM049 175. Scale bar=2mm in C and G, 1 .6mm in D and F, and 1 .28mm in all others. Abbreviations: h, heart; hv, principal pulmonary vein: i, intestine; kdl, primary ureter; kd2, secondary ureter; kp, pericardial lobe of kidney; kr, rectal lobe of kidney; me, mantle collar; mg, mantle gland. DIAGNOSIS. Shell minute, white, subglobose to globose with elevated spire, mean H/D 0.77. Whorls regularly coiled, mean whorl count 4.63. Teleoconch with numerous, prominent, slightly protractively sinuated, moderately widely spaced radial ribs, mean ribs/mm 9.04. Umbilicus closed or reduced to a tiny chink. Lip thickened, occasionally with a baso-columellar thickening. Penis with a centrally located longitudinal pilaster and an apical, accessory circular pad. DISTRIBUTION AND HABITAT. From N of Casino (Richmond Range), NE NSW to S of Maryborough (Mt Guyra), SE Qld; under logs and among rocks in subtropical notophyll vine forest, microphyll vine forest and vine thicket. REMARKS. The combination of tiny, white, strongly ribbed, globose shell that has a predominantly radially ribbbed protoconch and closed to almost-closed umbilicus readily distinguishes W globosa from other eastern Australian charopids hitherto examined (Stanisic, unpubl. data). In general shell form, W. globosa resembles Cralopa stroudensis (Cox, 1 864) but differs in lacking distinct spirals on the protoconch, having less sinuate, less numerous NEW CHAROPID LAND SNAILS 263 TABLE 3. Shell measurements for species of Macrophallikoropa , El sot her a, Egilomen and Decoriropa. (Abbreviations as in Table 1). Measurement Macrophallikoropa belli Macrophallikoropa stenoumbilicata Macrophallikoropa depressispira Elsothera brazieri Egilomen cochlidium Decoriropa lirata N 31 (*30) 6(*5) 19 C* 1 8; **17; *♦*16) 10 (*9; **8; ***7) 4 55 (*54, **53) D 1.43-2.22(1.76) ± 0.22 1.61-2.22(1.945) ± 0.234 1.97-2.37 (2.161) ± 0.112 2.72-3.99(3.37) ± 0.37 3.89-4.6 (4.203) ± 0.311 1.83-2.63(2.19) ± 0.22 PW 1.5-1.8(1.68) ± 0.1 * 1.5-1.8(1.683) ± 0.117 1.6-1.8(1.735) ± 0.07 ** 1.5-1.8(1.657) ± 0.098 *** 1.6-1.7(1.65) ± 0.058 1.2-1.8(0.63) ± 0.09 TW 2.1-3.7 (2.78) ± 0.44 * 2.2-3 (2.65) ± 0.327 3-3.4(3.224) ± 0.13 ** 2.3-2 J (2.486) ± 0.168*** 2.4-3. 1 (2.85) ± 0.332 1. 9-3.6 (2.56) ± 0.35 NW 3. 8-5. 5 (4.46) ± 0.43 4-4.7 (4.333) ± 0.273 4. 6-5. 2 (4.959) ± 0.173** 3. 8-4.4 (4.17) ± 0.21 ** 4-4.8 (4.5) ± 0.383 3.6-5. 3 (4.2) ± 0.36 H 0.73-1.12(0.92) ± 0.11 0.77-1.19(1.003) ± 0.147 0.9-1.14(1.063) ± 0.068 1.64-1.94(1.8) ± 0.1 * 2.29-2.67 (2.42) ± 0.175 0.92-1.4(1.13) ± 0.12 ** SP 0-0.14(0.06) ± 0.04 * 0.03-0.13 (0.085) ± 0.036 - 0-0.12(0.015) ± 0.042 ** 0-0.01 (0.005) ± 0.006 0-0.2 (0.08) ± 0.04 AH/AW 1.05-1.46(1.27) ± 0.11 * 0.97-1.27(1.09) ± 0.12 1.09-1.63(1.33) ± 0.15 0.97-1.3 (0.14) ± 0.1 ** 1.17-1.4(1.29) ± 0.13 1.07-1.43(1.22) ± 0.08 UMB 0.42-0.84 (0.64) ± 0.09 0.53-0.73 (0.627) ± 0.079 0.76-0.91 (0.84) ± 0.04 ** 0.8-1.31 (0.97) ± 0.16* 1.13-1.41 (1.278) ± 0.118 0.5-0.91 (0.69) ± 0.11 * RIB 102-248(179) ± 31.48 *147-197(171) ± 20.671 165-254(218) ± 20.607 *** 136-206(162) ± 24.48 * 65-85 (73.5) ± 9.292 36-109(57.82) ± 16.72 RIBS/MM 14.69-42.19 (32.65) ± 6.19 *26.43-32.66 (28.94) ± 2.601 37.97-26.39 (32.24) ±3.172 *** 13.70-17.96 (15.32) ± 1.773 ** 4.823-6.301 (5.571) ± 0.652 5.775-11.148 (8.310) ± 1.744 H/D 0.48-0.56 (0.52) ± 0.02 0.48-0.54(0.51) ± 0.02 0.46-0.52 (0.49) ± 0.02 0.49-0.58 (0.52) ± 0.03 * 0.56-0.59 (0.58) ± 0.01 0.48-0.58 (0.52) ± 0.02 ** ■ — D/U 2.31-3.43 (2.76) ± 0.26 3.04-3.2(3.11) 2.34-2.72(2.57) ± 0.07 ± 0.11 * 3.05-4.07(3.57) ± 0.3 * 3.06-3.44 (3.3) ± 0.18 2.81-3.81 (3.19) ± 0.25 * and more widely spaced ribs on the teleoconch and in lacking a true apertural sinus. Material obtained subsequent to Stanisic (1990) has not altered either the concept of the species or details of its distribution. However, it has provided information for a more detailed analysis of the species’ preferred habitat which shows that while W globosa lives in moist forest in the south it is found in drier rainforest (microphyll vine forest, vine thicket) as its distribution attenuates northwards. And though this may initially suggest that the species has a relatively wide environmental tolerance, personal observation by one of us (JS) indicates that tV. globosa is environmentally selective at the microhabitat level in the southern part ot its distribution. For example, in the Lamington NP, SE Qld tV globosa has only been found on the Araucaria Circuit walking track, near Binna Burra which is a much drier rainforest (microphyll vine forest) than the moist notophyll vine forest dominating most parts of Lamington NP. Such local micro-environmental selectivity on the part of IV globosa may help explain the apparently patchy distribution of the species over its range. Decoriropa gen. nov. ETYMOLOGY. Latin deconis, beautiful; referring to the elegant shell. TYPE SPECIES. Helix lirata Cox, 1868; herein designated. DIAGNOSIS. Shell golden brown, with evenly coiled whorls and a weak apertural sinus. Protoconch with crowded, evenly spaced, curved radial ribs (approx. 60) and numerous regularly crowded, low, continuous spiral cords that cross the radial ribs. Teleoconch with numerous prominent, weakly to strongly protractively sinuated, moderately crowded to widely spaced radial ribs; secondary sculpture of low, thin microradial ribs that have weak periostracal blades, ribs continuous on the major ribs, crossed by generally more prominent microspiral cords that form rounded beads at their intersection with the tnicroradials. Umbilicus moderately wide to very wide U-shaped. Kidney moderately bilobed with strongly reflexed pericardial lobe. Penis with two longitudinal pilasters. DISTRIBUTION AND HABITAT. NE NSW (near Kempsey) in a broad range of habitats ranging from temperate and subtropical 264 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 22. Distribution of Els other a ^ Coricudgia , Marilyniropa , Cralopa and Gouldiropa. Key: o Elsothera brazier i, ■ Coricudgia wollemiana , □ Marilyniropa jenolanensis ▲ Cralopa stroudensis , • Gouldiropa carlessi , A Gouldiropa kaputarensis. rainforest (humid and dry) to dry vine thickets and wet and dry sclerophyll forest where it lives under logs and rocks. REMARKS. Decoriropa has a similar shell profile to Egilomen but differs in having more crowded, thinner radials on the proto- conch that are crossed by continuous spiral cords (discontinuous periostracal wrinkles in Egilomen ), and a greater number of ribs on the teleoconch. The shell of Decoriropa also bears some similarity to those of Rhophodon and Egilodonta but lacks apertural barriers and has a smaller, less open (broad V-shaped) umbilicus (mean diameter 0.69mm). Most significantly the apical shell sculpture of these two ‘dentate 'groups differs from that of Decoriropa in having crowded radial ribs that become more crowded at the protoconch /teleoconch boundary (evenly spaced in Decoriropa ); and apical spiral elements that consist of discrete cordlets that do not cross the radial ribs (continuous cords crossing the radials in Decoriropa). The single species, D. lirata (Cox, 1868) comb, nov., occurs across a range of geologies from sandstone to volcanic rocks and limestone. In many ways it shows a similar environmental adaptability to Macrophatlikoropa belli (Cox, 1 864) comb. nov. However, in contrast to the range of that species D. lirata does not appear to extend into far northern NSW or SE Qld. Decoriropa lirata (Cox, 1864) comb. nov. (Figs 10, 13-18, 20-21, 33-34; Table 3) Helix lirata Cox, 1864: 38: 1868: 13, pi. 8, fig. 1. Helix ( Patula) lirata (Cox); Cox, 1909: 44. Egilomen liranim (Cox); Ircdale, 1937: 328; 1941a: 269. Egilomen lirata (Cox); Smith, 1992: 187. MATERIAL. NEOTYPE: AMSC427984. Height of shell= 1.2mm, diameter=2 24mm, H/D=0.53, D/U=3.03, number of whorls=4.3, ribs on last whorl=70. KEY MATERIAL. All NSW. QMM029722. 10SC/28RC, Georges R. banks, Warwick Farm (33°55 ,n S, 150°56’E), degraded cw, under logs, bark and rubble, 31.xii.l989, JS, JC; QMM042 1 09, 8SC/8RC, Bungonia Gorge, ENE Goulbum (34°48’2 1 "S, 1 50W47"S), lm/ew, 1 3.x. 1 992, JS GI; AMSC356052, 3RC, Sydney, Rose Bay, just S of Queens Bell, Hermitage Foreshore Park, off Hermitage Trail (33°51.75 , S. 151°16.09 E), under litter at base of retaining wall, 07.i. 1 999, MS; AMSC 1 54792, 20+RC, S of Braid wood. Wyanbene Caves (35°48.rS, 149°40.9 , E), lm, litter from cave entrance, 1 71 1981, WFP, W.F. Ponder ( Jnr); AMSC 136842, 12RC, W ofTaree, Manning R, Wingham Brush (31°52.25’S, 152°22.85 , E), rainforest, in litter, 26.viii.1982, JS; QMM052755, RC, Kcmpscy, W, c.4km NEW CHAROPID LAND SNAILS 265 W Sherwood, Sherwood- Willi Willi Rd (31°03’24"S, 152°4r06 M E), drf/lm, 40m, litter, 28.xii.1993, JS, JC; QMM064764, 20RC, Wombeyan Caves, NNE Goulbum (34°18’35"S, 149°57’46"E), hn/vt, litter, 13x1992, JS, GI; AMSC 154791, 3RC, S of Forster, E of Smiths Lake, Bald Head (32.382°S, 1 52.53 1°E), under low scrub on dunes, in litter, x, 1981 , OG; AMSC157311, 13RC. W of Taree, Wingham, Wingham Brush (31.871 1> S, 15238l°E), leal' litter i in subtropical lowland floodplain rf \ 05.ii.1982, *MS. OTHER MATERIAL. Northeastern NSW: QMMO 12745, MO44S30 MO 16758, M016763, M037514. M017274, M( >31909, M031935, MO32066, M037733, MO32072, M049478, M056147, MO52701, M056136, M056142, M059891, M032291, MO37106, M049395, M055875, M055893. MO56012, M056050, M056080, M059794, MO60143. MO60154, MO60477, MO60491, MO60498, AMSC 168693, C339712, C411598. Sydney Basin: QMMO29640, M034783, M034973. M037389, M017818, M042136, M044783, AMSC63774, Cl 40472, C3 19043. C3 19058. C3 19059, C3 19080, C3 19087, C3 19088, C319096, C319098, C319118, C319120, C3 19125, C3 19126, C345665, C356051. C3S6080. Southeastern NSW: QMM029271. M037791, M037812, MO37980, M042I24, M068010, AMSC3 1 8792, C63729, C157294, C168663, C157316, Cl 68673, Cl 68676, C358326, C358328, C358330, C346061. DIAGNOSIS. Shell very small, golden brown with evenly coiled whorls and a weak apertural sinus. Spire and early apex Hat to slightly elevated. Teleoconch with numerous prominent, weakly to strongly protractively sinuated, moderately crowded to widely spaced radial ribs (mean ribs/mm 8.31. Umbilicus moderately wide to very wide V-shaped, diameter range 0.50-0.9 1mm, mean D/U 3.19. Penis with two longitudinal pilasters. DESCRIPTION. Shell very small, golden brown, with 3. 6-5. 3 (mean 4.2) evenly coiled whorls, the last dccending in front; weak apertural sinus present. Shell diameter 1.83-2. 63mm (mean 2.19mm). Apex and early spire flat to slightly elevated. Height of shell 0.92-1.40mm (mean 1.13mm), H/D 0.48-0.58 (mean 0.52). Protoconch of 1.2- 1.8 whorls with sculpture of regularly spaced, moderately crowded, curved radial ribs, width of interstices (in multiples of rib width) 2-4 at apex. Secondary sculpture of low, continuous spiral cords (23-24), crossing radial ribs. Teleoconch sculpture of crowded, regularly spaced (increasing near end of last whorl), weakly protractively sinuated radial ribs; 36-109 (mean 58) ribs on the last whorl. Ribs/mm 5.78-11.15 (mean 8.31). Ribs wide, height less than or equal to width, straight in section, rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 4-6 ribs; interstices on the penultimate whorl equal to width of 3-6 ribs. Interstitial sculpture of weak microradial ribs, 4-10 between ribs on first post-nuclear whorl; 7-13 between ribs on penultimate whorl; continuous on major radials, with weak periostracal blades. Secondary microsculpture of more prominent low, microspiral cords that cross the major ribs and form strong tranversely elongate to rounded beads at their intersections with the nicroradials. Umbilcus very wide V-shaped, diameter 0.50-0.91mm (mean 0.69mm), D/U 2.81-3.81 (mean 3.19). Whorls flattened above and rounded below a weakly rounded periphery, sutures impressed. Aperture ovately lunate ovate, parietal callus present. Based on 55 measured specimens (AMSC356052, Cl 54792, Cl 36842, SC427984, QMM052755, M064764, ). Genitalia with ovotestis containing two clumps of alveoli, with two or more alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penial retractor muscle less than half the length of the penis; inserting at the junction of the penis and epiphallus. Epiphallus longer than penis, narrower than the vas deferens, entering penis through a simple pore; internally with longitudinal pilasters. Penis tubular with sheath, internally with two longitudinal pilasters. Vagina shorter or equal in length to penis. Pallial cavity with moderately bilobed kidney, pericardial lobe strongly reflexed at apex. Radula with central tooth strongly tricuspid; lateral teeth tricuspid; marginal teeth skewed towards the centre; ectocone of outer marginal teeth split into three teeth; endocone not split. Number of lateral teeth: 4; marginal teeth: 10; radular rows: 73. Based on 3 dissected specimens (QMM029722, MO42109). DISTRIBUTION AND HABITAT. Decoriropa lirata is widely distributed ranging from the Wyanbene Caves, SE NSW to the Macleay River valley, NE NSW. The species lives in a variety of forest-types ranging from humid subtropical rainforest to dry rainforest (vine thickets) and littoral rainforest, as well as dry and wet sclerophyll forest. The widespread nature of the distribution and habitat preference suggests an inherent adaptability that is probably a contributing factor to the species’ continued survival in suburban Sydney. REMARKS. Decoriropa lirata looks superficially like Marilyniropa jenolanensis sp. nov. but can be distinguished by its smaller shell that has narrower whorls (smaller cross section), a weak 266 MEMOIRS OF THE QUEENSLAND MUSEUM apertural sinus, more dense ribbing and usually, a less elevated spire. D. lirata does not appear to occur on the Jenolan or Tuglow limestones but is found on the nearby Church Creek and Colong Caves limestones where it is sympatric with M. jenolan - ensis. Juveniles of the latter species are easily confused with D. lirata but are readily separated by having fewer (approx. 45), less crowded radials on the protoconch. D. lirata is also apparently absent from the nearby, Abercrombie limestones but is present on the more distant Wombeyan limestones. Specimens from Wombeyan Caves have a lower rib count, slightly higher spire and narrower umbilicus than usual, but otherwise agree with the general concept of the species which is remarkably stable across a wide and variable landscape. In the absence of type material (Smith, 1992) a neotype was designated. The specimen chosen comes from inner Sydney close to the original type locality (Ruschcutters Bay). A key reason for choosing the Bellevue Hill locality was that the species is still readily found there alive. Coricudgia gen. nov. ETYMOLOGY. For Mt Coricudgy, NSW. TYPE SPECIES. Coricudgia wollemiana sp. nov.; herein designated. DIAGNOSIS. Shell white with a relatively large number of tightly coiled whorls and moderately elevated spire. Protoconch with very crowded (approx. 90) curved radial ribs and continuous crowded microspiral cords. Teleoconch with weakly sinuated, widely spaced, bold radial ribs; secondary sculpture of prominent microradial ribs with weaker microspiral cords that form rounded beads at their intersection. Umbilicus open U-shapcd. Kidney evenly bilobed with large wedge-shaped lobes; pericardial lobe only weakly reflected. Penis with 2 longitudinal pilasters. DISTRIBUTION AND HABITAT. Restricted range centred on Mt Coricudgy, Wollemi NP, NSW. Mostly found living among rocks in rainforest. REMARKS. Coricudgia gen. nov. displays several shell characters that distinguish it from other E Australian charopids with bold ribs and an open umbilicus. The high whorl count, large number of radial ribs and continuous spiral cords on the protoconch separate it from Egilomen (lower whorl number, few apical radials with NEW CHAROPID LAND SNAILS 267 vague spiral wrinkles) which has a similarly coiled, whitish shell with moderately wide umbilicus and moderately elevated spire. It is distinguished from Decoriropa chiefly by the more elevated spire, the greater number of straighter ribs on the protoconch, a greater number of tightly coiled whorls and the much less sinuate ribs on the teleoconch. The greater number of tightly coiled whorls distinguish it from Cralopa Iredale, 1941, Gouldiropa gen. nov., Sharniropa gen. nov. and Marilyn iropa gen. nov.which all have comparatively fewer whorls, bolder ribs and relatively looser coiling. Coricudgia wollemiana sp. nov. (Figs 10, 14-17, 19, 33-34; Table 4) ETYMOLOGY. For Wollemi NP, the type locality. MATERIAL. HOLOTYPE: AMSC205164, E of Rylstone, Mount Coricudgy, southern slope, NSW (32°50.82’S, 150°2rE), in litter off road, 2.x. 1983, PHC. Height of shells 1.95mm, diameter=3.73mm, H/D=0.52, D/U=3.59, number of whorls=6. 1 , ribs on last whorl=46. PARATYPES: All NSW AMSC162143, 18RC. same data as holotypc; QMM028659, 1 3SC/9RC, slopes of Mt Coricudgy, W of Rylstone (32°5LS, 150°2rE), under rocks, 15. i. 1990, JS, JC; QMMO44708, IRC, Mt Coricudgy, NE Rylstone, summit (32°49 , 48 M S, 150°2r24”E), remnant rainforest, under rocks, lO.i.1993, JS, JC; QMM059868. 4SC/7RC Rylstone, E, at Mt Coricudgy (32°5P25"S, 150°2r00”E), wtrf/basalt, under logs and rocks, 0 Li. 1996, JS, JC; QMM044725, 1RC/11SC, Mt Coricudgy, NE Rylstone, c.O.lkm E summit road. (32°51°00"S, 150°21T2"E), ws/wtrf, on roots of plants under rocks, 10.i.l993, JS, JC. OTHER MATERIAL. QMM059859. DIAGNOSIS. Shell very small, white, with moderately elevated spire (mean spire height 0.26mm) and a relatively large number of tightly coiled whorls (mean whorl count 4.9, range to 6. 1 whorls). Protoconch with crowded (approx. 90) curved radial ribs and continuous crowded microspiral cords. Teleoconch with weakly sinuated, widely spaced, bold radial ribs, mean rib count 4.1 ribs/mm. Umbilicus open U-shaped, mean D/U 3.49. Penis with two longitudinal pilasters. DESCRIPTION. Shell very small, white, with 3.6- 6. 1 (mean 4.9) tightly coiled whorls, the last descending more rapidly in front. Whorls weakly rounded above and rounded below a rounded periphery. Shell diameter 1.62-4. 02mm (mean 3.30mm). Apex and early spire weakly elevated. Height of shell 0.83-2.2 1mm (mean 1.69mm), H/D 0.45-0.55 (mean 0.51). Protoconch of 1.7- 1.8 whorls with sculpture of crowded. regularly spaced, slightly curved radial ribs (approx. 90); width of interstices (in multiples of rib width) 3 at apex. Secondary sculpture of continuous spiral cords that cross the radials. Teleoconch with widely but regularly spaced, weakly protractively sinuated, bold radial ribs. Ribs on last whorl 28-47 (mean 40), ribs/mm 3.01-5.50 (mean 4.09). Height of ribs equal to width; straight in section; rounded on top, continuous on major radials. Rib interstices on the first post-nuclear whorl equal to width of 6-7 ribs. Interstitial sculpture of high microradial ribs, 9 between major ribs on first post-nuclear whorl; 16 between ribs on penultimate whorl. Secondary sculpture of low microspiral cords forming strong rounded beads at their intersection with the microradials. Umbilicus open, U-shaped, diameter 0.55- 1.1 5mm (mean 0.94mm), D/U2.95-3.80 (mean 3.49). Aperture broadly ovately lunate, parietal callus present. Based on 10 measured specimens (AMS Cl 62 143, C205164). Genitalia with ovotestis containing two clumps of alveoli, with more than two alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penial retractor muscle less than half the length of the penis; inserting onto the penis head; entering penis through a simple pore. Penis tubular; with two longitudinal pilasters present, one very large and irregular in shape’ the other much smaller and regular in shape). Vagina shorter than penis. Pallial cavity with strong, evenly bilobed kidney; apex of pericardial lobe weakly reflexed. Mantle gland present. Radula with central tooth strongly tricuspid; lateral teeth tricuspid; first and second laterals equal in width. Mesocone of first lateral tooth slender, slightly diamond-shaped; marginal teeth skewed towards the centre. Ectocone of outer marginal teeth not split; endocone not split; number of lateral teeth: 7; marginal teeth: 11; radular rows: 100. Based on 2 dissected specimens (QMM028659). DISTRIBUTION AND HABITAT. Known only from the Rylstone area, NSW but particularly the upper parts of Mt Coricudgy, Wollemi NP, NSW where the species lives among volcanic rocks in warm temperate rainforest. REMARKS. Coricudgia wollemiana sp. nov. is easily recognisable by its white shell with very high whorl count, tightly coiled whorls and moderately elevated spire. Some specimens attain greater than 6 whorls which is a 268 MEMOIRS OF THE QUEENSLAND MUSEUM comparatively high whorl count in the context of moderately sized (i.e. approx, shell diameter 3. 5-4. 5mm) eastern Australian charopids. However some other, smaller charopids, including some dealt with in this study, e.g. Rhophodon spp. and Letomola spp., have higher whorl counts. The discovery of this species on the heights of Mt Coricudgy once again points to the unusual nature of this rainforest/volcanic refugium among an otherwise dry sclerophyll/sandstone land- scape. Mt Coricudgy is also home to the geographically restricted Macrophallikoropa stenoumbilicata sp nov. suggesting that the locality has broad significance as an historical refugium for mesic communities. A single specimen of C. wollemiana has also been found at a lowland locality nearCapertee but the viability of any lowland populations needs to be more accurately assessed by undertaking additional collecting in the area. It is quite possible that the Capertee specimen was stream drift material washed from the mountain slopes. Relatively little land snail collecting has been under- taken in the Wollemi NP and the full extent of the distribution of C. wollemiana has yet to be determined. Marilyniropa gen. nov. ETYMOLOGY. For Marilyn Joy McGrady. DIAGNOSIS. Shell very small, straw to beige in colour, with evenly coiled whorls; apertural sinus absent. Protoconch with evenly spaced, moderately crowded, curved radial ribs (approx. 55) that are crossed by crowded, low continuous spiral cords; radial ribs becoming bolder toward the protoconch/tel coconch boundary. Teleoconch with bold, widely spaced, strongly sinuate ribs (mean ribs/mm 4.62) that become more crowded on the latter half of the last whorl. Umbilicus wide V-shaped. Kidney moderately bilobed, pericardial lobe reflexed. Mantle gland present. Penis with five longitudinal pilasters. DISTRIBUTION AND HABITAT. Known only from limestone outcrops SSE Oberon, NSW. The single species, Marilyniropa jenolanensis sp. nov., lives among limestone talus. REMARKS. Marilyniropa gen. nov. is grossly similar to Decoriropa in having an open umbilicus and bold ribbing but differs in having fewer, apical radial ribs, an elevated spire, no apertural sinus and fewer, variably spaced, more curved radial ribs on the teleoconch. Anatomically, Marilyniropa differs by having greater number of penial pilasters and shorter epiphallus than Decoriropa. Unfortunately it is not possible to NEW CHAROPID LAND SNAILS 269 put the significance of these character shifts in reproductive anatomy into phyletic context at this time except to say that they do represent major departures from the patterns seen in the somewhat conchologically similar species belonging to Decoriropa, Sharniropa, Gouldiropa, Acheronopa and Cralopa. The combination of more tightly coiled whorls and larger umbilicus differentiates Marilyniropa from the sympatric Sharniropa and Gouldiropa. The current restriction of Marilyniropa to a few habitats surrounding and including Jenolan Caves, once again highlights the biogeographical importance of these limestones to land snail evolution. Marilyniropa jcnolanensis sp. nov. (Figs 10, 14-17, 19, 33-34; Table 4) ETYMOLOGY. For the type locality. MATERIAL. HOLOTYPE: AMSC205165, Jenolan Caves, NSW (33°49.3’S, 150°1.2’E), 6.vi.l982. O.L. Griffiths. Height of shell- 1.77mm, diameter-3 .46mm, H/D-0.51, D/U=3.33, number of whorls=4.5. ribs on last whorl-42. PARATYPES: All NSW. AMSC 144224, many RC, same data as holotvpe; AMSC 1 42966, Jenolan Caves, ridge on top of Grand Arch (33°49.24’S, 150°1.32’E), 12.xii.1979, JS. QMMO9730, 10 IRC, Jenolan Caves, above Grand Arch, lm, litter among rocks, 1 1 .xii. 1 979, JS; QMM028777. 2SC/4RC, Jenolan Caves, lm, under rocks, 9.i. 1990, JS, JC; QMM037776. 7RC, Jenolan Caves (33°49'S, 150°0rE), lm, in litter, 9.U990, JS. JC; QMM037465, 4SC/36RC, Jenolan Caves (33°49*S, 150°0rE), lm, on limestone rocks among dirt, 10.i.l990, JS, JC. OTHER MATERIAL. All NSW. QMMO70485, MO70486, MO62450. AMSC63736, C63852, C121716, Cl 24272, Cl 54773, Cl 54774, Cl 54775, Cl 54776, Cl 54777, Cl 57333, Cl 68669, C346748, C346750. DIAGNOSIS. Shell light brown to beige in colour with evenly coiled whorls, mean whorl count 4.5 1 . Apex flat, early spire elevated, mean spire height 0.25mm, mean H/D 0.48. Proto- conch with evenly spaced, moderately crowded, curved radial ribs (approx. 55) that are crossed by crowded, low continuous spiral cords. Teleo- conch with bold, moderately widely spaced, strongly sinuate ribs (mean ribs/mm 4.62) that become more crowded on the latter half of the last whorl. Umbilicus wide V-shaped, mean diameter 0.89mm, mean D/U 3.67. Penis with 5 short, longitudinal pilasters. DESCRIPTION. Shell very small, straw to light golden brown in colour with 4. 1-4.8 (mean 4.5) evenly coiled whorls, the last descending in front; apertural sinus not developed. Shell diameter 2. 96-3. 65mm (mean 3.24mm). Apex flat, early spire weakly elevated. Height of shell 1.33-1. 97mm (mean 1.55mm), H/D 0.43-0.55 (mean 0.48). Protoconch with 1.4- 1.7 whorls. Apical sculpture of crowded regularly spaced, slightly curved radial ribs (approx. 45); width of interstices (in multiples of rib width) 4-6 at apex. Secondary sculpture of narrow, continuous spiral cords that cross the microradial ribs; ribs becoming bolder toward the protoconch/teleo- conch boundary. Teleoconch with prominent, regularly spaced, strongly protractively sinuated radial ribs, 35-6 1 (mean 47) ribs on the last whorl. Ribs/mm 3.76-6.08 (mean 4.62). Ribs greater in height than width; straight in section; rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 4-7 ribs. Secondary micro- sculpture of more prominent low, microspiral cords that cross the major ribs and form strong tranversely elongate to rounded beads at their intersections with the nicroradials. Whorls flattened above and rounded below a rounded periphery, sutures impressed. Umbilicus open, wide V-shaped. Umbilical width 0.73- 1.07mm (mean 0.89mm), D/U 3.33-4.05 (mean 3.67). Based on 23 measured specimens (AMSC 1 44244, C205165). Genitalia with ovotestis containing two clumps of alveoli, with more than two alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penial retractor muscle less than half the length of the penis, inserting at the junction of the penis and cpiphalius; epiphallus short. Penis tubular with 5 short, longitudinal pilasters. Vagina shorter than penis. Pallial cavity with moderately bilobed kidney, pericardial lobe reflexed. Mantle gland present. Radula with central tooth strongly tricuspid; lateral teeth tricuspid with first lateral slightly wider than second lateral. Mesocone of first lateral tooth slender, slightly diamond-shaped; marginal teeth skewed towards the centre. Ectocone of outer marginal teeth not split; endocone not split. Number of lateral teeth: 4; marginal teeth: 18; radular rows: 100. Based on 1 dissected specimen (QMM037465). DISTRIBUTION AND HABITAT. Known only from the Jenolan Caves region (Jenolan, Tuglow and Colong limestones), Blue Mountains, W of Sydney; living among limestone talus. REMARKS. Marilyniropa jenolanensis sp. nov. most closely resembles Decoriropa lirata (Cox, 1864) comb. nov. in general appearance but can be distinguished from that species by its larger size, more elevated spire and fewer, more widely 270 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 25. Dorsal view of shells. A, Cralopa stroudensis (SEQ), QMMO 16857. B, Cralopa stroudensis (Sydney), QMM029638. C, Gouldiropa carles si, AMSC003640. D, Shamiropa wollondillyana , QMMO70399. E, Shamiropa borenorensis , AMSC205166. F, Shamiropa xanana , QMMO70398. G, Acheronopa attunga , QMM070400. H, Hedleyropa yarrangobillyensis, QMM070401. 1, Scleridompa sarahjaneae (Yarrangobilly), QMM070402. J, Scleridompa nandewar ; QMM070403. K, Diphyoropa saturni , AMSC205 1 67. L, Diphyoropa macleayana , AMSC 168685. Scale bars = 0.5mm in J; 0.8mm in A-H, L; 1 mm in K. NEW CHAROPID LAND SNAILS 271 spaced, bolder apical radial ribs and fewer, less crowded ribs on the teleoconch. M. jenolanensis broadly resembles Sharniropa wollondillyana sp. nov. in general appearance of the shell (shape, sculpture) but the latter is larger, has more loosely coiled whorls, smaller umbilicus and bolder, more widely spaced ribs on the teleoconch. Cralopa Iredale, 1941 Cralopa Iredale. 1941a: 267: Kershaw, 1956b: 8: Burch. 1976b: 132; Stanisic, 1990: 147 (in part); Smith, 1992: 184 (in part). TYPE SPECIES. Helix stroudensis Cox, 1 864; by original designation. DIAGNOSIS. Shell very small with moderately loosely coiled whorls, deep sutures and a well-developed apertural sinus. Protoconch with very crowded, relatvely narrow, curved radial ribs (approx. 80) crossed by numerous regularly spaced, crowded, low, continuous spiral cords. Teleoconch with numerous, prominent and very strongly protractively sinuated, crowded radial ribs that have weak periostracal blades. Second- ary sculpture of prominent microspiral cords that cross the microradials and form round beads at their intersection. Umbilicus closed or only very slightly open. Kidney moderately to strongly bilobed with strongly reflexed pericardial lobe. Penis wih two longitudinal pilasters. DISTRIBUTION AND HABITAT. Cralopa Iredale, 1941 ranges from southern NSW to SE Qld. The single species lives under logs and rocks in moist rainforest and wet sclerophyll forest. REMARKS. Cralopa was introduced by Iredale (1941 a) for Helix stroudensis Cox, 1 864. Iredale (1941b) added an additional species, Cralopa intensa Iredale, 1941. Stanisic (1990) revised Cralopa and in doing so reassigned C. intensa to Sinployea Solem, 1983 and included two additional new species (C. kaputarensis , C. carlessi) on the basis of shared shell and anatomical characters (closed to almost closed umbilicus, presence of an apertural sinus and large cylindrical penis). Analysis of protoconch sculpture consisted of noting the common presence of prominent radial ribs and low spiral cords. A difference in apical whorl size was noted between both C. carlessi and C. kaputarensis and C. stroudensis , the latter two species having a smaller protoconch than the former species. While conceding the cohesiveness of his new concept of Cralopa , Stanisic ( 1 990) did however, make a number of observations relevant to a possible future revision of the genus once additional material became available for study. Notable among these was firstly, that the biogeography of the genus was at odds (east-west) with the general pattern displayed by most other genera reviewed (north-south); secondly, that some individuals of C. stroudensis from the limestones of the Macleay Valley, W of Kempsey were unusual for their large size in what was an otherwise conchologically conservative, widespread species; and thirdly, that there was a large degree of variability displayed by C. carlessi in material from widespread localities (Wee Jasper area, SE NSW to Inverell, NE NSW). At the time this variability in shell features was considered to be local differences in a species with unusually wide- spread, disjunct populations. A number of these populations are herein excluded from the definition of C. carlessi (see below). Cralopa Iredale, 1941 is redefined on the basis protoconch sculpture. In contrast to both C. carlessi and C. kaputarensis and C. stroudensis have comparatively smaller protoconchs with more numerous and more crowded apical radial ribs (Stanisic, 1990). Cralopa is herein restricted to include only the type and C. kaputarensis (Nandewar Range Bioregion). C. carlessi (herein restricted to localities in the Attunga-Inverell area of the New England Bioregion, NSW) is reassigned to Gouldiropa gen. nov. Specimens of C. stroudensis from the limestone outcrops of the Macleay Valley, W of Kempsey cited in Stanisic (1990), viz. AMSC121680 [Yessabah], AMSC152198 [Natural Arch], are excluded from the revised definition of C. stroudensis and will be considered elsewhere (Stanisic, in prep.). Cralopa Iredale, 1941 as redefined herein differs from that of Stanisic (1990) by the removal of C. carlessi Stanisic, 1990. Although the shell of this species bears a number of similarities to that of C. stroudensis these must be considered examples of convergent or parallel evolution. The smaller protoconch and denser apical ribbing of C. stroudensis and C. kaputarensis are considered key characters that serve to generically separate these species from C. carlessi. ‘ Charopa 4 colliveri Gabriel, 1947 from SE Victoria was included here by Smith & Kershaw (1979) without detailed reasons but presumably on the basis of the reduced umbilicus and bold radial ribs on the teleoconch. A casual inspection of a co-type of ‘C. ' colliveri in the QM (M035761) from the Stan Colliver Collection reveals that there are characters such as the 272 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 26. Protoconch of shells from above. A, Cralopa stroudensis (SEQ), QMMOl 6857. B, Cralopa stroudensis (Sydney), QMM029638. C, Gouldiropa carlessi . AMSC003640. D, Sharniropa wollondillyana , QMMO70399. E, Sharniropa borenorensis, AMSC205166. F, Sharniropa xanana , QMMO70398. G, Acheronopa attunga , QMM070400. H, Hedleyropa yarrangobillyensis, QMM070401. I, Scleridoropa sarahjaneae ( Yarrangob illy). QMM070402. J, Scleridoropa nandewar , ; QMM070403. K, Diphyoropa saturni , AMSC205I67. L, Diphyoropa macleayana , AMSC 168685. Scale bars=0.15mm in K, L; 0.2mm in A-J. NEW CHAROPID LAND SNAILS 273 considerably fewer number of radial ribs on the protoconch, the absence of an apertural sinus and less sinuated radial ribs that suggest this species is not congeneric with C. stroudensis. ‘C. ' colliveri also differs from C. stroudensis in the nature of the umbilical closure which in the latter is partially due to relection of the columellar margin of the aperture over the umbilical opening rather than constriction as is the case in ‘CV colliveri. All these differences indicate generic incompatability, however, the generic placement of this species must await a future review that will probably need to include other Victorian and possibly Tasmanian charopids. Cralopa stroudensis (Cox, 1864) (Figs 10, 20-22, 25-29, 34; Table 4) Helix stroudensis Cox, 1864: 20; Tryon, 1887: 25, pi. 4, figs 28-29. Gyrocochlea stroudensis (Cox). I ledley, 1 924: 2 1 7, pi. 3, figs 25-27; Ircdale, 1937: 323. Cralopa stroudensis (Cox). Ircdale, 1941a: 269; Kershaw, 1956b: 8; Stanisic, 1990: 150, figs 95-97 (part); Smith, 1992: 184. MATERIAL. HOLOTYPE: AMSC63500, E of Dungog. Stroud, NSW: (32°24.5'S, 151°58’E), Rev. R.L.King. Height of shell- 1.62mm, diameter=2.56mm, H/D=0.63, D/U=8.30, ribs on last whorl =109, whorls=4. PARATYPES. AMSC225, 4, same data as holotype. OTHER MATERIAL. NSW: QMM029638, MO 12281. MO 16857. M 049237, MO42140, MO20294. M031855, MO32109, MO36809, M037544, MO56140, MO38770, M049239, M055877, M055931. SE Qld: QMM029796, M045I73, M028453. DIAGNOSIS. Shell light golden brown, with moderately loosely coiled whorls, a well developed apertural sinus and strongly impressed sutures. Apex and early spire weakly elevated. Teleoconch with numerous, prominent and very strongly protractively sinuated, crowded radial ribs; mean ribs/mm = 10.08. Umbilicus closed or only very slightly open, mean D/U 7.81. Penis with two longitudinal pilasters. DESCRIPTION. Shell very small, light golden brown, with 2. 8-4.3 (mean 4.0) moderately loosely coiled whorls. Whorls weakly rounded to shouldered above and rounded below a weakly rounded periphery, well developed apertural sinus present. Shell diameter 1.63- 1.33mm (mean 2.58mm). Apex and early spire very weakly elevated. Height of shell 1.02-1. 73mm (mean 1 .53mm), H/'D 0.53-0.66 (mean 0.66mm). Protoconch of 1 .5- 1 .7 whorls with very crowded, regularly spaced, curved radial ribs (approx. 80); ribs of medium width; height less than or greater than width; spacing uniform; width of interstices (in multiples of rib width) 3-5 at apex. Secondary sculpture of low, crowded microspiral cords that cross the microradials. Teleoconch with prominent, crowded, strongly protractively sinuated, bold radial ribs. Ribs on the last whorl 68-126 (mean 88); ribs/mm 7.90-16.50 (mean 10.83); rib height less than to greater than width; straight in section; rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 3-5 ribs; on the penultimate whorl equal to width of 4-5 ribs. Interstitial sculpture of weak, low to very low microradial ribs; 7-8 between ribs on first post-nuclear whorl; 6-13 between ribs on penultimate whorl. Microradial ribs with weak periostracal blades. Secondary microsculpture of prominent spiral cords that cross the microradials and form round beads at their intersection. Sutures strongly impressed. Umbilicus reduced to a pinhole and partially occluded by a reflection of the columellar edge of the aperture; occasionally closed. Umbilical width when open 0.14-0.44mm (mean 0.34mm), D/U 5.68-11-64 (mean 7.81). Aperture ovately lunate; parietal callus present. Based on 15 measured specimens (QMM029638, M016857, M049237). Genitalia with ovotestis containing two clumps of alveoli, with more than two alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penial retractor muscle less than half the length of the penis; insering onto the penis head, or at the junction of the penis and epiphallus; enters penis through a simple pore. Penis tubular; two longitudinal; pilasters present. Pallial cavity with kidney moderately bilobed; apex of pericardial lobe reflexed. Mantle gland present. Radula with central tooth strongly tricuspid; lateral teeth tricuspid; first and second laterals equal in width, or first lateral wider than second lateral. Mesocone of first lateral tooth triangular, or slender, slightly diamond-shaped; marginal teeth skewed towards the centre. Ectocone of outer marginal teeth not split; endocone not split; number of lateral teeth; 5-6; marginal teeth: 13-14; radular rows: 88-91. Based on 6 dissected specimens (QMMO 12281, M029638). DISTRIBUTION AND HABITAT. Sydney area, west to the Blue Mountains, central NSW and N to the Bunya Mts, SE Qld; living under logs and rocks in subtropical and warm temperate rainforest. 274 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 27. Lateral view of shells. A, Cralopa strvudensis (SEQ), QMMO 1 6857. B, Cralopa stroudensis (Sydney), QMM029638. C, Gouldiropa carles si, AMSC003640. D, Sharniropa wollondillyana , QMMO70399. E, Shamiropa borenorensis , AMSC205166. F, Sharniropa xanana, QMMO70398. G, Acheronopa attunga , QMM070400. H, Hedleyropa yarrangobillyensis , QMM070401. I, Scleridoropa sarahjaneae (Yarrangobilly), QMM070402. J, Scleridoropa nandewar ; QMM070403. K, Diphyoropa saturni , AMSC205167. L, Diphyoropa macleayana , AMSC205274. Scale bars=lmm in A-I, K, L; o.5mm in J. NEW CHAROPID LAND SNAILS 275 REMARKS. The definition of Cralopa stroudensis presented herein differs from that of Stanisic ( 1 990) by the exclusion of material from the limestones of the Macleay Valley, W of Kempsey. NSW (AMSC121680, C128902, Cl 52 198). The Macleay Valley specimens have much less crowded radials on the protoconch (possibly affiliated with Gouldiropa ; Stanisic, pers. obs.) but are not dealt with in this study. C. stroudensis does, however, occur in the Kempsey region where it has been found in subtropical rainforest and wet sclerophyll forest adjacent to, but not on limestone outcrops. Cralopa stroudensis shows some differences in shell features across its range (teleoconch rib spacing, shell size) but these variations display little geographic consistency and are also within the limits of observed intrapopulation variability. Most significantly this conchological variation is not reflected in any detectable change in reproductive anatomy. Shells of C stroudensis can be distinguished from those of some other east Australian eharopids with a closed to almost closed umbilicus eg. Elsothera sericatula (Pfeiffer, 1849)[Sydney region], E. nautilodea (Cox, 1866) [Grafton area, NE NSW], Whiteheadia globosa (Stanisic, 1 990) comb. nov. [NSW/Qld Border Ranges] and Gouldiropa gen. nov. by a combination of the following features: numerous, very crowded radial ribs on the protoconch (fewer and more widely spaced in W globosa and Gouldiropa); very crowded, strongly sinuated radial ribs on the teleoconch (more crowded and very weakly sinuated in Elsothera. widely spaced and almost straight in W. globosa ); and a well-developed apertural sinus (absent in Elsothera , W. globosa and Gouldiropa). Cralopa kaputarensis Stanisic, 1990 (Figs 10, 20-22, 25-29) Cralopa kaputarensis Stanisic. 1990: 153: 98; Smith 1992: 184. MATERIAL. HOLOTYPE. QMMO 16753, Mt Kaputar, summit, NSW (30°16’S, 1 50° 1 O' E), among grass and litter, 9.xi. 1 983, JS, DP, PHC. J. Short. Height ofshell=0.90mm, diameter=1.66, H/D=0.54, whorls=4.0, ribs on last whorl=80. PARATYPES. All NSW. QMMO 16742, 1SC/2RC, Dawson Springs, Mt Kaputar (30 ,, 16 , S, 1 50° 1 O’ E), under logs, in fem gully alt. 1 300m, 8.xi. 1 983, JS, DP, PHC. J. Short; QMMO 1 6750. 1 RC, Jokers Spring, Mt Kaputar (30“16'S, 150°10’E), edges of rocky creek bed, among lichens, alt. 1 100m, 8.ix. 1983, JS. DP, PHC, J. Short. OTHER MATERIAL. QMM049167, M049175. DIAGNOSIS. Shell very small, dark brown with moderately loosely coiled whorls and a barely visible apertural sinus; whorls flattened above a rounded periphery. Spire and early apex barely elevated. Teleoconch with numerous, crowded, prominent strongly protractively sinuated radial ribs (mean ribs/mm 13.94). Umbilicus reduced to a slight chink. Penis with two longitudinal and bifurcated pilasters present. DISTRIBUTION AND HABITAT. From Mt Kaputar, Nandewar Ranges, NSW; in moist woodland living under rocks, logs and among lichen near springs. REMARKS. Cralopa kaputarensis differs from G. carlessi by its smaller size, in having a more closed umbilicus and a greater number of radial ribs on theprotoconch and teleoconch. C. kaputarensis is still only known from summit area of Mt Kaputar. However, this could be an artefact of collecting effort since most areas in and around these ranges still need to be investigated. Additional specimens collected since 1990 have not altered the species profile of Stanisic (1990). Gouldiropa gen. nov. ETYMOLOGY. In honour of the late Stephen Jay Gould. TYPE SPECIES. Cralopa carlessi Stanisic, 1990; herein designated. DIAGNOSIS. Shell very small, dark brown, with moderately loosely coiled whorls and strongly impressed sutures; apertural sinus absent or only weakly developed. Protoconch with relatively widely spaced, crowded curved radial ribs (approx. 40) and numerous, regularly crowded, low continuous spiral cords that cross the radial ribs. Teleoconch with numerous, bold, weakly to very strongly protractively sinuated, moderately crowded to very widely spaced radial ribs. Interstitial sculpture with microspiral cords stronger than microradial ribs, beading at their intersection barely noticeable, obscured by strength of microspiral cords. Umbilicus closed or only slightly open. Columellar edge of aperture weakly reflected across the umbilical opening. Kidney moderately to strongly bilobed with reflexed apex on pericardial lobe. Penis with two longitudinal pilasters that may be divided. DISTRIBUTION AND HABITAT. New England Bioregion. Species occur on limestone outcrops associated with vine thicket and in settered moist refugia on volcanic rocks. The species live under rocks among vine thicket and in wet sclerophyll forest. 276 MEMOIRS OF THE QUEENSLAND MUSEUM J K L FIG 28. Ventral view of shells. A, Cralopa stroudensis (SEQ), QMMO 16857. B, Cralopa stroudensis (Sydney), QMM029638. C, Gouldiropa earless i, AMSC003640. D, Sharniropa wollondillyana , QMMO70399. E, Sharnirbpa bo re no re ns is, AMSC205166. F, Sharniropa xanana , QMMO70398. G Acheronopa attunga , QMM070400. H, Hedleyropa yarrangobillyensis, QMM070401 . 1, Scleridoropa sarahjaneae (Yarrangobilly), QMM070402. J, Scleridompa nandewan QMM070403. K. Diphyoropa saturni , AMSC205 1 67. L. Diphyoropa macleayana , AMSC 168685. Scale bars=0.5mm in J; 0.8mm in D; 1mm in A-C, E-I, K, L. NEW CHAROPID LAND SNAILS 277 REMARKS. Gouldiropa sp. nov. is similar to Cralopa but differs most significantly in having fewer and less crowded ribs on the protoconch; fewer and less crowded teleoconch ribs that are bolder and much less sinuated than in Cralopa ; and in lacking an apertural sinus. Gouldiropa carlessi (Stanisic, 1990) comb. nov. (Figs 10, 20-22, 25-29, 33-34; Table 4) Cralopa carlessi Stanisic, 1990:156, figs 101-103 (in part); Smith. 1992: 184 (in part). MATERIAL. HOLOTYPE: NSW: AMSC123538, Inverell (29°46.5'S, 151°6.8’E), C.T.Musson. Height of shell=2.04. diameter=3.62, H/D=0.56, number of whorls=4.5, ribs on last whorl=70. PARATYPES. AMSC153716, C3640, C28496, C123610. OTHER MATERIAL. AMSC164886, QMM049I95, M052694, M032090, MO49207. DIAGNOSIS. Shell very small, brown with moderately loosely coiled whorls and a barely visible apertural sinus; whorls flattened above a rounded periphery. Spire and early apex slightly elevated. Teleoconch with numerous prominent strongly protractively sinuated radial ribs (mean ribs/mm 6.03). Umbilicus very slightly open, diameter range 0.26-0.43mm, mean D/U 5.49. Penis with two longitudinal pilasters. DESCRIPTION. (Amended from Stanisic, 1990). Shell very small golden brown with 3.8-4. 1 (mean 4.0) moderately loosely coiled whorls; apertural sinus very weakly developed. Shell diameter 2.96-3.44mm (mean 3.20mm). Apex and early spire weakly elevated. Height of shell 1.6 1-1. 88mm (mean 1.76mm), H/D 0.49-0.61 (mean 0.56). Protoconch with 1.5 whorls. Apical sculpture primarily radial consisting of approximately 40 regularly and widely spaced, curved radial ribs; ribs with height less than width; width of interstices (in multiples of rib width) 4-8 at apex. Secondary sculpture of continuous, regualrly crowded spiral cords, crossing the radial ribs. Teleoconch with prominent strongly protractively sinuated, variably spaced radial ribs that become more spaced on the last whorl; 46-74 (mean 60) ribs on the last whorl. Ribs/mm 5.14-6.91 (mean 6.03). Ribs wide, height equal to width, straight in section, rounded on top. Interstices on the 1st post-nuclear whorl equal to width of 4-6 ribs; on the penultimate whorl, greater than or equal to width of 6 ribs. Interstitial sculpture of very low microradial ribs and stronger microspiral cords: 10-12 microradials between ribs on 1st post-nuclear whorl; 12-16 between ribs on penultimate whorl. Microspiral cords crossing microradials but not forming distinct beads at their intersection. Umbilicus very narrowly open, U-shaped. Umbilical width 0.26-0.43mm (mean 0.32mm), D/U 5.21-5.77 (mean 5.49). Aperture ovately lunate. Whorls rounded above and below a weakly rounded periphery, sutures moderately impressed; parietal callus present. Based on 8 measured specimens (AMSC3640). Genitalia with ovotestis containing two clumps of alveoli, with two alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penial retractor muscle less than half the length of the penis, inserting at the junction of the penis and epiphallus. Epiphallus entering penis through a simple pore; shorter than penis; narrower than the vas deferens. Penis tubular, internally with two longitudinal pilasters. Vagina shorter than penis. Based on 8 measured specimens (AMSC3640). Pallial cavity with moderately bilobed kidney, pericarial lobe reflexed. Mantle gland present. Radula: central tooth strongly tricuspid; lateral teeth tricuspid; ectocone of outer marginal teeth split into 2-3 teeth; endocone not split. Lateral teeth: 5; marginal teeth: 14; radular rows: 106. Based on 1 dissected specimen (QMM032080). DISTRIBUTION AND HABITAT. N New England Tableland, NSW; under rocks on limestone outcrops covered in vine thicket or in moist sclerophyll forest. REMARKS. Gouldiropa carlessi Stanisic ( 1 990) comb. nov. is herein restricted to include only populations from the northern part of the New England Tableland, NSW. Stanisic (1990) considered that this species had an extensive range from Wee Jasper, SE NSW to Inverell, NE NSW wherein intraspecific variability included considerable differences in rib spacing, spire protrusion and umbilical width. That study, however, was constrained by a paucity of material for examination. Since that time additional specimens have become available for scrutiny and have enabled a more detailed analysis of these characters to be undertaken. A closer inspection indicates that a number of species were involved. Unfortunately, and some- what hastily, some of these were made paratypes for the nominal species, a salutary lesson for using only topotpes in the formal type series. Specimens cited by Stanisic (1990) as coming from limestones on Glenrock Station (AMSC340138, formerly AMSC 144200) are assigned to Sharniropa xanana. Specimens from SE NSW (Tuglow Caves [AMSC1 24279], 278 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 29. Radula. A-C, Cralopa stroudcnsis, QMM029638. D-F, Gouldiropa carles si, QMM032080. G-I, Gouldiropa kaputarensis , QMM049175. J-L, Sharniropa wollondillyana, QMM029241. M-O. Acheronopa attunga , QMM049218. Scale bars=0.01mm in A-L; 0.02mm in M-O. NEW CHAROPID LAND SNAILS 279 Jenolan Caves [QMM017295], Wee Jasper [AMSC142961] and Wombeyan Caves [AMSC 142967]) probably represent several species and are excluded. They will be dealt with elsewhere once more material becomes avilable for study. Gouldiropa carlessi is distinguished from G. kaputarensis by its larger size, flatter whorls, slightly more open umbilicus, more elevated spire and by having fewer ribs on the teleoconch. Sharniropa xanana from the southern part of the New England Tableland has shouldered whorls, fewer ribs on the teleoconch and a more open umbilicus. G. carlessi is sympatric with the similarly shaped Acheronopa attunga but is readily distinguished from that species by its smaller size, almost closed umbilicus (widely open in A . attunga) and in having more crowded radial ribs on the teleoconch. Sharniropa gen. nov. ETYMOLOGY. For Sham Rose. TYPE SPECIES. Sharniropa wollondillyana sp. nov.; herein designated. DIAGNOSIS. Shell very small, dark brown, with moderately loosely coiled whorls and strongly impressed sutures; apertural sinus absent. Protoconch with relatively widely spaced, crowded curved radial ribs (approx. 45-50) and numerous, regularly crowded, low continuous spiral cords that cross the radial ribs. Teleoconch with numerous, bold, weakly to very strongly protractively sinuated, crowded to very widely spaced radial ribs. Interstitial sculpture with microspiral cords stronger than microradial ribs, beading at their intersection weak to barely noticeable, obscured by strength of microspiral cords. Umbilicus narrowly open U-shaped. Columellar edge of aperture not reflected across the umbilical opening. Kidney moderately to strongly bilobed with reflexed apex on pericardial lobe. Penis internally with two longitudinal pilasters. DISTRIBUTION AND HABITAT. N part of the Southern Tableland Bioregion to the S end of the New England Tableland Bioregion; only on limestone outcrops living among limestone talus. REMARKS. Sharniropa gen. nov. differs from Gouldiropa mainly in the coiling pattern of the shell. In Sharniropa narrowing of the umbilicus is caused by constriction, in Gouldiropa it is a combination of constriction and columellar reflection. Acheronopa differs in having a greater number of apical radial ribs (approx. 60), a larger, flatter shell with evenly coiled whorls and a wider umbilicus caused by gradual decoiling. Sharniropa wollondillyana sp. nov. (Figs 1 0, 23, 25-29, 3 1 -34; Table 5) ETYMOLOGY. For the Wollondilly River. MATERIAL. HOLOTYPE. QMMO70399, Abercrombie Caves, SW Oberon, NSW, (33°54’42"S, 149°2r36"E), remnant vt 1 lm, under logs and rocks, 13.i.l993, JS, JC. Height of shell= 1 .43mm, diameter=2.57mm. H/D=0.56, D/LM.51, number of \vhorls=3.9, ribs on last whorl=31. PARATYPES. All NSW. QMM04477 1 , 48RC. same data as holotype; QMM029241, IRC, QMM03778L 10RC, Abercrombie Caves, SW Oberon, lm, under rocks and in litter, 8.i.l990, JS, JC: AMSC 157323, 7RC, SW of Bathurst, Bombah, 100m S of Abercrombie Caves camping ground (33°54.8’S, 149°2L5'E), lm, under rocks and soil on limestone hill, 6.vi.l982, MS; AMSC1 58364, manv RC, S of Blayney, Abercrombie Cave, near Bushranger Cave (33°55'S, 149°2rE), vii.1982, O.L. Griffiths. OTHER MATERIAL. AMSC 137245. DIAGNOSIS. Shell very small, brown with moderately loosely coiled whorls; whorls weakly shouldered above a rounded periphery. Teleoconch with widely spaced, prominent, strongly protractively sinuated radial ribs (mean ribs/mm 3.93). Umbilicus open, very narrow U-shaped; umbilical diameter range 0.55- 0.73mm, mean D/U 4.43. Penis with two longitudinal pilasters. DESCRIPTION. Shell very small, colour brown with 3. 7-4. 3 (mean 4.0) moderately loosely coiled whorls. Shell diameter 2. 27-3. 34mm (mean 2.84mm). Apex and early spire barely elevated. Height of shell 1.21- 1.54mm (mean 1.43mm), H/D 0.45-0.58 (mean 0.51). Protoconch of 1.5- 1.8 whorls with sculpture of approximately 45-50, crowded, curved, regularly spaced radial ribs, ribs of medium width; height equal to width, width of interstices (in multiples of rib width) 6-8 at apex. Secondary sculpture of low continuous, eroded and regularly spaced spiral cords that cross the radial ribs. Teleoconch sculpture of widely and regularly spaced (becoming more crowded at the end of the last whorl), strongly protractively sinuated radial ribs; 26-34 (mean 35) ribs on the last whorl, ribs/mm 2.79-5.00 (mean 3.93). Ribs wide; height greater than width; straight in section; rounded on top; interstices on the first post-nuclear whorl equal to width of 8-10 ribs; interstices on the penultimate whorl equal to width of 8 ribs. Interstitial sculpture of weak microradial ribs 20 between ribs on first post-nuclear whorl; 20 between ribs on 280 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 30. Radula. A-C, Hedleyropa yarrangobillyensis , QMMO39840. D-F, Scleridoropa sarahjaneae (Yarranuobilly), QMM039974. G-I, Scleridoropa nandewar ; QMM049158. J-L, Diphyoropa macleayana, AMSC 162 1 84. Scale bars=0.0 1 mm. penultimate whorl. Secondary sculpture of more prominent, low microspiral cords with weak beading at their intersection with the microradial ribs. Umbilicus moderately wide U-shaped, diameter 0.55-0. 73mm (mean 0.64mm), D/U 3.99-5.03 (mean 4.43). Sutures impressed, whorls weakly shouldered above and rounded below a rounded periphery. Aperture ovately to roundly lunate. Based on 19 measured specimens (QMM044771, MO70399). Genitalia with ovotestis containing two clumps of alveoli, with two alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penial retractor muscle less than half the length of the penis, inserting at the junction of the penis and epiphallus. Epiphallus shorter than penis; narrower than the vas deferens; entering penis through a simple pore. Penis tubular, two longitudinal pilasters present. NEW CHAROPID LAND SNAILS 281 Pallial cavity with kidney moderately bilobed; apex of pericardial lobe reflexed. Mantle gland present. Radula with central tooth strongly tricuspid; lateral teeth tricuspid; ectocone of outer marginal teeth not split; endocone not split. Number of lateral teeth: 6; marginal teeth: 12; radular rows: 97. Based on 3 dissected specimens (QMM02924 1 ). DISTRIBUTION AND HABITAT. Aber- crombie limestones, NSW; under logs and rocks. REMARKS. Sharniropa wollondillyana sp. nov. differs from S. boronorensis in having fewer radial ribs on the teleoconch and a larger umbilicus. S. wollondillyana is superficially similar to the sympatric Decoriropa l i rata but differs primarily in having fewer apical radial ribs. D. lirata is also smaller than S. wollondillyana , has a relatively larger umbilicus and more crowded ribs on the teleoconch. Sharniropa borenorensis sp. nov. (Figs 23, 25-28; Table 4) ETYMOLOGY. For the type locality. MATERIAL. HOLOTYPE: AMSC205166, Borcnore Caves, W side of large cave entrance W of Borenorc, W of Orange, NSW (33°15.14'S, 148°56.19’E), in soil and grass among limestone boulders, vi. 1 983, MS. Height of shcll= 1.50mm, diamcter=2.77mm, H/D=0.54, D/U=5.54, number of whorls=3.7, ribs on last whorl=50. PARATYPES. All NSW. AMSC 1 57300, 1 3RC same data as holotype; QMM044745, 12RC, Borcnore Caves, c. 18.6km WNW Orange (33°15*06"S, 148°55’42 M E), lm/ew, in litter iunder rocks, 1 Li. 1993. JS, JC; QMM037496, 30RC, Borcnore Caves, c. 1 0km W Orange (33°15’S. 148°56*E), remnant vt/lm, among rocks, 13.i. 1991, JS. DIAGNOSIS. Shell very small, brown, with moderately loosely coiled; whorls weakly shouldered above a rounded periphery. Teleo- conch with crowde, prominent, strongly protractively sinuated radial ribs (mean ribs/mm 7.13). Umbilicus open, very narrow U-shaped; umbilical diameter range 0.50-0.71 mm, mean D/U 4.95. Anatomy unknown. DESCRIPTION. Shell very small, brown, with 3. 7-4. 2 (mean 4.0) moderately loosely to evenly coiled whorls. Shell diameter 2.64-3. 11 mm (mean 2.84mm). Apex and early spire barely elevated. Height of shell 1.33-1. 54mm (mean 1.42mm), H/D 0.45-0.54 (mean 0.50). Protoconch of 1.5- 1.7 whorls with curved, regularly spaced radial ribs; ribs of medium width, height less than width, width of interstices (in multiples of rib width) 5 at apex. Secondary sculpture of low, continuous regular spaced spiral cords that cross the radial ribs. Teleoconch sculpture of crowded, regualrly spaced (but becoming more crowded toward the end of the last whorl), strongly protractively sinuated radial ribs; 50-79 (mean 64) ribs on the last whorl, ribs/mm 5.75-8.09 (mean 7.13). Ribs wide; height equal to width; straight in section; rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 3-4 ribs. Interstitial sculpture of microradial ribs, 5-6 between ribs on first post-nuclear whorl; 8 between ribs on penultimate whorl. Secondary microsculpture of of prominent low, regularly spaced microspiral cords that cross the microradial ribs without froming distinct beads. Umbilicus very narrow U-shaped, diameter 0.50-0.7 1mm (mean 0.58mm), D/U 4.34-5.54 (mean 4.95). Sutures impressed, whorls weakly shouldered above and rounded below a weakly rounded periphery; Aperture roundly to ovately lunate; parietal callus present. Based on 8 measured specimens (AMSC 157300, C205166). Anatomy unknown. DISTRIBUTION AND HABITAT. Known only from the type locality at the Borenore Caves limestone outcrop; found under grass amongst limestone boulders. REMARKS. Sharniropa borenorensis sp. nov. is similar to S. wollondillyana from the Wombeyan and Abercrombie limestone outcrops. It differs mainly from the latter species in having a greater number of radial ribs on the last whorl (50-79, mean 64 compared with 26-34, mean 35 for S. wollondillyana) and a smaller umbilicus. Sharniropa xanana sp. nov. (Figs 23, 25-28; Table 4) Cralopa carlessi Stanisic, 1990:156 (in part), figs lOld-e; 102a-e; Smith, 1992: 184 (in part). ETYMOLOGY For Xanana Gusmao, President of East Timor. MATERIAL. HOLOTYPE: QMMO70398, Timor Caves, via Murrunmdi, NSW (31°41'S, 151°08'E), lm/remnant vt, in litter, 25.L\1989. JS, DP. Height of shell= 1.74mm, diameter=3.34mm, II/D 0.52, D/U 5.96, number of whorls=4.1, ribs on last whorl=18. PARAT YPES. All NSW. QMM037735, 29RC, AMSC205159, 5RC, same data as holotype; QMMO32065, 1 juvenile SC/6RC, in soil among rocks, otherwise same data as holotype. OTHER MATERIAL. QMM056149, AMSC 144200. DIAGNOSIS. Shell very small, light golden brown, with moderately loosely coiled whorls 282 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 31. Reproductive system. A, Shamiropa wollondillyana , QMM029241. B, Acheronopa attunga , QMM049218. C, Hedleyropa yarrangobi l lyensis , QMMO39840. D, Scleridoropa nandewar, ; QMM049158. E. Scleridoropa sarahjaneae (Yarrangobilly), QMM039974. F, Diphyoropa macleayara , AMSC162184. Scale bar=2.5mm in A and C, 2mm in B, and 1 .6mm in all others, dg, prostate; e, epiphallus; g, ovotestis; gd, hermaphrodite duct; gg, albumen gland; gt, talon: p, penis; pirn, penis retractor muscle; s, spermatheca; ss, spermathecal stalk; ut, uterus; uv, free oviduct; v, vagina; vd, vas deferens. and a barely visible apertural sinus; whorls strongly shouldered above a rounded periphery. Spire and early apex flat to barely elevated. Teleoconch with relatively few, widely spaced, prominent, strongly protractively sinuated radial ribs (mean ribs/mm 1.86). Umbilicus open, very narrow U-shaped; umbilical diameter range 0.45-0. 62mm, mean D/U 6.41. Anatomy unknown. DESCRIPTION. Shell very small, light golden brown, with 3.9-4.5 (mean 4.09) loosely coiled whorls. Shell diameter 3.04-3.6 1mm (mean 3.3 1 ). Apex and early spire very weakly elevated. Height of shell 1.45- 1.86mm (1.60mm), H/D 0.44-0.52 (mean 0.48). Protoconch relatively large with sculpture of approximately 40, regularly and widely spaced, weakly curved radial ribs; ribs of medium width; height less than width; width of interstices (in multiples of rib width) 3-5 at apex. Secondary sculpture of crowded, continuous microspiral cords that cross the microradial ribs. Teleoconch with prominent, regularly spaced, strongly protractractively sinuated, bold radial ribs, 16-25 (mean 19) ribs on the last whorl. Ribs/mm 1.65-2.20 (mean 1 .86). Ribs greater in height than width; straight in section; rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 1 3 ribs; on the penultimate whorl, equal to width of 13 ribs. Interstitial sculpture of very low microradial ribs and stronger microspiral cords; 30 microradials between ribs on first post-nuclear whorl, becoming almost impossible to count on penultimate whorl; beading indistinct due to NEW CHAROPID LAND SNAILS 283 TABLE 4. Shell measurements for species of Coricudgia , Marilyniropa , Cralopa , Gouldiropa and Sharniropa. (Abbreviations as in Table 1). Measurement Coricudgia wollemiana Marilyniropa jenolanensis Cralopa stroudensis Gouldiropa carlessi Sharniropa borenorensis Sharniropa xanana N 10 (*9; **4; ***3) 23 15 10 8 9 D 1.62-4.02 (3.304) ± 0.75 2.96-3.65 (3.24) ± 0.194 1.63-3.13(2.58) ± 0.35 2.66-3.84(3.21) ± 0.32 2.64-3.11 (2.841) ± 0.186 3.04-3.61 (3.307) ± 0.204 PW 1.7-1.8(1.775) ± 0.05 ** 1.5-1.7(1.587) ± 0.069 1.5-1.7(1.59) ± 0.07 1.5 1.5-1.7(1.563) ± 0.074 1.5-1.7(1.589) ± 0.06 TW 1. 8-4.3 (3.1) ± 1.253 *** 2. 6-3.2 (2.922) ± 0.165 1.3-2.7(2.38) ± 0.33 2.2-2.6 (2.43) ± 0.12 22-2.1 (2.425) ± 0.158 2.3-2. 8 (2.5) ± 0.141 NW 3.6-6. 1 (4.9) ± 1.253 *** 4.1-4.8(4.509) ± 0.165 2. 8-4.3 (3.97) ± 0.36 3.7-4. 1 (3.93) ± 0.12 3.7-4.2 (3.987) ± 0.155 3. 9-4.5 (4.089) ± 0.169 H 0.83-2.21 (1.691) ± 0.435 1.33-1.79(1.549) ± 0.146 1.02-1.73 (1.53) ±0.17 1.31-2.03(1.74) ± 0.19 1.33-1.54(1.421) ± 0.071 1.45-1.86(1.6) 1 ± 0.133 SP 0.06-0.47 (0.259) ± 0.14 0.11-0.41 (0.25) ± 0.084 0.05-.18 (0.13) ± 0.04 0.1-0.33 (0.24) ± 0.07 0.09-0.21 (0.154) ± 0.038 0.06-0.3 (0.211) ± 0.079 AH/AW 0.75-1.14(0.92) ± 0.11 0.78-0.98 (0.9) ± 0.05 0.81-1.11 (0.9) ± 0.08 0.77-0.93 (0.84) ± 0.06 0.77-0.91 (0.84) ± 0.05 0.65-0.79 (0.74) ± 0.05 UMB 0.55-1.15(0.938) ± 0.177 0.73-1.07(0.888) ± 0.087 0.14-0.44 (0.34) ± 0.07 0.26-0.54 (0.36) ± 0.1 0.5-0.71 (0.58) ± 0.083 0.45-0.62 (0.522) ± 0.059 RIB 28-47 (40.222) ± 6.36 * 35-61 (47.087) ± 7.366 68-126 (88.73 ) ± 16.45 28-74 (56.4) ± 13.62 50-79 (63.625) ± 8.193 16-25(19.444) ± 3.206 RIBS/MM 3.008-5.502 (4.089) ± 0.695 * 3.764-6.078 (4.616) ± 0.588 7.904-16.50 (10.83) ± 2.241 3.35-6.85 (5.56) ± 1.13 5.746-8.086 (7.125) ± 0.735 1.649-2.204 (1.862) ± 0.201 H/D 0.45-0.55 (0.51) ± 0.03 0.43-0.55 (0.48) ± 0.03 0.53-0.66 (0.6) ± 0.04 0.49-0.58 (0.54) ± 0.02 0.45-0.54 (0.5) ± 0.03 0.44-0.52 (0.48) ± 0.03 D/U 2.95-3.8 (3.49) ± 0.23 3.33-4.05 (3.67) ± 0.22 5.68-11.64(7.81) ± 1.63 5.22-13.23 (9.57) ± 2.55 4.34-5.54 (4.95) ± 0.4 5.29-7.85(6.41) ± 0.91 strength of microspiral cords. Sutures impressed, whorls strongly shouldered above and rounded below a weakly rounded periphery. Sutures strongly impressed. Umbilicus very small, narrow U-shaped. Umbilical width 0.45-0. 62mm (mean 0.52mm), D/U 5.29-7.85 (mean 6.41). Aperture ovately lunate, parietal callus present. Based on 9 measured specimens (QMMO32065, M037735. MO70398). DISTRIBUTION AND HABITAT. Limestone outcrops at Timor Caves and Glcnrock Stn, NE of Scone, NSW; in litter among limestone talus. REMARKS. Sharniropa xanana sp. nov. differs from its congeners by its strongly shouldered whorls, and relatively sparse telcoconch ribbing. Superficially the species bears a greater similarity to Acheronopa attimga. Both have a flattened shell with bold, widely spaced radial ribs. However, .S’, xanana is smaller, has a larger protoconch, fewer, more loosely coiled and more shouldered whorls, microsculpture with comparatively indistinct beading and a more closed umbilicus. This species was considered to be conspecific with Gouldiropa carlessi Stanisic, 1990 comb. nov. by Stanisic (1990). The Timor and Glenrock limestone outcrops are among open woodland. Vegetation on the outcrops is degraded due mainly to human activity (fires, land clearing) but there is evidence that vine thicket may have been present on these in the past (Stanisic, pers. obs.). Acheronopa gen. nov. ETYMOLOGY. Greek Acheron, river in the nether world; alluding to the underground streams that have formed many of the caverns present in limestone formations. TYPE SPECIES. Archemnopa attimga sp. nov.; herein designated. DIAGNOSIS. Shell very small, brown, strongly depressed (mean H/D 0.40), with flattened, evenly coiled whorls. Apical sculpture of relatively crowded, curved radial ribs (approx. 60) and numerous, weaker, continuous low microspiral cords. Teleoconch with strongly sinuated, very widely spaced, bold radial ribs. Secondary microsculpture of low microspiral cords and microradial riblets forming beads at their intersection. Kidney moderately bilobed with apex of pericardial lobe reflexed. Umbilicus 284 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 32. Pallial cavity. A, Shamiropa wollondillyana , QMM029241 . B, Acheronopa attunga , QMM0492 1 8 . C, Hedleyropa yarrangobillyensis, QMMO39840. D, Scleridoropa nandewar, ; QMM049158. E, Scleridoropa sarahjaneae , QMM039974. F, Diphyoropa macleayana , AMSC 1 62 1 84. Scale bar=2mm in A and F, 1 .6mm in B, and 1.28mm in all others. Abbreviations: h, heart; hv, principal pulmonary vein; i, intestine; kdl, primary ureter; kd2, secondary ureter; kp, pericardial lobe of kidney; kr, rectal lobe of kidney; me, mantle collar; mg, mantle gland. wide V-shaped. Penis with 3 longitudinal pilasters. DISTRIBUTION AND HABITAT. Yarra- manbully limestones near Attunga; between limestone talus on limestone karst covered in remnant vine thicket REMARKS. Acheronopa gen. nov. is most similar to Shamiropa gen. nov from the Timor/ Glenrock limestone outcrops and Gouldiropa gen. nov. However, it differs significantly in having more crowded microradial ribs on the protoconch and in having more flattened whorls. NEW CHAROPID LAND SNAILS 285 Acheronopa also differs from the sympatric Gouldiropa in having an open rather than closed umbilicus and in the more prominent, sparser teleoconch ribbing. The unusually depressed shell shape may be a functional adaptation to living between limestone slabs. Acheronopa attunga sp. nov. (Figs 10, 23, 25-29, 31-34; Table 5) ETYMOLOGY. For the New England town of Attunga. MATERIAL. HOLOTYPE: QMM070400, Manilla, c. 20.4km ESE, Manilla-Yarramanbully Rd, NSW (30°5 1 WS, 1 50°50’54 M E), lm/vt, under rocks in talus pile, 25.ix. 1 993, JS, JC. I leight of shell= 1 .8 1 mm, diameter=4.27, H/D=0.42, D/U=3.21, number of \vhorls=4.9, ribs on last whorl=23. PARATYPES: All NSW. QMM049218, 42SC/49RC, same data as holotype; QMM049187, 7RC, Manilla, c.6.6km ESE at Amaroo Homestead (30 t, 46’06 M S, 1 50°47’36"E), vt/lm, under rocks abd logs and in litter, 24.ix. 1993, JS, JC: QMM049193, 8RC, Manilla, c. 19.9km ESE, Manilla-Yarramanbully Ck Rd (30°50 , 42 M S, 150.51T2 M E), lm/vt, under rocks and logs, 24.ix.1993, JS. JC; QMM052693, 18RC, Manilla, c.20.4km ESE, Manilla-Yarramanbully Rd, (30°5r00"S, 1505L54E), lm/vt, in litter, 25.ix.1993, JS, JC: QMMO32089, 100RC, AMSC205 1 60, I0RC, Attunga, c.l2km N, Woodbum- Yarramanbul ly Rd (30°51’S, i50°51’E), Im, under and betw een rocks, 26.xi.1989, JS, DP. DIAGNOSIS. Shell very small, with strongly depressed, evenly coiled whorls. Teleoconch with strongly sinuated, very widely spaced, bold radial ribs, mean ribs/mm 1.72; secondary sculpture of prominent microspiral cords and low radial ridges, forming prominent beads at their intersection. Umbilicus wide V-shaped. Penis with three longitudinal pilasters. DESCRIPTION. Shell very small, depressed, brown, with 4.6-5. 2 (mean 5.0) evenly coiled whorls. Shell diameter 3. 92-6. 28mm (mean 4.65mm). Apex and early spire depressed. Height of shell 1.59-2.1 1mm (mean 1.85mm), H/D 0.30-0.48 (mean 0.40). Protoconch of 1.5- 1.7 whorls with sculpture of crowded, regularly spaced, slightly curved radial ribs, width of interstices ( in multiples of rib width) 5-7 at apex. Secondary apical sculpture of crowded, regularly spaced, continuous microspiral cords. Teleoconch sculpture of strongly sinuated, very widely spaced, bold radial ribs, increasing in spacing toward aperture, ribs on last whorl 20-3 1 (mean 25), ribs/mm 1.22-2.12 (mean 1.72). Ribs wide, height less than width; straight in section, rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 3-5 ribs; interstices on the penultimate whorl equal to width of 6-8 ribs. Interstitial sculpture of low microradial ribs, 7-9 between major ribs on first post-nuclear whorl; 16-20 between ribs on penultimate whorl. Secondary sculpture of prominent low microspiral cords that cross the major radial ribs and form very strong, raised beads where they intersect with the microradials. Sutures strongly impressed, whorls fattened above and weakly rounded below a flattened periphery. Aperture broadly ovately lunate. Umbilicus open, wide V-shaped, diameter 1.22-1.7 1mm (mean 1.48mm), D/U 2.89-4.21 (mean 3.16). Based on 23 measured specimens (QMM049218, M070400). Genitalia with ovotestis containing two clumps of alveoli, with more than two alveolar lobes per clump. Talon stalk 1-3 times the diameter of the talon; talon circular. Penal retractor muscle less than half the length of the penis; inserting at the junction of the penis and epiphallus. Muscle fibres also attached to the base of the penis, close to the vagina/atrium. Epiphallus narrower than the vas deferens, entering penis through a simple pore; shorter than penis. Penis pear shaped, internally with three longitudinal pilasters, one much smaller than the other two. Vagina very short. Pallial cavity with moderately bilobed kidney, apex of pericadial lobe reflexed. Mantle gland present. Radula with central tooth strongly tricuspid; lateral teeth tricuspid; first and second laterals equal in width. Mesocone of first lateral tooth slender, slightly diamond-shaped; marginal teeth skewed towards the centre; ectocone of outer marginal teeth not split; endocone not split. Number of lateral teeth: 6; marginal teeth: 18; radular rows: 94. Based on 3 dissected adults (QMM049218). DISTRIBUTION AND HABITAT. Acheronopa attunga sp. nov. is known only from the Yarramanbully limestones near Attunga; living between limestone talus on limestone karst covered in remnant vine thicket. REMARKS. Acheronopa attunga sp. nov. is most similar to Sharniropa xanana sp. nov from the Timor/Glenrock limestone outcrops. However, it differs significantly from that species in being relatively much larger, having more crowded microradial ribs on the protoconch and in having more fattened whorls. A. attunga difiers from the sympatric Gouldiropa carlessi (Stanisic, 1990) comb. nov. most noticeably in having an open rather than closed umbilicus and 286 MEMOIRS OF THE QUEENSLAND MUSEUM NEW CHAROPID LAND SNAILS 287 in havimg more prominent, sparser teleoconch ribbing. The unusually depressed shell of A. attunga may be a functional adaptation to living between limestone slabs. Hedleyropa gen. nov. ETYMOLOGY. Named in honour of Charles Hedley. TYPE SPECIES. Hedleyropa yarrangobillyensis sp. nov.; herein designated. DIAGNOSIS. Shell very small, light brown in colour. Apex and early spire flat. Protoconch with very densely crowded, curved radial ribs (approx. 90), crossed by low, crowded spiral cords. Teleoconch with protractively sinuated radial ribs and a secondary sculpture of low radial ridglets and more prominent, crowded micro- spiral cords; microsculpture continuous on the major radials. Kidney moderately bilobed with weakly reflected pericardial lobe. Penis with two longitudinal pilasters. DISTRIBUTION AND HABITAT. Yarrangobilly limestones, Kosciusko NP, SE NSW and surrounding environs. However, there are other undescribed species in the QM and AM collections from the region which may belong to this genus and it is quite probable that the range of the genus could be extended significantly particularly in other parts of the Kosciusko NP and NE Vic. REMARKS. Hedleyropa gen. nov. is similar to Cralopa in having crowded apical radial ribs and numerous bold, relatively crowded sinuated teleoconch ribs, but differs in having a greater number of microradial ribs on the protoconch, no apertural sinus, finer and much less sinuate teleoconch ribs and a very narrow U-shaped umbilicus rather than one closed by columellar reflection. There is also a subtle difference in whorl profile between the two genera with Hedleyropa having more rounded whorls compared with that of Cralopa. Hedleyropa varrangohillvensis sp. nov. (Figs 10, 23, 25-28, 30-34; Table 5) ETYMOLOGY. For the type locality. MATERIAL. HOLOTYPE. QMM070401, Yarrangobilly, Yarrangobilly R, c.lkm S bridge, NSW (35°39°28"S, 148°27’4rE), lm/ew, under rocks, 8.x. 1992, JS, GI. Height of shell=2.20mm, diameter=3.74, H/D=0.54, D/TJ=4.35, number of whorls=4.5, ribs on last whorl=123. PARATYPES. All NSW. QMM039973, 1 SC/50+RC, same data as holotypc; QMMO39840, 5SC, Tumut, c.42.3km SE, Tumut - Yarrangobilly Rd (35°35’ 1 6"S, 1 48°20°04 M E), ew, under rocks. 8.x. 1 992, JS, GI; QMM049419, IRC, Yarrangobilly, c.4.4km N turnoff to caves, Tumut - Kiandra Rd (35°4 T00"S, 148°2ff54"E), ew/lm, under logs and rocks, 03.L1994, JS, JC; QMM039975, 5SC/3RC, Yarranuobilly, Yarrangobilly Caves Reserv e (35°43'45 M S, 148°29T6 M E), ew/lm, under rocks, 8.x/ 1992. JS, GI; AMSC1 62152, 6RC. Kosciusko NP. Yarrangobilly Caves, Village Cave System [?=Y-1 10] (35°43.5*S, 148°29.5'E), on wet moss in overhang, 6.vi.l980, Ken Keck. DIAGNOSIS. Shell very small, with evenly coiled whorls and an almost flat spire. Teleoconch with bold, relatively crowded, protractively sinuated radial ribs mean ribs/mm 9.16). Umbilicus narrow U-shaped, mean D/U 4.3; whorls rounded at the periphery and evenly coiled. Penis with two longitudinal pilasters. DESCRIPTION. Shell very small, light brown in colour with 4.2-4.7 (mean 4.5) evenly coiled whorls, the last weakly descending in front. Shell diameter 3.21 -4.52mm (mean 3.82mm). Apex and early spire low to flat. Height of shell 1.63-2.5 1mm (mean 1.98mm), H/D 0.48-0.58 (mean 0.52). Protoconch of 1.6- 1.9 whorls with sculpture of approximately 90, weakly curved, uniformly spaced radial ribs; ribs of medium width with height less than width; width of interstices (in multiples of rib width) 3-4 at apex. Secondary sculpture of much lower, narrow, low continuous spirals (approx. 37); height of spirals less than width. Teleoconch sculpture of strongly protractively sinuate, variably spaced radial ribs; spacing increasing towards aperture; 86-136 (mean 109) ribs on the last whorl. Ribs/mm 8.17-1 0.72 (mean 9.16). Rib height equal to width; straight in section; rounded on top; interstices on the first post-nuclear whorl equal to width of 4-6 ribs; interstices on the penultimate whorl equal to 4-5. Interstitial sculpture of low radial ridges, 8-9 between ribs on first FIG. 33. Protoconch microsculpture. A, Rhophodon megradyorum , QMMO70392. B, Egilodonta paucidentata , QMMO70396. C, Letomola lanalittieae , QMMO70397. D, Macrophallikoropa depressispira , AMSC205163. E, Elsothera brazieri , QMM028775. F, Decoriropa lirata (Taree), AM SC 136842. G, Coricudgia wollemiana , AMSC205164. H, Marilyniropa jenolanensis , AMSC205165. 1, Gouldiropa carlessi (Inverell), AMSC3640. J, Shamiropa wollondillyana , QMMO70399. K, Acheronopa attunga , QMM070400. L, Hedleyropa yarrango- billyensis , QMM070401. M, Scleridoropa sarahjaneae (Abercrombie Caves), AMSC162165. N-P. Diphyoropa macleayana , AMSC 1 68685. Scale bars=30pm in A, D; 40pm in B, C, E, F; 50pm in G, H, J, M, O; 60pm in I, K, L, N, P. 288 MEMOIRS OF THE QUEENSLAND MUSEUM post-nuclear whorl; 8 between ribs on penultimate whorl; crossed by more prominent, very low microspiral cords. Beading weak. Umbilicus narrow U-shaped, diameter 0.70-1. 20mm (mean 0.90mm), D/U 3.72-5.05 (mean 4.30). Sutures strongly impressed, appearing weakly channeled; whorls slightly shouldered above and rounded below a rounded periphery. Aperture roundly lunate. Umbilicus open, narrow U-shaped. Based on 20 measured specimens (QMM039973, M070401). Genitalia with ovotestis containing two clumps of alveoli. Talon stalk slightly longer than the diameter of the talon; talon circular. Penial retractor muscle less than half the length of the penis; inserting at the junction of the penis and epiphallus. Epiphallus present; entering penis through a simple pore; slightly shorter than penis; narrower than the vas deferens. Penis tubular with two longitudinal pilasters. Vagina shorter than penis. Pallial cavity with kidney moderately bilobed; apex reflexed. Radula with central tooth strongly tricuspid; lateral teeth tricuspid; ectocone of outer marginal teeth not split (but the ectocone of the second last tooth is split into three); endocone not split. Number of lateral teeth: 4, marginal teeth: 15; radular rows: 85. Based on 1 dissected specimen (QMMO39840). DISTRIBUTION AND HABITAT. Yarrangobilly limestones, SE Talbingo, SE NSW; living among among limestone slabs. REMARKS. Hedleyropa varrangobillyensis sp. nov. is readily distinguished from other sympatric charopids by a combination of its dense radial ribbing on both the protoconch and teleoconch and the comparatively reduced umbilicus. Scelidoropa sarahjaneae sp. nov. has a radially ribbed teleoconch but differs in having a larger flammulated shell with wide cup-shaped umbilicus and a protoconch with microdaradial ribs in the interstices of the major ribs. Anatomically the two species are markedly distinct. Hedleyropa yarrangobillyensis displays a tendency for reduction in the size of the umbilicus. The general shell form (weakly channelled sutures, bold ribbing, numerous apical radial ribs) recalls Cralopa stroudensis (Cox, 1864). However, the greater number of apical microradials, weaker sinuosity of the teleoconch ribs, lack of an apertural sinus and narrow but open U-shaped umbilicus all indicate that H. yarrangobillyensis is a member of Hedleyropa. Scelidoropa gen. nov. ETYMOLOGY. Greek skelidos, rib; alluding to the interstitial microradial ribs on the protoconch. TYPE SPECIES. Scelidoropa sarahjaneae sp. nov. DIAGNOSIS. Small, light brown shell with darker radial flammulations with slightly tightly coiled whorls. Spire flat to slightly elevated. Protoconch with crowded, evenly spaced, slightly curved radial ribs (approx. 90). Secondary sculpture of low continuous microspiral cords and low slightly curved microradial ribs that form beads at their intersection. Teleoconch with slightly sinuated, crowded to very crowded ribs; microsculpture of microradial ribs and microspiral cords equal in strength, forming strong beads at their intersection. Umbilicus wide saucer-shaped. Penis with 0-4 pilasters. Kidney almost unilobed. Central radula tooth weakly tricuspid; lateral teeth with a smaller additional cusp between ectocones of the tooth. DISTRIBUTION AND HABITAT. From SE NSW to the southern edge of the New England Tableland, NSW (Nundle area) and west to the Nandewar Range, NSW. The species live in litter among rocks. REMARKS. The shell of Scelidoropa gen. nov. is quite distinctive among the east coast charopid genera. The flammulated shell with elevated spire, bold and prominent teleoconch ribs and very wide umbilicus are a combination of characters that are seen in other charopid groups. However, the microradial ribs on the protoconch immediately identify it among the east-coast fauna. Species in Victoria that are currently questionably classified in Pernagera Iredale, 1939 (Smith, 1992) series also have apical radial sculpture, bold teleoconch ribs and flammulated shells, but, and most significantly, lack the microradial ribs on the protoconch. Some species of Elsothera Iredale, 1933, superficially resemble Scelidoropa but the latter can be distinguished by its much larger size, bolder, weakly sinuated ribs, wider cup shaped umbilicus and most significantly, by the present of intersitial microradials on the protoconch. Anatomically the two genera are immediately separable by the form of the epiphallus which is long and unusually coiled in Elsothera but shorter and more typically reflexed in Scelidoropa. The lateral teeth of the radula of NEW CHAROPID LAND SNAILS 289 Scelidoropa are also distinctive in the context of all the other eastern Australian charopids hitherto reviewed in possessing an additional cusp. The breadth of the range of Scelidoropa is matched only by that of Maavphallikoropa gen. nov. However, in contrast to the latter, the species’ distributions do not include rainforests and encompass only very localised mesic refugia such as rocky outcrops (limestone and granite), and mountain tops (Mt Kaputar, Mt Coricudgy and Mt Canoblas). The species is absent from the Sydney Basin and the limestones of the Macleay Valley. Scelidoropa sarahjaneae sp. nov. (Figs 10, 24-28, 30-34; Table 5) ETYMOLOGY. For Sarah Jane Roberts. MATERIAL. HOLOTYPE. QMM070402, Yarrango- billy, Yarrangobilly R, c.lkm S bridge (35°39’28"S, 148®27’4r’E), ew/lm, in litter, 08.10.1992, JS, GI. Height of shcll=L42mm, diameter=3.18mm, H/D=0.45, D/U=2.26, number of whorls=4.6, ribs on last whorl=122. PARATYPES. All NSW. QMM039974, 13SC/50RC, same data as holotype; QMM049418, 4RC, Yarrangobilly, c.lkm S on Tumut - Kiandra Rd (35°39’30"S, 148°27’42”E), ew/lm, under rocks, 03.i.l994, JS, JC; QMM049421, 5RC, QMM052775, 34RC, Yarrangobilly, c.4.4km N T’off to caves, Tumut - Kiandra Rd (35°4 1 W'S, 1 48°28 , 54"E), ew/lm, under logs and rocks, 03, i. 1994, JS, JC; QMM039979. 7SC/50RC, Yarrangobilly, Yarrangobilly Caves Reserve (35°43'45"S, 148°29 , 16 M E), ew/lm, in litter, 08.x. 1992, JS, Gl; QMMO68013, 50RC, Yarrangobillv, Yarrangobilly R, c.lkm S bridge (35°39'28"S, 148 0 27’41 M E), lm, in litter, 03.i. 1 994, JS, JC; AMSC153724, IRC, Kosciusko NP, Yarrangobilly Caves, beside trail to River Cave ( Y-27) (35.725°S, 148.492°E), in litter, 31.X.1980, WFP, j.Half; AMSC 1 68695, 1 RC, Kosciusko NP. Yarrangobilly Caves, Village Cave System (?= Y-l 1 0) (35.725°S, 148.492°E), on wet moss in overhang , 06.vi.1980, Ken Keck; AMSC 162 150, many RC, Kosciusko NP, Yarrangobilly Caves, beside trail to River Cave (Y-27) (35.725°S, 1 48.492°E), in litter, 31.x. 1980, WFP, J.Hall; AMSC 162151, 1 RC, Kosciusko NP, Yarrangobilly Caves (35.725°S, 148.492°E), 3 1.x. 1980, WFP, J.Hall. OTHER MATERIAL. NSW: AMSC 168626, C355262, C367084, Cl 62 140. C355269. Timor Caves: QMMO32063, M037731, AMSC 1 57326, C171504, C162162, Cl 62 1 85, C355267, C355268, C355266, Mt Coricudgy: QMMO44707, M028663, M044714. M044722, M059872, AMSC 162 144. Jenolan Caves: QMM09687, MO 16941. M028778, M037772, AMSC63851, M062446, M035764, M044798, AMSC 124276, Cl 24277, 057334, C162157, 062168, 021717, 054813, C354933, C355270. C411537, 044225. Wombcyan Caves: AMSC3 1 9705. Cl 573 1 7, C 355260, C384164. Abercrombie Caves; QMM 029242 . MO37780, M044766, AMSC157322, 058365, 062165. Orange district: QMMO37490, M039782, AMSC 168647, 068697. SE NSW: QMMO68018, M039986, M039991, M049429, AMSC162159, C354882, 068646, C354917, C354923, C354919, C354921, C’354926, C355263, C355261, C355265, C357298, C357300, C357301, 0 62148, C355264, 062147. DIAGNOSIS. Shell small, golden brown, with darker flammulations. Spire slightly elevated. Teleoconch with crowded, bold slightly sinuate ribs. Mean ribs/mm 10.43. Umbilicus very wide saucer-shaped, mean D/U 2.44. Penis with 4 longitudinal pilasters. DESCRIPTION. Shell very small, golden brown, with darker flammulations (very faint to very dark), with 3. 7-5. 3 (mean 4.5) evenly coiled whorls. Shell diameter 2. 65-4. 27mm (mean 3.35mm). Apex and spire flat. Height of shell 1.30-2. 17mm (mean 1.65mm). H/D 0.44-0.54 (mean 0.49). Protoconch: flat of 1.1 -1.9 whorls. Apical sculpture of bold, regularly spaced slightly curved radial ribs. Ribs bold, medium to wide; height greater than or equal to width; interstices equal to the width of two to three ribs at apex; equal to the width of 4-6 ribs at post-nuclear boundary; ratio of rib spacing (apex to post-nuclear boundary): 0.3-0.5. Microradials with periostracal blades; 1-3 microradials between major ribs at apex; 6-8 microradials between ribs at post-nuclear boundary. Secondary sculpture of low (height less than width) microspiral cords crossing the major radials and forming beads at their intersection with the microradials. Teleoconch with bold prominent, closely and regularly spaced, slightly protectively sinuated radial ribs. Ribs on last whorl 70-176 (mean 109), ribs/mm 7.10-16.05 (10.43). Ribs medium size, straight in section, rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 4-5 ribs; interstices on the penultimate whorl equal to width of 3-5. Microsculpture of microradial ribs and microspiral cords, about equal in strength. Microradials low, 7-11 between ribs on first postnuclear whorl; 7-11 between ribs on penultimate whorl. Microspirals low; crossing the microradials and forming srong beads at their intersection with the microradial ribs; beading strong, always round, or elongate on the first post-nuclear whorl, becoming round towards the aperture. Umbilicus wide saucer-shaped, diameter 0.93-1. 79mm (mean 1.38mm), D/U 2.12-3.55 (mean 2.44). Sutures weakly impressed, whorls rounded above and below a rounded periphery. Aperture narrowly roundly ovate; parietal callus present. Based on 121 measured specimens (AMSC355266, C63851, Cl 62 165, Cl 71504, C355260, QMM039974, M052781, M070402). 290 MEMOIRS OF THE QUEENSLAND MUSEUM NEW CHAROPID LAND SNAILS 291 Ovotestis containing three to seven clumps of alveoli, with more than two alveolar lobes per clump. Talon stalk shorter than talon, or absent; talon circular. Penial retractor muscle half the length of the penis or greater, inserting onto the penis head or at the junction of the penis and epiphallus. Epiphallus present; shorter than or equal to the penis length, wider than the vas deferens; equal in width to penis; internally with two longituduinal pilasters. Penis tubular with 4 longitudinal pilasters. Vagina shorter than or equal length to penis. Pallial cavity with kidney almost unilobed; apex slightly reflexed. Mantle gland absent, or present (present in one specimen; orange, thick). Radula with central tooth weakly tricuspid; lateral teeth tricuspid, with an additional smaller cusp formed between the two ectocones. Ectocone of outer marginal teeth split into three teeth, or split into four or more teeth; endocone not split. Number of lateral teeth: 4-6, marginal teeth: 10-11; radular rows: 90-94. Based on 10 dissected specimens (QMM039974, M03999L MO32063, AMSC354933, C354923). DISTRIBUTION AND HABITAT.From the Kosciusko NP in S NSW to the S New England Region, N NSW but is rare in the Sydney Basin. REMARKS. Scelidoropa sarahjaneae sp. nov. differs from S. nandewar sp. nov. (Nandewar Range) in having a coarser shell sculpture, elevated rather than flat spire and a relatively smaller umbilicus. Marilyniropa jenolanensis gen. et sp. nov. from the Jenolan Caves area bears some superficial resemblance to this species but has more widely spaced ribs, smaller umbilicus and a protoconch without microradial ribs. The wide distribution of S. sarahjaneae broadly mirrors that of some other species discussed in this study. However in contrast to widespread species such as Macrophallikoropa belli (Cox, 1864), Cralopa stroudensis (Cox, 1 864) and Egilomen lirata (Cox, 1 868) this species is barely represented in the Sydney Basin. Reasons for this are not obvious but S. sarahjaneae may have a disliking for sandstone habitats. Scelidoropa nandewar sp. nov. (Figs 10, 24-28, 30-32; Table 5) ETYMOLOGY For the Nandewar Range. MATERIAL. HOLOTYPE: NSW. QMM070403, Nandewar Ra, Mt Dowe (30°16'54"S, 150 o 09’48”E), snow gums, 1440m, under logs and bark, 23.ix.1993, JS, JC. Height of shcll= 1 ,38mm, diameter=3.07mmm, H/D=2.22. D/U=2.21, number of whorls=4.9, ribs on last whorl= 1 89. PARATYPES. NSW: QMM049158, 4SC/5RC, same data as holotype, Nandewar Ra, Mt Dowe (30°16'54"S, 150°09’48"E) t snow gums, 1440m, under logs and bark, 23.ix.1993, JS, JC; QMMO 16743, 3SC/3RC, Dawsons Spring, Mt Kaputar (30°16’S, 150°10’E), 1300m, under moss, 08.xi.1983, JS, DP, PHC, J. Short: QMMO 16754, IRC, Mt Kaputar, summit (30°1 6’S, 1 50°10'E), among rocks, grass and litter, 1 500m, 09.xi. 1 983, JS, DP, PHC, J. Short; QMM049166, 8RC/4SC, Nandewar Ra, c. 1.2km SW Mt Kaputar, summit (30 U I6'36"S, 150°08’48"E), dsf, 1420m, under bark and rubbish, 23.ix.1993, coll JS, JC; QMMO65450, ISC, Nandewar Ra. Mt Kaputar NP, E of Bark Hut Camping Ground (30° 1 7’30"S, 150°08T8"E), dsf, 1180m, under logs, 23.ix.1993, JS, JC. DIAGNOSIS. Small golden brown shell with darker flammulations. Teleoconch with very crowded, bold slightly sinuate radial ribs. Ribs/mm c. 19.60. Umbilicus very wide saucer- shaped, D/U c.2.21. Penis with four longitudinal pilasters. DESCRIPTION. Shell very small, light brown, with darker flammulations, having c.4.9 slightly tightly coiled whorls. Shell diameter c.3.07mm, spire and apex flat. Shell height c. 1.38mm, H/D c.2.22. Protoconch of c.2.0 whorls with apical sculpture of bold, regularly spaced slightly curved radial ribs. Ribs bold, medium to wide; height greater than or equal to width; interstices equal to the width of 2.5 ribs at apex; equal to the width of 3 ribs at post-nuclear boundary; ratio of rib spacing (apex to post-nuclear boundary): 0.3-0.5. Microradials present; 4 between each pair of major ribs. Secondary sculpture of low (height less than width) microspiral cords that FIG. 34. Teleoconch microsculpture. A. Rhophodon palethorpei , QMM070390. B, Rhophodon duplicostatus , QMMO70393. C, Egilodonta baimsdalensis (Yarrangobilly Caves, Kosciusko National Park), AMSC 142959. D, Letomola lanalittleae , QMMO70397. E, Macrophallikoropa depressispira , AMSC205I63. F, Elsothera brazieri , QMM028775. G, Decoriropa lirata (Taree), AMSC 136842. H, Coricudgia wollemiana , AMSC205 164. I. Marilyniropa jenolanensis, AMSC205165. J, Cralopa stroudensis, Sydney. K, Gouldiropa carlessi (Inverell), AMSC3640. L, Sharniropa wollondil/yana , QMMO70399. M, Acheronopa attunga , QMM070400. N, Medley ropa yarrangobi l lye ns is , QMM070401. 0, Scleridoropa sarahjaneae (Abercrombie Caves), AMSC 162 165. P, Diphyoropa saturn i, AMSC 142234. Scale bars=30pm in B and E; 40pm in A, C; 50pm in O; 60pm in D, G, K, L, N, P; 70pm in J; 80pm in H; 100pm in F; 130pm in I, M. 292 MEMOIRS OF THE QUEENSLAND MUSEUM cross the major radials and form beads at their intersection with the microradials. Teleoconch with bold prominent, very crowded and regularly spaced, slightly protractively sinuated radial ribs. Ribs on last whorl c.189, ribs/mm c. 19,60. Ribs medium; in size; straight in section, rounded on top. Rib interstices on the First post-nuclear whorl equal to width of 2-3 ribs; interstices on the penultimate whorl equal to width of 2-3 ribs. Microsculpture of microradial ribs and microspiral cords, about equal in strength. Microradials low, 4-5 between ribs on first postnuclear whorl; 3-4 between ribs on penultimate whorl. Microspirals low; crossing the microradials and forming srong beads at their intersection with the microradial ribs; beading strong, always round. Umbilicus wide saucer-shaped, diameter c. 3. 07mm, D/U c.2.21. Sutures weakly impressed, whorls rounded above and below a rounded periphery. Aperture narrowly roundly ovate; parietal callus present. Based on the measured holotype (QMM070403). Genitalia with ovotestis containing 6 clumps of alveoli, with 2-8 alveolar lobes/clump. Talon stalk very short; talon circular. Penial retractor muscle <1/2 penis length; inserting at junction of the penis and epiphallus. Epiphallus shorter than penis; wider than the vas deferens; equal in width to penis; internally with 2 longituduinal pilasters. Penis tubular with 4 longitudinal pilasters. Vagina equal length to penis. Pallial cavity with kidney almost unilobed; apex reflexed. Mantle gland present (orange in colour). Radula with central tooth weakly tricuspid; lateral teeth tricuspid, with a smaller additional cusp present between the ectocones. Ectocone of outer marginal teeth split into three teeth; endocone not split. Number of lateral teeth: 4, marginal teeth: 11; radular rows: 88. Based on 1 dissected specimen (QMM049158). DISTRIBUTION AND HABITAT. From Mt Kaputar, NSW; wet sclerophyll forest among rocks. REMARKS. Scelidoropa nandewar sp. nov. is similar to the widespread S. sarahjaneae but can be distinguished by the more closely spaced ribs, flatter spire and relatively smaller umbilicus. The geographical restriction of S. nandewar contrasts with that of S. sarahjaneae and highlights the refugial significance of the Nandewar Ranges in land snail evolution. Diphvoropa gen. nov. ETYMOLOGY Greek diphyes, double nature; referring to the dual nature of the protoconch sculpture. TYPE SPECIES. Helix satumi Cox, 1 864 DIAGNOSIS. Very small brown shell with evenly coiled whorls and flat spire. Protoconch sculpture bimodal with the first two-thirds of the protoconch having broad, irregularly spaced spiral lirae that weaken in the last third wher broad, curved radial ridges develop and eventually dominate. Teleoconch sculpture of prominent, almost straight, moderately widely to closely spaced radial ribs. Umbilicus wide to very wide V-shaped. Kidney moderately bilobed. Epiphallus long. Penis tubular up to five longitudinal pilasters. DISTRIBUTION AND HABITAT. Southern Tablelands Bioregion to the Macleay Valley, W of Kempsey, NE NSW. Species live under logs, rocks and leaf litter chiefly in eucalypt forest. REMARKS. Diphyoropa gen. nov. is characterised by its unusual apical sculpture that features an initial prominence of spiral cords which give way to a dominance of curved radial ribs and ridges on the last 1/3 of the protoconch. This protoconch sculpture is unusual in the context of apical sculptures of Australian Charopidae (Iredale, 1937, 1941a,b; Kershaw 1955, 1956a,b; Smith & Kershaw, 1979, 1981; Stanisic, 1990) and as such is diagnostic for the genus. However, considering the many intricate variations on grossly similar apical sculptural themes that have been reported in this study, care needs to be taken in interpreting the condition seen in Diphyoropa as monophyletic. This peculiar apical sculpture may be merely a grade of organisation similar to the reticulate, spiral or radial patterns that have thus far been reported for many charopid species. Undescribed taxa in the collections of the QM from other parts of NSW and SE Qld have grossly similar protoconch sculpture to D. satumi and D. macleay ana. These may or may not belong to this clade. Diphvoropa saturni (Cox, 1864) comb. nov. (Figs 23, 25-28; Table 5) Helix saturni Cox. 1864a: 35; 1868: 14, pl.6, fig. 1 1 . Helix costulata Cox, 1864b: 184; non costulata Lamarck, 1822. Endodonta (Charopa) saturni (Cox). Cox 1909: 69. Egilomen saturni (Cox). Iredale, 1937: 328; Iredale, 1941a: 269. NEW CHAROPID LAND SNAILS 293 Dentherona satumi (Cox). Smith & Kershaw, 1979: 173. Dentherona (Dentherona) satumi (Cox). Smith, 1992: 187. MATERIAL. LECTOTYPE. AMSC205167. Height of shell— 1.18mm, diameter=2.22mm, H/D=0.53, D/U=3.08, number of whorls=4.2, ribs on last whorl=59. PARALECTOTYPE. AMSC 136900, same data as lectotvpe. OTHER MATERIAL. All NSW, Sydney Basin: AMSC8899, Cl 1809, Cl 36900, Cl 54787, Cl 54909, Cl 549 19, Cl 54920, Cl 54921, Cl 54924, Cl 54956, C3 18793, C377420, C3 18807, C3I8811, C3 18908, C318911, C318912, C318913. C318914, C318916, C318918, C3 18926, C3 18931, C3 18932, C3 18933, C3 18934, C3 18935, C3 18941, C3 18942, C3 18943, C3 18944, C3 18946, C3 18947, C319714, C345664, C345675, C345720, C345721. C346756, C346761, C346765, C346767, C346769. C346773, C348735, C354869, C3 18928, C3 18929, C3 18936, C3 18938, C346771, C63819, Cl 54954, QMM028233, M029635, M029639. M029721, MO37970, M034784, M034785, MO34809, M034974, MO37390, M060057, M028224, M042125. Lord Howe I: QMM064I25, M064218, M064133, AMSC204488, C204496, C204581, C397669, C398539, C400485. C389819, C389821, C389824, C389825, C389826, C389827, C389828, C389829, C389830, C389831, C389832, C389833, C389834, C389835, C390217. Vic.: AMSC 1553 19. DIAGNOSIS. Shell very small, with evenly coiled whorls and flat spire. Protoconch sculpture bimodal, with the 1st 2/3 of protoconch having broad, irregularly spaced, spiral lirae weakening in the last 1/3 where broad, curved radial ridges develop and eventually dominate. Teleoconch sculpture of prominent, almost straight, widely spaced radial ribs. Umbilicus very wide V-shaped. Penis with several longitudinal pilasters. Epiphallus about half penis length. DESCRIPTION. Shell very small, light brown in colour with 4. 2-4. 6 (mean 4.33) evenly coiled whorls. Shell diameter 3. 02-3. 59mm (mean 3.26mm). Spire flat. Height of shell 1.18-2.03mm (mean 1.70mm), H/D 0.54-0.57 (mean 0.55). Protoconch of 1.6- 1.7 whorls with sculpture consisting of broad irregularly spaced, spiral lirae for the first two-thirds of the protoconch; weakening in the latter third of the protoconch where broad curved radial ridges develop and gradually become more prominent than the spirals; spirals crossing radial ridges. Ridges wide height less than width; spacing uniform; width of interstices (in multiples of rib width): 2-3. Teleoconch sculpture of bold, almost straight, moderately widely spaced radial ribs; 58-85 (mean 72) ribs on the last whorl, ribs/mm 6.11-7.93 (mean 7.00). Ribs medium in size, height equal to width; straight in section, rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 6-8 ribs; interstices on the penultimate whorl equal to width of 6-7 ribs. Microsculpture of microradial ribs and stronger microspiral cords that cross the microradials and form strong elongate to round beads at their intersection. Microradials low; 9-1 1 between ribs on first postnuclear whorl; 11-12 between ribs on penultimate whorl. Umbilicus very wide V-shaped, diameter 0.72-1. 27mm (mean 1.04mm), D/U 2.82-3.17 (mean 2.9). Sutures weakly impressed, whorls shouldered above and rounded below a rounded periphery. Aperture roundly lunate, parietal callus present. Based on 6 measured specimens (AMSC 142234, C318912, C205167). Genitalia (not figured) with ovotestis containing two clumps of alveoli; with more than two alveolar lobes per clump. Talon stalk equal to talon diameter; talon circular. Penial retractor muscle less than half the length of the penis; inserting onto the penis head. Epiphallus wider than the vas deferens, equal to a half to two thirds the length of the penis. Penis tubular with a prominent tapering distal; internally, upper chamber with several longitudinal pilasters. Pallial cavity (not figured) with kidney moderately bilobed; apex slightly reflexed. Radular morphology unknown. Based on four dissected specimens (QMM028233, M060057, M028244, MO37390). DISTRIBUTION AND HABITAT. Sydney Basin Bioregion but extending to the N South- eastern Tablelands Bioregion (Wombeyan Caves). Presumed introduced populations in Victoria and on Lord Howe Island; under rocks, litter and logs, or discarded household debris in urban situations. REMARKS. Diphyoropci satumi (Cox, 1864) comb. nov. was placed in either Egilomen or Dentherona by various authors (Iredale, 1937; Smith & Kershaw, 1979) without explanation. The protoconch of Egilomen has widely spaced thick radial ribs with weak spiral creases while that of Dentherona (type: Helix ( Charopa) dispar Brazier, 1871) is reticulate with comprising strong radials and somewhat weaker (less raised) spirals. D. satumi is almost exclusively confined to the Sydney Basin where it persists in urban situations. Neither sculptural pattern is considered to be gencrically compatible with the condition in D. satumi. 294 MEMOIRS OF THE QUEENSLAND MUSEUM The almost exclusive occurrence of D. saturni within the Sydney basin suggests strongly that the extralimital records from Victoria and Lord Howe Island are introductions. A single record from a Gloucester garden, N of Barrington Tops (AMSC 154954) could also be a translocation since it is the only record N of the Hunter River in an area where much collecting has taken place in surrounding forests. Diphyoropa saturni has spread prolifically in Lord Howe Island, even outdoing introduced European species such as Cantareus aspersus (Muller, 1774) and Vallonia pulchella (Muller, 1774). A single specimen of D . saturni has been recorded from inner Melbourne, Vic but it could be more prominent in that city and an examination of material in the Museum of Victoria is needed to establish its local abundance more accurately. Populations do not show much conchological variation throughout the natural range ol the species. Locally the species is most likely to be confused with Elsothera brazieri (Cox, 1868) which differs from it primarily in having a protoconch with dominant radial ribs throughout that are crossed by weak, continuous spiral cords. The decision not to figure the anatomy of this species was made to expedite publication. Very little spirit material of this species was available for study, and while this material was of sufficient quality to allow for observation of major anatomical features, it was not suitable for illustration. Diphyoropa macleavana sp. nov. (Figs 10, 24-28, 30-34; Table 5) ETYMOLOGY. For the Macleay Valley. MATERIAL. HOLOTYPE. AMSC205274, Kempsey, WN W, c. 1 .5km E Mt Sebastopol, Kempsey-Carrai Rd, NSW (30°57 , 00 M S, 152 o 28’40"E), drflm. 350m, on rocks in liner, 28.xi. 1 989, JS, DP. Height of shell= 1 .59mm, diameter=2.81mm, H/D=0.57, D/V=3. 35, number of whorls=4. 1 .ribson last whorl=106. PARATYPES. All NSW. QMM056395, IRC, same data as holotvpe; QMMO5601 1, 2RC. Kempsey, WNW, c. 1.5km E Mt Sebastopol, Kempsey-Carrai Rd (3()°57’00"S. 152°28’40"E). drtflm, 350m. in liner. 08.U995, JS. JC; AMSC 168685, 5RC, W of Kempsey, on W side Yessabah Caves limestone outcrop (31.095°S, 152.689 n E), in soil under leaf litter in Lantana infested rainforest, iii. 1 983. MS. OTHER MATERIAL. QMMO37098. M056399, M049382, M055876, M056392, MO60236, AMSC 140247, C140258, C1621S4. DIAGNOSIS. Shell very small, with evenly coiled whorls and flat spire. Protoconch sculpture bimodal with the first 2/3 of the protoconch having broad, irregularly spaced spiral lirae weakening in the last 1/3 where broad, curved radial ridges develop and eventually dominate. Teleoconch sculpture of prominent, almost straight, moderately widely spaced radial ribs. Umbilicus very wide V-shaped. Penis tubular with upper chamber muscular and lower part short, thin, tapering; internally with 5 longitudinal pilasters; epiphallus as long as main section of penis. DESCRIPTION. Shell very small, light brown, with c.4.1 evenly coiled whorls. Shell diameter c.2.8mm. Spire flat to slightly depressed. Height of shell c.2.81mm, H/D c.0.57. Protoconch of c. 1 .8 whorls with bimodal sculpture consisting of broad irregularly spaced, weakly curved, spiral lirae for the first two-thirds of the protoconch; weaking in the latter third of the protoconch where broad curved radial ridges deveolop and become more prominent than the spirals; spirals crossing radial ridges. Ridges wide, height less than width; spacing uniform; width of interstices (in multiples of rib width): 3-6. Teleoconch sculpture of bold, almost straight, closely spaced radial ribs; c. 106 ribs on the last whorl, ribs/mm c.13.4. Ribs medium in size, height equal to width; straight in section; rounded on top. Rib interstices on the first post-nuclear whorl equal to width of 4-5 ribs; interstices on the penultimate whorl equal to width of 3-4 ribs. Microsculpture of microradial ribs and stronger microspiral cords that cross the microradials and form strong elongate to round beads at their intersection. Microradials low; 36 between ribs on first post- nuclear whorl; 48 between ribs on penultimate whorl. Umbilicus very wide V-shaped, diameter c. 0.84mm, D/U c.3.35. Sutures weakly impressed, whorls shouldered above and rounded below a rounded periphery. Aperture roundly lunate, parietal callus present. Based on the measured holotype. Genitalia with epiphallus equal to penis length. Penis tubular internally with 2 longitudinal pilasters. Pallial cavity with kidney moderately bilobed; apex slightly reflexed. Radula with central tooth strongly tricuspid, only slightly smaller than first lateral tooth; lateral teeth tricuspid. Mesoconc of lateral teeth slender, slightly diamond-shaped. Marginal teeth tricuspid; ectocone of outer marginal teeth not split; endocone not split. Number of lateral teeth: 8; marginal teeth: 5; radular rows: 98. Based on 2 dissected specimens (AMSC 1 62 1 84). NEW CHAROPID LAND SNAILS 295 TABLE 5. Shell measurements for species of Acheronopa , Hedleyropa , Scleridoropa and Diphyoropa. (Abbreviations as in Table 1). Measurement Acheronopa attunga Hedleyropa yarrangobillyensis Scleridoropa euryscopa Diphyoropa saturni Diphyoropa macleayana 23 20(*19; **17) 119 5 * r D 3.92-6.28 (4.654) ± 0.468 3.21-4.52(3.82) ± 0.354 2.65-4.27 (3.35) ± 0.32 3.02-3.59 (3.264) ± 0.274 2.81 PW 1.5-1. 7 (1.6) ± 0.067 1.6-1.9(1.671) ± 0.077** 1.1-1.9(1.71) ± 0.09 1.6-1.7(1.66) ± 0.055 1.8 TW 3-3.6 (3.37) ± 0.177 2.3-3. 1 (2.806) ± 0.205 ** 2-3.6 (2.78) ± 0.27 2.5-2. 9 (2.7) ± 0.187 2.3 NW 4.6-5.2 (4.97) ± 0.172 4. 2-4.7 (4.476) ± 0.156** 3.7-5.3(4.48) ± 0.26 4. 2-4.6 (4.36) ± 0.219 4.1 H 1.59-2.11 (1.845) ± 0.159 1.63-2.51 (1.981) ± 0.209 1.3-2.17(1.65) ± 0.18 1.62-2.03 (1.806) ± 0.182 1.59 SP 0.11-0.41 (0.263) ± 0.078 0.12-0.41 (0.208) ± 0.076 0-0.32 (0.11) ± 0.07 0-0.01 (0.002) ± 0.004 - AH/AW 0.65-0.88 (0.74) ± 0.05 0.83-1.07 (0.96) ± 0.07 1.09-1.45(1.26) ± 0.07 1.18-1.44(1.3) ± 0.11 1.23 UMB 1.22-1.71 (1.477) ± 0.142 0.7- 1.2 (0.896) ± 0.125 0.93-1.79(1.38) ± 0.18 0.98-1.27(1.102) ± 0.145 0.84 RIB 20-31 (25) ± 3.162 86-136(108.9) ± 13.408 * 70-176(109.29) ± 21.34 58-85 (74.2) ± 11.032 106 RIBS/MM 1.216-2.118(1.721) ± 0.238 8.170-10.72 (9.156) ± 0.736 * 7.102-16.05(10.43) ± 2.082 6.113-7.933 (7.216) ± 0.699 13.4 H/D 0.3-0.48 (0.4) ± 0.04 0.48-0.58 (0.52) ± 0.03 0.44-0.54 (0.49) ± 0.02 0.54-0.57 (0.55) ± 0.01 0.57 D/U 2.89-4.21 (3.16) ± 0.27 3.72-5.05 (4.3) ± 0.36 2.12-3.55 (2.44) ± 0.18 2.82-3.17(2.98) ± 0.16 3.35 DISTRIBUTION AND HABITAT. Macleay River Valley, NE NSW; living in litter in a range of vegetation types including eucalypt woodland and dry rainforest on limestone. Closer to the coast D. macleayana occurs in eucalypt forest and littoral rainforest. REMARKS. Diphyoropa macleayana sp. nov. is smaller than D. saturni and has more crowded ribs on the teleoconch. However a key difference exists in the reproductive anatomy in which the epiphallus of D. macleayana is as long as the penis compared with an epiphallus half to two-thirds the length of the penis in D. saturni . D. saturni also has a distal tapering extension ot the penis which is not seen in D. macleayana. Elongation of both the penis and in particular the epiphallus together with the bimodal protoconch sculpture are key departures from the more typical patterns seen in eastern Australian charopids hitherto invesitgated. An analogous situation of combined conchological/anatomical character shifts was reported for Rotacharopa Stanisic, 1990 by Stanisic (1990). However, with so much of the charopid fauna yet to be studied, it is difficult to gain a true phylogenetic perspective of the significance of these features. PHYLOGENETIC ANALYSIS The characters and character states (Table 6) forming the data matrix (Table 7) lead to the strict consensus tree of the 250 most parsimonious trees (length 127, consistency index 0.56, retention index 0.65) (Fig. 35), with bootstrap values shown for each clade. In the strict consensus tree, the Rhophodon , Letomola , Elsothera and Scleridoropa clades were well supported, with bootstrap values of 90% or greater. Macrophallikoropa was also moderately well supported (bootstrap value of 80%). These groups were therefore accepted as genera. Most other clades in the tree were poorly supported. The remaining taxa were either placed into monotypic genera, or united in genera with taxa not included in the analysis (based on shell characters). Two exceptions to this are Cralopa and Egilodonta. Cralopa was not well supported by bootstrap values. One species ( carlessi ) was removed from this group into a new genus; however, the other two taxa were retained in Cralopa. There are at least two undescribed species that appear to be closely related to C. kaputarensis (JS, pers. obs.), and a study of these, 296 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 6. List of characters and character states used in the phylogenetic analysis. Character number Character States 1 Shell colour 1 : absent, 2: flecked. 3: flammulated, 4: plain 2 Coiling pattern 1: loose, 2: even, 3: tight. 3 Whorl profile above periphery 1: flattened, 2: rounded, 3: shouldered 4 Supraperipheral sulcus 1: absent. 2: present 5 Apertural sinus 1: absent, 2: shallow. 3: deep 6 Spire height 1 : apex and early spire elevated, 2: apex depressed and early spire elevated, 3: apex and early spire flattened. 4: apex and early spire depressed 7 Protoconch shape 1: exsert, 2: flattened 8 Umbilicus shape 1 : cup or saucer shaped, 2: V-shaped, 3: U-shaped, 4: closed 9 D/U ratio 1: less than 3, 2: 3-4, 3: 4-4.5, 4: more than 4.5 10 Protoconch radial ribs 1 : absent, 2: present 11 Protoconch - regular radial ribs 1: absent, 2: present 12 Protoconch microradials 1 : absent. 2: present 13 Protoconch spirals 1 : absent, 2: present 14 Protoconch pits 1: absent, 2: present 15 Protoconch radial ribs - spacing 1: evenly spaced. 2: increasing towards teleoconch, 3: decreasing towards teleoconch 16 Protoconch radial ribs - number 1 : 45 or fewer. 2: 46-65. 3: 66-85, 4: 85 or more 17 Protoconch spirals - form 1: wrinkles. 2: threads, 3: lirae 18 Protoconch spirals - crossing radial ribs 1: spirals do not cross radial ribs, 2: spirals cross radial ribs 19 Teleoconch radial ribs 1 : absent, 2: present 20 Tcleoconch rib count 1: low (69 or fewer), 2: medium (70-149), high (149 or more) 21 Teleoconch radial rib width 1 : narrow, 2: medium, 3: wide 22 Teleoconch rib curvature strength 1 : strong. 2: weak 23 Teleoconch rib sinuation 1: convex, 2: concave, 3: straight 24 Kidney 1 : almost unilobcd. 2: moderately bilobed, 3: strongly bilobed 25 Secondary ureter 1: closed, 2: open 26 Epiphallus and vas deferens junction 1 : bulbous, 2: no-bulbous 27 Epiphallus coiling 1: simple, 2: complex 28 Penis pilasters 1: simple longitudinal. 2: circular, 3: absent 29 Number of longitudinal pilasters 1 : 1 pilaster, 2: 2 pilasters. 3: 5 or more pilasters, 4: 3-4 pilasters 30 Vergic structure 1 : absent, 2: absent 31 Ovotestis - general shape 1 : many follicles in line along duct; 2: few follicles arranged in clumps (usually 2) 32 Ovotestis lobes 1 : 1 lobe, 2: 2 lobes, 3: more than 2 lobes 33 Preputial tube 1 : absent, 2: absent 34 Radula - lateral tooth structure 1 : continuous, 2: discontinuous 35 Radula central tooth structure 1 : tricuspid, 2: almost unicuspid 36 Radula - marginal teeth 1: saw-like (Rectangular). 2: similar in shape to laterals 37 Apertural barriers - microdentition 1 : triangular. 2: rectangular 38 Parietal barrier A 1 : absent, 2: present 39 Parietal barrier B 1 : absent, 2: present 40 Parietal barrier C 1: absent, 2: present 41 Palatal barrier A 1 : absent. 2: present 42 Palatal barrier C 1 : absent, 2: present 43 Palatal barrier E 1 : absent, 2: present 44 Basal barrier A 1: absent, 2: present NEW CHAROPID LAND SNAILS 297 TABLE 7. Data matrix for the phylogenetic analysis. when more material becomes available, may show that the latter should be in a separate genus, leaving Cralopa as a monotypic taxon. Egilodonta had a bootstrap value of 46%. There was greater support for the group [ Egilodonta + Rhophodott], which had a bootstrap value of 74%. These groups have been united before (Stanisic, 1990). However, the 4 Egilodonta taxa are united by characters such as wide rib spacing on the teleoconch and a penial 298 MEMOIRS OF THE QUEENSLAND MUSEUM Minidonta hendersoni Aaadonta constricta constricta Rhophodon kempseyensis Rhophodon palethorpis Egilodonta bairnsdalensis Egilodonta wyanbenensis Egilodonta bendethera Egilodonta paucidentata Letomola contortus Letomola lanahttlae Macrophallikoropa belli Macrophallikoropa stenoumbilicata Scleridoropa sarahjaneae Scleridoropa nandewar Elsothera brazieri Elsothera sericatula Diphyoropa satumi Whiteheadia globosa Coricudgia wollemiana Cralopa stroudensis Cralopa kaputarensis Gouldiropa carlessi Sharniropa wollondillyana Hedleyropa yarrangobillyensis Decoriropa lirata Marilyniropa jenolanensis Acheronopa attunga FIG. 35. Strict consensus tree of 27 taxa comprised of the 250 most parsimonious trees (length 127, consistency index 0.56, retention index 0.65). Bootstrap values are marked at the base of each clade. verge, and are widely separated geographically from the other Rhophodon taxa. For these reasons, Egilodonta is established as a separate genus. This study contains only a small cross-section of the Australian charopid fauna, and some of the low branch support may be due to the fact that key taxa are missing from the analysis. Until more Australian charopid taxa are described or thoroughly revised, many relationships within the group will remain unresolved. DISCUSSION The study of the eastern Australian Charopidae is still in its infancy and the descriptive task that faces researchers cannot be overstated. Stanisic (1990) analysed conchological and anatomical variation in 50 species of eastern Australian Charopidae and attempted to place these into regional and local context. In doing so the in- adequacy of many previous species descriptions and generic diagnoses based solely on comparatively gross shell charcters (Iredale, 1937, 1 94 1 a,b; Smith & Kershaw, 1979, 1981) was exposed. Only the studies of Solem (1984) [on two northern Australian species] and Smith & Kershaw (1985) [on three Tasmanian species] were considered of sufficient standard to enable detailed com- parisons to be undertaken with species extralimital to that study. The current study has attempted to build on the work of Stanisic (1990) and consequently has resulted in a change in some of the taxonomic concepts developed in the earlier study. The species included in the current study were specifically selected on the basis that the species were generally well differentiated on shell features. As a result, they comprise only a limited range of morphotypes among a very large east-coast fauna. Remarks on the relative significance of patterns of morphological variation need to be tempered by the knowledge that additional "unusual’ forms will emerge as this fauna is investigated. This is especially so in regard to the many species occurring on the high mountains of the Wet Tropics, NE Qld (Stanisic et al., 1994b) which display a range of shell variation greater than seen in the remainder of the east coast fauna (Stanisic, 1993a; unpubl. data). Intergeneric variation is discussed under the relevant generic treatments but inferred relationships between genera are still largely tentative. In summary, it would appear that quite different lineages appear to have developed grossly similar patterns of shell form (size, shape and sculpture) probably in response to environ- mental selection. Living among seasonally dry, limestone talus would seem to favour species with dentate apertures and those with bold postnuclear sculpture. Anatomical variation was largely conservative with little in the way of identifiable character shifts. Where such shifts did occur (. Macrophallikoropa , Diphyoropa) these coincided with major shifts in shell NEW CHAROPID LAND SNAILS 299 patterns, particularly protoconch sculpture. Subtle differences in shell features such as fine structure of protoconch sculpture, architecture of post-embryonic sculpture, whorl profile and coiling pattern provided a means of discrim- inating between the grossly convergent patterns and in identifying significant generic level trends. Most useful was the difference in protoconch sculpture. A casual review of the generic taxa previously reviewed by Stanisic (1990, 1993a, b; 1996) shows that in nearly all cases, this feature is generically diagnostic. In two exceptions, Cralopa and Egilomen , greater attention to differences in protoconch sculpture would have avoided some of the problems rectified in this study. It would appear that this single feature will be a key guide to generic level classification within this family but needs to be interpreted in microscopic rather than macroscopic detail. Hence, terms such as radial or spiral are in themselves of limited use. BIOGEOGRAPHY The Charopidae are a Gondwanan group (Stanisic, 1998a) which has historically been regared as having greatest diversity in the moist forests of southeastern Australia. However, significant charopid faunas have been reported from the high mountains of the Queensland Wet Tropics (Stanisic et al., 1994) and the semi-arid Brigalow Lands of Qld (Stanisic, 1 998b). In each of these cases though, the charopids are associated with rainforest refugia reinforcing the contention of Stanisic (1990) that the association of charopids and rainforests in eastern Australia has had a long history, probably extending beyond the Miocene. Our knowledge of rainforest evolution since the Miocene (A.P. Kershaw, 1981: Webb & Tracey, 1981; Kershaw' & Nix, 1988; Nix, 1991) suggests that rainforest has been contracting in the face of increased continental aridity. Rainforests were more widespread when Australia was wetter, possibly extending into the interior of the continent (Martin, 1981), and hence it is reasonable to speculate that the charopid fauna would have been more widely distributed. Martin (1987, 1997) has shown that rainforest was present in the inland Lachlan and Darling Rivers as recently as the early to mid-Pliocene. As climatic conditions on the Australian continent shifted to a drier regime, rainforest receded into moist refugia and the overall range of the Charopidae would have contracted. At the individual species level this climate-induced restriction of rainforest would have led to the fragmentation of populations in the first instance and extinctions in many cases. The surviving Charopidae are now found mainly in suitable refugia. This longterm isolation of such fragmented charopid populations would have provided numerous opportunities for allopatric speciation. Subsequent dispersal and recolonisation may have occurred as climatic conditions (and hence vegetation systems) fluctuated between w'et and dry. Allopatric speciation in the Charopidae has been documented in the larger rainforest massifs (Stanisic 1987, 1993a,b). But the phenomenon should be even more pronounced in areas where small patches of rainforest were separated by the rapidity developing, drier eucalypt communities. Hitherto there has been little published evidence to show the effects of climatic change on the distribution of snails that exist in the ‘drier’ areas of eastern Australia. Rainforest is used here in a broad sense to include dry vine thickets and scattered remnants sometimes represented by only a few key fioristic elements. In semiarid areas rainforest survives chiefly on rocky outcrops, in gully heads and along watercourses mainly because these habitats provide a more stable moisture supply than the surrounding countryside. Rock outcrops in particular are important microhabitats for both rainforest plants and land snails. These ‘rock piles' may consist of basalt talus, granite boulders or limestone karst, and preserve environmental moisture in the crevices between the rocks They also provide important shelter from fire. Limestone outcrops are particularly significant for land snails in eastern Australia (Stanisic, 1 997 a,b). Besides providing shelter and moisture, the limestone also provides a ready supply of calcium for shell production. Collecting has shown that in the majority of cases these outcrops are snail-rich ‘islands’ in a sea of otherwise snail-poor countryside. They often support quite distinctive land snail communities of which examples have been highlighted by Stanisic (1997a,b). The rainforest on many of these outcrops may be the result of recent opportunistic seeding but in others the association may have greater continuity. Such is probably the case" in the Macleay Valley where rainforest, besides growing on the limestone outcrops, is also a dominant fioristic feature of the surrounding countryside. And, probably not through mere coincidence, here there is a proliferation of land 300 MEMOIRS OF THE QUEENSLAND MUSEUM snails in numbers not seen elsewhere in eastern Australia (Stanisic, 1997a). Not all limestone outcrops have as spectacular faunas as those in the Macleay Valley, but they are distinctive nonetheless. In some cases, less than half a dozen species may be present, but they may still include restricted endemics. Because this study covers a very small component of a much larger and undescribed charopid fauna, biogeographic hypotheses emerging from this work remain limited. Never- theless one fact is evident from this studyand that is that limestone outcrops have been important historical refugia for the Charopidae in E NSW. They have been of primary importance where some genera have relied on this archipelago of habitats for survival. This appears to be true for Egilodonta , Letomola , Achenoropa. Shantiropa and Marilyniropa. In both Letomola and Sham iropa a 1 1 opa tri c speciation following frag- mentation of a wider ancestral population that probably encompassed non-limestone areas is indicated. Isolation of species has probably been longterm. In contrast Achenoropa attunga and Marilyniropa jenolanensis are localised limestone endemics, in each case scattered across several geographically proximate, but none- theless isolated limestone outcrops. They may be recently fragmented populations of limestone dwellers that have yet undergone speciation. In contrast, Egilodonta exemplifies the principle of population fragmentation followed by allopatric speciation. In other groups the limestone outcrops appear to have formed a small but nonetheless integral part of a wider network of mesic refugia in which these tiny litter snails were able to withstand the vagaries of a drying landscape. Such is the case with Macrophallikoropa , Scelidoropa and Rhophodon where species also occur outside the limestone outcrop archipelago. Macro- phalli - koropa belli has the most widespread distribution of the species investigated herein, yet has restricted sister species on the Jenolan Caves limestones and on the volcanically capped Mt Coricudgy. The biogeo- graphic importance of the Jenolan outcrops is further highlighted by the presence of the restricted Marilvnimpa jenolanensis. The widespread distributions of such species as Decohropa l i rata. Macrophallikoropa belli and Scelidoropa sarahjaneae across the limestone archipelago suggests that the the adaptability of individual species may also have been influential in the determination of some of the distribution patterns discussed herein. At a regional level, general trends are few. Perhaps such trends will become more obvious when additional species are studied. The absence of the otherwise widespread Scelidoropa sarahjanae from much of the Sydney Basin (in contrast to situation with D.lirata , Cralopa stroudensis and M. belli) is unusual and needs to be further investigated. Secondary areas of biogeographic significance appear to be volcanic peaks such as Mt Coricudgy (Macrophallikoropa stenoumbilicata , Coricudgia wollemiana ), Mt Kaputar (Scelidoropa nandewar , Cralopa kaputarensis) and Glenugie Peak (Rhophodon duplicostatus). Again these refugia preserve mesic communities chiefly because they are largely rocky outcrops that conserve moisture. The lack of many major radiations within this study is probably a function of the choice of species chosen for study whereby highly differentiated forms were selected. However, it probably also reflects the relictual nature of this fauna. Many charopids have quite localised distribututions and opportunistic survival in such small refugia appears to have been relatively commonplace. At the same time there would have been many instances of extinction. CONSERVATION Some of the larger limestone outcrops (e.g. Jenolan, Wombeyan) mentioned in this study have been afforded legislative protection because of their value as caving sites and this security has provided an inadvertent safe haven for the resident biota. But many lesser known outcrops have no formal protection at all and the plight of their fauna, including the unusual charopids, is uncertain. As this study has shown, many of the limestone-associated charopids are localised endemics, often restricted to single outcrops. A conservation strategy that aims to preserve this fauna by listing all limestone outcrops on the National Estate should be considered. ACKNOWLEDGEMENTS This project was supported in large part by a grant from the taxonomy project of the NSW Biodiversity Strategy, arranged through the Australian Museum, and was supervised by Dr Winston Ponder. One of us (JS) is also grateful to the Australian Biological Resources Study for funding survey work on the New South Wales NEW CHAROPID LAND SNAILS 301 limestones in 1992-1994. We thank Kylie Stumkat for completing the initial scanning electron micrographs. These provided the impetus for the successful completion of this project. We also thank Darryl Potter, QM and Ian Loch, AM for their help in supplying material for study. Thanks are also due to Sham Rose for completing all the anatomical drawings, to Alison Miller for producing the maps, to Winston Ponder for assistance with the cladistic analysis and to Andrew Roberts, who contributed to the project through the Immortals Program at the Australian Museum. LITERATURE CITED ADAM, P. 1992. Australian rainforests. (Clarendon Press: Oxford). BELBIN. L. 1994. PATN: pattern analysis package. (CSIRO Division of Wildlife and Ecology: Canberra). BURCH, J.B. 1976a. Mollusca. Pp. 109-124. In Broadbent, J. & Clark, S. (eds) A faunal survey of east Australian rainforests. Interim Report. (Australian Museum: Sydney). 1976b. Outline the classification of Australian terrestrial molluscs (native and introduced) Journal of the Malacological Society of Australia 3: 127-156. COX, J.C. 1866. Description d'espcces nouvelles provevant d' Australie ct des iles Solomon et Norfolk. Journal de Conchyliologic 14: 45-48. 1868. A monograph of Australian land shells. 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A survey of the land snails of the Queensland Brigalow Lands: their distribution, conservation status and close association with vine thickets. Unpubl. report for the Queensland Deparment of Environment. (Queensland Museum: Brisbane). STANISIC, J., EDDIE, C., HILL, A. & POTTER, D. 1994. A preliminary report on the distribution of land snails occurring within the Wet Tropics area. Unpubl. report for the Wet Tropics Management Authority. Queensland Museum: Brisbane). SWOFFORD, D.L. 2001. PAUP*: Phylogenetic Analysis Using Parsimony. (Sinauer Associates, Inc.: Sunderland, Massachusetts). WEBB, L.J. & TRACEY, J.G. 1981. Australian rainforests: patterns and change. Pp. 605-694. In Keast, A. (cd.) Ecological biogeography of Australia. (W. Junk: The Hague). A NEW CANDONINAE GENUS (CRUSTACEA: OSTRACODA) FROM SUBTERRANEAN WATERS OF QUEENSLAND, WITH A CLADISTIC ANALYSIS OF THE TRIBE CANDONOPSINI I VAN A KARANOVIC Karanovic, I. 2005 01 10: A new Candoninae genus (Crustacea: Ostracoda) from subterranean waters of Queensland, with a cladistic analysis of the tribe Candonopsini. Memoirs of the Queensland Museum 50(2): 303-319. Brisbane. ISSN 0079-8835. Pioneercandonopsis hancocki gen. nov., sp. nov. was collected from subterranean waters of Pioneer Valley, Queensland, and it is the first subterranean ostracod from this state. The new genus belongs to the tribe Candonopsini Karanovic, in press b, which also includes Candonopsis Vavra, 1891, Caribecandona Broodbakker, 1983. and Cubacandona Broodbakker, 1983. Characteristics of the hemipenis and prehensile palps suggest that Pioneercandonopsis is more closely related to the latter two genera than to Candonopsis. A cladistic analysis based on two sets of morphological characters was performed on all 32 species of the tribe Candonopsini, with Cryptocandona dudichi (Klie, 1930) chosen as outgroup taxon. Resulting cladograms support a monophyletic origin for the Candonopsini and close phylogenetic relationship between the new genus and Caribecandona and Cubacandona. □ Pioneercandonopsis, new genus, Candonopsini, Candoninae, cladistic analysis. Ivana Karanovic. Western Australian Museum, Francis Street, Perth 6000 WA, Australia (e-mail: karanovic@museum.wa.gov.au); 13 A total of 19 freshwater free-living ostracods are known from Queensland (Sars, 1889; De Deckker, 1978, 1979, 1981 ,1983; DeDeckker& McKenzie, 1981) and all are surface water inhabitants. The subterranean ostracod fauna of Queensland is completely unexplored. Recently, the Queensland Department of Natural Resources and Mines started an extensive sampling of the subterranean waters of Pioneer Valley. I had the opportunity to examine these samples and discovered the new ostracod described in this paper. This is the first sub- terranean ostracod recorded from Queensland and is also the 10th ostracod genus described from Australian subterranean waters. The other 9 genera are from groundwaters of Western Australia (Karanovic & Marmonier, 2003; Karanovic, 2003a, b, in press a, b). All 10 genera belong to the Candoninae Kaufmann, 1900. This subfamily has 1 1 extant genera in Australia (10 endemic, plus Candonopsis Vavra, 1891), with an additional 20 genera worldwide. Initially, the Candoninae was divided into tribes Namibcypridini Martens, 1992 with two genera, and Candonini Kaufamnn, 1900 containing 16genera (Martens, 1992). However, discovery of a highly diverse Candoninae fauna in WA, with striking morphological features, necessitated revision of the Candoninae with the aim of better understanding the phylogenetic September 2004. relationships between genera. Karanovic (in press a, b) described the Humphreyscandonini Karanovic, in press a, and Candonopsini Karanovic, in press b. The former tribe includes 7 genera known only from subterranean waters of the Pilbara region (WA). while the Candonopsini encompasses Candonopsis , Caribecandona Broodbakker, 1983, and Cubacandona Broodbakker, 1983. The new genus from Queensland is assigned to this tribe. The Candonopsini contains 32 Recent species. Candonopsis is the largest, with 24 species, and is divided (Karanovic, in press b) into Candonopsis s. str. and Abcandonopsis Karanovic, in press b. The latter subgenus is an Australian endemic with 3 subterranean species: 2 from Kimberley and one from the Murchison Region (Karanovic, in press b). Candonopsis s. str. has 5 species in Australia, 4 endemic to subterranean waters of the Murchison Region (Karanovic & Marmonier, 2002). The remaining Candonopsis species are in Europe, South America and Africa. Cubacandona and Caribecandona are less diverse and have more restricted distributions. Cubacandona was represented only by the type species, C. cubensis (Danielopol, 1978), from Cuba (Danielopol, 1978, 1980), but Karanovic (in press b) assigned Candonopsis species from South America to that genus. Al l 3 species of Caribecandona are found 304 MEMOIRS OF THE QUEENSLAND MUSEUM in subterranean waters of the West Indies (Broodbakker, 1983). Cubacandona and Caribecandona share 3 very important morphological characteristics: 2 extensions of the lateral lobe of hemipenis, almost symmetrical prehensile palps and a very short terminal segment of the mandibular palp. In contrast, Candonopsis has only I extension of the lateral lobe, extremely asymmetrical prehensile palps and a long terminal segment of the mandibular palp. With respect to these characteristics, the new genus is more closely related to the Central and South American genera than it is to Candonopsis , but shares several setal reductions with the subgenus Abcandonopsis and some other Candonopsis s. stn species. To test the phylogenetic relationships in the Candonopsini, a cladistic analysis was performed. This analysis was based on species-level taxonomy, because this is more likely to represent evolutionary relationships (Walker-Smith & Poore, 2001 ) and included all 32 species of the tribe. MATERIAL AND METHODS TAXONOMIC METHODS. Samples were collected with haul-nets (mesh size 250 or 350 pm) from bores. Haul-nets are actually simple plankton nets of a different size suitable for the bore, which can range from 30-180 mm in diameter. Weighed nets were lowered down into the bore with one bottle screwed on its distal part then hauled through the water column, usually a number of times. Animals were stained with the ‘Rose Bengal 7 dye, for easier sorting. For observation under the light microscope, ostracods were dissected in a mixture of distilled water and glycerol (1:1) with fine entomological needles (mark 000). Dissected appendages were mounted in Faure’s medium and observed using a Leica DMLS brightfield compound microscope with C-plan achromatic objectives. All drawings were prepared using a drawing tube attachment on the same microscope. For the Scanning Electron Microscopy (SEM), soft parts were taken from their valves and then dehydrated in a graded series of ethanol: 70%, 80%, 95% and 100%. In each of the grades, soft parts were left for about 1 5 minutes. The carapace was kept in the 100% ethanol for only a few minutes. After that, the soft parts and carapace were mounted on a SEM stub using double-sided adhesive tape. All samples were sputter coated with gold and observed under the LEO FEG VPSEM micro- scope with in-lens detectors, working distance between 2-6mm, and accelerating voltage between 2-5kV. In the systematics, the length of all segments was measured along the midline, and length ratios are presented beginning with the proximal segment. Appendage terminology follows Martens (1998), while chaetotaxy of all limbs follows Broodbakker & Danielopol (1982), revised for the antenna by Martens (1987). i follow Martens’ (1987) terminology except that the shortest claw on the male terminal segment is considered homologous with the female GM claw, while the longest one is homologous with the female Gm claw. Setae on the third thoracopod are labeled according to Meisch (1996), while setae on the antennula are labeled according to Karanovic (in press c). Lobes on the hemipenis are designated according to Danielopol (1969). In front of the abbreviations tor the setae on the endopodal segments of the 3rd thoracopod and setae on the 1st thoracopod, the letter ‘T’ was added so that they are not confused with abbreviations for the hemipenis given below. ABBREVIATIONS, a - lateral lobe on hemipenis; A1 - antennula; A2 - antenna; CBl, CB2 - setae on the antennular coxobasis; b - medial lobe on hemipenis; dl, d2, dp - setae on the basal segment of the third thoracopod; Enl-6 - endopodal segments of the antennula; En a, b, c, d - endopodal setae on the antennula; Exl, 2 - exopodal setae on the antennula; Fu - furca; Gl, G2, G3, GM, Gm - antennal claws; H - height; L - length; LV - left valve; Md - mandibula; Mxl -maxillula; QM - Queensland Museum; RV . right valve; Ta, Ta\ Tb, Td - setae on the first thoracopod; Te, Tf, Tg. Th 1 , Th2, Th3 - setae on endopodal segments of the third thoracopod; Tl, T2, T3 - first, second and third thoracopods; 1 1 , t2 - medial setae on the second endopodal segment of the antenna; W - width; WAM - Western Australian Museum; Y, ya, yl, y2, y3 - aesthetascs; z 1 , z2, z3 - apical setae on the second endopodal segment of the antenna. CLADISTIC METHODS. A total of 32 species of the Candonopsini are included in the cladistic analysis, while Ctyptocandona dudichi (Klie, 1930) was chosen as an outgroup taxon. According to Ba I tanas et al. (2000), Ctyptocandona Kaufmann, 1900 represents a primitive lineage in the Candoninae. All known species of Cryptocandona have the highest number of setae found in the Candoninae on the Al, A2, and all thoracopods, which makes them A NEW OSTRACOD GENUS FROM QUEENSLAND 305 closely related to the Candonopsini. However, the characteristics of Md, the prehensile palps, furca and hemipenis exclude Cryptocandona from the Candonopsini and the genus is placed in the Candonini (Martens, 1992). Cryptocandona dudichi was chosen because it was recently redescribed in detail (Namiotko et al., 2001 ) and both sexes are known. The morphological characters used in this analysis were selected from information in the literature. Characters were polarised so that the outgroup taxon values all remained zero. This gives the least increase in the length of the overall cladogram (Kitching et al., 1998). The ‘0’ state also means a plesiomorphic character state, while 4 1 ’ means an apomorphy. Unknown or missing values are coded Only the carapace shape and the total number of the Al segments were multistate characters (with states 0, 1 , and 2); all other characters used in this analysis were binary. Carapace shape was coded as a non-additive (representing unordered multistate character as a linked series of binary character) character, while the total number of segments on A 1 was coded as additive (representing ordered multistate character as a linked scries of binary characters). Two data matrices were created, characters coded, optimised and weighted using the computer program WinClada, version 1.00.08 (Nixon, 2002), and then analyzed using NONA, version 2 (Goloboff, 1999). All 24 characters from the first data matrix were analyzed as equally weighted, using the heuristic and the Rachet Island Hopper searches with the WinClada default parameters. For the heuristic search the default parameters are: hold 100; mult*l; hold/1 (hold 100 trees in memory, perform tree bisection and reconstruction (TRB) branch swapping on 1 random additional replicates and hold 1 starting tree in memory). For the Rachet Island Hopper search the default parameters are: 200 replications; 1 tree to hold; 3 characters to sample; 1 0 random constraint level and amb-poly= ( amlv collapses a branch if the ancestor and descendant have different states under same resolutions of multistate characters or of poly=treats trees as collapsed). Characters in the second data matrix were weighted so that the highest weight ( 1 ) was given to the characters from 0-15. Those were characters of the tribe, genera, subgenera, and some characters of the species groups. Characters 16-27 were weighted 0.75, majority of these are homoplastic. The lowest value (0.5) was given to the characters from 28-33, i.e. to the carapace shape and to the almost all ambiguous characters. When analysing with the Rachet Island search the same default parameters were used, while for the Heuristic search the following options were chosen: hold 10000; mult* 100; hold/10. On all trees obtained unsupported branches were collapsed and characters were optimised using the fast optimisation option in WinClada. REFERENCE SOURCES FOR THE OUTGROUP AND INGROUP TAXA (indicates original paper is a reference source). 1. Cryptocandona dudichi (KJie, 1930): Namiotko et al. (2001); 2. Candonopsis (Abcandonopsis) aula Karanovic, in press b*; 3. C. (A.) indoles Karanovic, in press b*; 4. C. (A.) williami Karanovic & Marmonier, 2002*: Karanovic (in press b); 5. C. (Candonopsis) africana Klie, 1944*: Rome (1962); Martens (1984); 6. C (C.) anteroarcuata Rome, 1962*; 7. C. (C.) boui Danielopol, 1978*: Danielopol (1980); 8. C. (C.) bujukuensis Loftier, 1968* 9. C. (C.) dani Karanovic & Marmonier, 2002*; 10. C. (C.) hummelincki Broodbakkcr, 1983*; 11. C. (C.) kingsleii (Brady and Roberston, 1870): Petkovski (1977) 12. C. (C.) kimberleyi Karanovic &Marmonier, 2002*; 13. C. (C.) mareza karanovic &Petkovski, 1999*; 14. C. (C.) murchisoni Karanovic &Marmonier, 2002 *; 15 . C. (C.) nama Daday, 1913*; 16. C. (C.) navicula Daday, 1910*: Klie (1935); Rome (1962); Karanovic (in press b); 17. C. (C.) putealis Klie, 1932*; 18. C. (C.) scourfieldi Brady, 1910: Petkovski & Meisch (1995); 19. C. (C.) solitaria Vavra, 1895*: Klie (1936) 20. C. (C.) sumatrana Klie, 1932*: Harding (1962); Victor & Fernando (1978); Karanovic (in press b); 21. C. (C.) tenuis (Brady, 1886)*: Sars (1896); Karanovic and Marmonier (2002); 22. C. (C.) thienemanni Schafer, 1945*; 23. C. (C.) trichota Schafer, 1945*; 24. C. (C.) urmilae Gupta, 1988*; 25. C. (C.) westaustraliensis Karanovic & Marmonier, 2002 *; 26. Cubacandona cubensis (Danielopol, 1978*): Danielopol (1980); Broodbakkcr (1983); 27. C. anisitsi (Daday, 1905)*: Klie (1930); 28. C. columbiensis (Mehes, 1913)*; 29. C. falclandica (Vavra, 1898)*; 30. Caribecandona trapezoidea Broodbakker, 1983*; 31. C. auricularia Broodbakker, 1983*; 32. C. ansa Broodbakkcr, 1983*. 306 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 1. Pioneercandonopsis hancocki gen. et sp. nov. A, G, H, Holotype (<3); B-F, Allotype (9). A, carapace, lateral view from the left side; B, carapace, lateral view from the left side; C, LV, internal view; D, Mxl palp and three endites; E, Tl; F, rake-like organ; G, left prehensile palp; H, right prehensile palp. Scales=0.1mm. A NEW OSTRACOD GENUS FROM QUEENSLAND 307 SYSTEMATICS Family CANDONIDAE Kaufmann, 1900 Subfamily CANDONINAE Kaufmann, 1900 Tribe CANDONOPSINI Karanovic, in press b Pioneercandonopsis gen. nov. TYPE SPECIES. Pioneercandonopsis hancodd sp. nov. ETYMOLOGY. Prefix Pioneer (after the Pioneer Valley, Queensland where the species was collected) and the genus name Candonopsis. Gender feminine. DIAGNOSIS. Carapace subreniform, surface smooth (Fig. 5A,B). Calcified inner lamella narrow posteriorly, wider anteriorly. Muscle scar imprints typical for Candoninae, although poorly visible. A1 4-segmented. Segments En2, En3, En4 and En5 fused. Small suture and seta En3a between En3 and En4. A2 in females with 4-segmentcd endopod, while 5-segmented in males. Male sexual bristles (t2 and t3). Claws G 1 and G3 male A2 reduced into setae; zl and z2 transformed into claws, former one considerably shorter; z3 seta-like (Fig. 5C). Md with 6 rays in vibratory plate. Md-palp 4-segmentcd; second segment with 1 seta externally, 3+2 setae in bunch internally; penultimate segment with 3 setae extero-distally. Terminal segment of Md-palp with broadly fused central claw (Fig. 5D); L:W of same segment equals 1.1:1. Mxl palp with rectangular terminal segment, and with 6 appendages. T1 with symmetrical palps, subterminal sclerified structures thin, only 1 well developed, another 1 very small and hardly visible (Fig. 2D). T2 5-segmented, basal segment without any seta. T3 5-segmented; basal segment with all setae (dp, dl and d2); setae Te and Tg present, while seta Tf missing; terminal segment with 2 long (Th2 and Th3 ) and 1 short seta (Th 1 ). Fu without posterior seta, anterior setae present (Fig. 5F), both claws present and long. Hemipenis with all lobes developed. Lobe V with 2 extensions: 1 dorsal and 1 distal; dorsal much smaller than distal. Lobe fc h' small and rounded; lobe *b’ with 1 ventral, finger like part. Part ‘g’ very poorly sclerified. Zenker’s organ with 7 whorls of spines. Genital field rounded, without extensions. Pioneercandonopsis hancocki gen. et sp. nov. (Figs 1-5) ETYMOLOGY. For Peter Hancock, Department of Natural Resources and Mines who collected the material. MATERIAL. Holotype 6 (dissected on slide, QM W27255), allotype 9 (dissected on slide, QM W27256); paratype 6 (dissected on slide, WAM C33460); paratype 6 and 2 paratype $ (all on SEM stub, WAM C33461); 1 paratype <3, 3 paratype 9 and 15 juveniles (nauplius larvae) (all in a test tube in 70% ethanol, QM W27257); 1 paratype <3, 2 paratype 9 and 15 juveniles (nauplius larvae) (all in a test tube in 70% ethanol, WAM C33462). TYPE LOCALITY. Australia, Queensland, Pioneer Valley, Bore No. 125004B, 148°35’24”E; 21°08’39"S; 26 June 2003; collector P. Hancock. DESCRIPTION. Holotype (RV (1); 3, total number of segments on Al: seven (0), six (1), four (2); 4, En2a seta: present (0), absent (1); 5, male sexual bristles: present (0), absent (1); 6, zl seta in males: long (0), short ( 1 ); 7, G2 claw in females: shorter than G 1 (0), as long as G1 (1); 8, number of setae on second segment of Md-palp: two(0), one( 1); 9, number of strong claws on terminal segment of Md-palp: two (0), one (1); 10, type of fusion of terminal claw on Md-palp: narrow (0), broad (1); 11, L:W ratio of terminal segment of Md-palp: less than 3:1 (0), more than 3:1 (1); 12, prehensile palps: asymmetrical (0), symmetrical (1); 13, subterminal sclerified structures on prehensile palps: both well developed (0), only one well 310 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 3. Pioneercandonopsis hancocki gen. et sp. nov. A, Holotype (cJ); B, C, Allotype (9). A, Fu; B, Fu and genital lobe; C, Md. Scale=0.1mm. A NEW OSTRACOD GENUS FROM QUEENSLAND 311 FIG 4. Pioneercandonopsis hancocki gen. et sp. nov. A-D, Holotype (d); E-F, Allotype (9). A, hemipenis; B, Zenker’s organ; C, T2; D, Furcal attachment; E, T3; F, Furcal attachment. Scales=0.1mm. 312 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Scanning Electron Microphotographs. Pioneercandonopsis hancocki gen. et sp. nov. A, B, E, Paratype ( 9 , 0.68mm, WAMC33461); C, D, F, Paratype ( 6 , 0.72mm, WAMC33461 ). A, carapace surface, frontal part; B, seta on the carapace surface; C, detail of A2; D, detail of terminal segment of Md palp; E, rake-like organ; F, detail of Fu. Scales: A=1 1.2p,m; B=666.6nm; C=4.9p.m; D=3.2p.m; E=3.69p.m; F=5.6|im. developed ( 1 ); 14, basal seta on T2: present (0), absent ( 1 ); 15, seta on second segment T2: present (0), absent (1); 16, Td2 seta: present (0), absent (1); 17, Te seta: present (0), absent ( 1 ); 1 8, Tf seta: present (0), absent ( 1 ); 19, number of short setae on terminal segment T3 : two (0), one ( 1 ); 20, posterior furcal seta: present (0), absent (1); 21, posterior furcal claw: well developed (0), reduced ( 1 ); 22, spines on furcal claws: absent (0), present ( 1 ); 23, number of ‘a’ lobe extensions: one (0), two (1); 24, internal hemipenis ducts: not coiled (0), coiled (1). The heurisitc analysis of the weighed characters from the second data matrix (Table 2) A NEW OSTRACOD GENUS FROM QUEENSLAND 313 TABLE 1. Data matrix for the species of the tribe Candonopsini and the outgroup. Characters correspond to the Fig. 6. Taxon 0 12 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 Crypior ant Iona dudkhi 000 0 000 000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 C. (A.) aula 0 0 10 1 0 1 1 0 1 1 1 - - 1 0 1 1 1 1 1 1 0 - - C. (A. ) indoles 00 10 1 - - 1 0 1 1 1 - - 1 1 1 1 1 1 1 0 0 0 1 C. (A.) williami 0 0 10 1 0 1 1 0 1 1 1 0 1 1 0 1 1 1 1 1 0 0 0 1 C. (C ) africana 0 0 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C. (C.) anteroarcuata 0 0 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 0 0 1 C. (C.) boui 0 0 0 0 1 0 1 0 0 1 1 1 0 1 0 0 0 0 0 1 1 0 0 0 1 C (C.) bujukuensis 0 0 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 l 0 1 C. (C.) dani 0 10 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C. (C.) hummelincki 0 00 0 1 - - 1 0 1 1 1 - 0 0 0 0 0 1 I 0 l - C. (C.) kingsleii 0 0 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 0 0 1 C. (C.) kiniberleyi 10 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 1 1 1 0 1 0 1 C. (C.) mareza 0 0 0 0 1 0 1 00 1 1 1 0 1 0 0 0 0 0 1 1 0 0 0 1 C. (C.) murchisoni 0 10 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C.(C.) nanui 0 0 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C. (C.) navicula 0 00 0 1 - - 1 0 1 1 1 - 0 0 0 0 0 I 1 0 1 C(C.) pu leal is 000 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C (C ) scourfieldi 0 0 0 0 1 0 1 0 0 1 1 1 0 1 0 0 0 0 0 1 1 0 0 0 1 C. (C.) solilaria 0 0 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C. (C.) sutnatratui 0 0 0 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C.(C.) tenuis 000 0 1 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 1 0 1 0 1 C. (C.) thienemanni 0 0 0 0 1 0 1 1 0 1 l 1 0 1 0 0 1 0 0 1 1 1 0 0 l C(C.) trirhota 0 00 0 1 - - 1 0 1 1 1 - 0 0 0 0 0 1 1 0 0 C. (C.) urmilae 0 10 0 1 0 1 0 0 1 1 1 0 1 0 0 0 0 0 1 1 0 0 0 1 C. (C.) westaustraliensis 1000 1 0 1 1 0 1 1 1 0 l 0 0 0 0 0 1 1 1 0 0 1 Cub. rubensis 0 0 0 0 1 1 1 - 0 1 1 0 1 1 0 0 1 1 0 1 1 0 0 1 0 Cub. anisitsi 000 0 1 0 1 1 0 1 1 0 1 0 0 0 1 0 0 1 1 0 0 1 0 Cub. ( olunibtensis 0 0 0 0 1 0 1 1 0 0 0 0 1 0 0 0 I 0 1 1 1 0 0 1 0 Cub. /airland ira 0 0 0 0 1 0 1 1 0 1 1 0 1 1 0 0 1 0 0 1 1 0 0 1 0 Carib. trapezoidea 0 10 1 1 0 1 1 0 1 1 0 1 1 0 0 0 0 0 1 1 1 0 l 0 Carib. aurirularia 0 10 1 1 0 1 1 0 1 1 0 1 1 0 0 0 0 0 1 1 I 0 1 0 Carib. ansa 0 10 1 1 0 1 1 0 1 1 0 1 1 0 0 0 0 0 1 1 1 0 1 0 Pioneer hanrorki 0 0 0 2 1 0 1 1 1 1 1 0 1 1 1 1 0 0 1 1 1 0 0 1 1 revealed 8849 equally parsimonious trees with L=21, Ci=80 and Ri=90. The strict consensus of all trees had the same length, Ci and Ri values. On the other hand, the Rachet Island Hopper analysis found only one most parsimonious tree, presented in the Fig. 7. This tree also had L=21, Ci=80 and Ri=90, and it is very similar to the strict consensus of 8849 trees except that it better resolves the relationships in the Candonopsis clades. Characters and their coding corresponding to the Table 2 matrix are as follows: 0, Posterior furcal seta: present (0), absent (1); 1, number of strong claws on terminal segment of Md-palp: two (0), one (1); 2, type of fusion of strong claws on tcmiinal segment of Md-palp: narrow (0), wide (1); 3, zl seta in males: long (0), short ( 1 ); 4, number of short setae on terminal segment T3: two (0), one ( 1 ); 5, number of ‘a’ lobe extensions: one (0), two (1); 6, number of segments on T3 : five (0), four (1); 7, subterminal sclerified structures on prehensile palps: both well developed (0), only one well developed ( 1 ); 8, L : W ratio of terminal segment of Md-palp: less than 3:1 (0), more than 3:1 (1); 9, dorsal view: LV=RV (0), LV>RV (1); 10, total number of segments on A 1 : seven (0) , six (1), four (2); 1 1, number of setae on second segment of Md-palp: two (0), one (1); 12, lateral view: symmetrical (0), asymmetrical (1); 13, spines on furcal claws: absent (0), present (1) ; 14, En2a seta: present (0), absent (1); 15, G2 claw in females: shorter than G1 (0), as long as G1 (1); 16, basal seta on T2: present (0), absent (1); 17, seta on second endopodal segment ofT2: present (0), absent (1); 18, Td2 seta: present (0), absent ( 1 ); 1 9, T f seta: present (0), absent (1); 20, Te seta: present (0), absent (1); 21, Ta’ seta: present (0), absent (1); 22, number of setae on A2 penultimate segment externally: two (0), one (1); 23, prehensile palps: asymmetrical (0), symmetrical (1); 24, internal hemipenis ducts: not coiled (0), coiled (1); 25, posterior furcal claw: well developed (0), reduced (1); 26, male sexual bristles: present (0), absent ( 1 ); 27, surface: smooth (0), ornamented ( 1 ); 28, carapace shape: subtriangular (0), subreniform (1), subtrapezoidal (2); 29, posterior margin: narrower than anterior (0), wider or equal to anterior ( 1 ); 30, CB2 seta: present (0), absent (1); 31, En2d seta: present (0), absent (1); 32, En3b seta: present (0), absent (1); 33, En4b seta: present (0), absent. DISCUSSION The Candonopsini has at least 2 evolutionary branches: 1 with Candonopsis Vavra, 1891, and the other with Caribecandona, Cubacandona and Pioneercandonopsis . The first branch, presented as a separate clade in all cladograms 314 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 2. Data matrix for the species of the tribe Candonopsini and the outgroup. Characters correspond to the Fig. 7. Taxon 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Cr. dudichi 0000000000 0 0 0 0 0 0 C. ( A .) aula 1 1 1 100- 1 1 0 0 0 0 1 1 C. (A.) indoles 1 1 • 1 - 0 - 1 1 0 0 0 0 1 1 C. (A.) willuuni 1 1 1 10 0 1 1 1 0 0 0 0 1 1 C (C.)africana 1 1 - 10 0 1 1 0 0 - 0 1 - - C. (C.) anleroart . 1 1 1 10 0 1 1 0 0 0 0 0 1 1 C (C) boui > 1 1 10 0 1 1 0 0 0 0 0 1 0 C. (C.) bujukuensis 1 1 1 100 1 1 0 0 - 0 1 - 1 C. (C.)dani 1 1 I 10 0 1 1 0 0 0 1 1 1 1 C. (C.) hummelinrki 1 1 - 1 - 0 - 1 0 0 0 0 1 1 1 C. (C.) kingsleii 1 1 1 10 0 1 1 0 0 0 0 0 1 1 C. (C.) kimberleyi 1 1 1 10 0 1 1 0 0 0 0 1 1 I C. (C.) rnare'XJ 1 1 1 10 0 1 1 0 0 0 0 0 1 0 C. (C.) murchisoni 1 1 1 10 0 1 1 0 0 0 1 1 1 1 C. (C.) narna 1 1 1 10 0 1 1 0 0 0 0 1 - I C. (C.) navicula 1 1 - 1 - 0 - 1 0 0 0 0 1 1 1 C (C) pit teal is 1 1 - 10 0 1 1 0 0 0 1 - 1 C. (C.) scourfieldi 1 I 1 10 0 1 1 0 0 0 0 0 1 0 C. (C.) solitoria 1 1 - 10 0 1 1 0 0 0 1 - - C.(C.) sumatrana 1 1 - 10 0 1 1 0 0 0 0 1 1 1 C. (C.) tenuis 1 1 1 10 0 1 1 0 0 0 0 1 1 1 C. (C.) thienetnanni 1 1 1 10 0 1 1 0 0 0 0 0 - 1 C. (C.) trichota 1 1 - 1 - 0 - 1 0 0 0 0 0 1 C. (C.) unnilae 1 1 1 10 0 1 1 0 0 0 1 0 1 0 C. (C.) westaustral 1 1 1 10 0 1 1 0 0 0 0 0 1 1 Cub. cubensis 1 1 1 110 1 0 0 0 0 0 0 1 - Cub. anisitsi 1 i - 1 10000 0 0 0 - - Cub. columbiensis 1 0 0 1 111000 0 0 0 0 - 1 Cub.falclandica 1 1 - 1 10 10 0 0 0 0 - 1 Car. trapezoidea 1 1 1 110 1 0 0 1 0 1 0 1 1 Car. auricularia 1 1 1 110 1 0 0 1 0 1 0 1 1 Car. ansa 1 1 1 1 10 10 0 1 0 1 0 1 1 Pioneer, hancocki 1 1 1 1 10 10 0 2 1 0 0 1 1 (Figs 6, 7), is defined with a long terminal segment on the Md-palp, a synapomorphy of all 24 Recent species of Candonopsis. In addition, all species are characterised with 2 sym- plesiomorphies: the asymmetrical prehensile palps, and only 1 extension of the lobe V on the hemipenis. However, compared with other Candoninae the palps and the hemipenis in Candonopsis are very characteristic and have little in common with other Candoninae. Within Candonopsis several distinct clades occur on all resulting cladograms. The largest is defined by one synapomorphic character (presence of a spine on the furcal claws), and it includes almost 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 1 1 - 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 - 1 0 - 0 1 1 1 1 1 1 1 0 1 1 1 1 1 0 1 0 0 0 1 1 0 1 1 1 . 0 0 0 0 - . 0 1 0 0 0 1 1 - - - * 0 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 - 1 0 1 0 0 0 1 1 0 1 1 1 - 0 0 0 0 - 0 0 1 0 0 0 1 1 - - - - 0 0 0 0 0 0 1 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 - - 0 - 0 1 1 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 1 0 0 0 0 0 0 0 I 0 1 1 0 1 0 0 1 2 0 0 1 1 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 - 0 0 1 0 0 0 1 1 - - • - 0 0 0 0 0 0 0 - - 0 - 0 1 1 0 0 0 0 - 0 0 0 0 - 1 0 1 0 0 0 1 1 * - - * 0 0 0 0 0 0 1 0 1 0 0 0 1 1 0 1 0 0 - 0 0 0 0 - - 0 1 0 0 0 1 1 - - - - 0 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 - 0 1 0 0 - - 0 1 0 0 0 0 0 - - - - - 0 0 0 0 - - - - 0 0 0 0 - - - - 0 0 0 0 0 - 1 0 1 0 0 0 1 1 - 1 1 0 0 0 0 0 0 1 0 0 1 1 0 1 1 1 0 0 1 1 0 0 1 0 1 - 1 1 0 0 1 0 2 1 0 1 1 0 - 0 0 0 - 1 0 0 0 2 1 * - - 0 1 0 - 1 1 0 0 0 0 1 1 - • - 0 0 0 - 1 0 0 0 0 1 1 0 0 0 0 0 0 1 1 0 1 0 0 2 0 0 1 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 1 0 0 2 0 0 1 0 0 1 1 1 t 0 1 1 1 1 0 0 0 1 1 1 1 1 1 all African, the only Central American and almost all Australian species of Candonopsis s. str. This correlates with Klie's (1932) zoogeographical groupings of the species based on the appearance of the furcal claws, in which almost all species from the tropics and subtropics have a spine at least on one of the furcal claws, while European species have both claws spineless. The exceptions are 1 species from Lake Tanganyika (G anteroarcuata Rome, 1962) and 1 from the Murchison Region (C. westaustraliensis Karanovic & Marmonier, 2002). The Australian species has, in addition, a reduced posterior furcal claw. Abcandonopsis A NEW OSTRACOD GENUS FROM QUEENSLAND 315 Karanovic, in press b is a clade defined by only 1 autapomorphic character - asymmetrical valves in dorsal view. All other features of this subgenus are homoplastic. For example, the 3 known species of Abccindonopsis have many setal reductions that are also recorded in Pioneer- candonopsis hancocki sp. nov., C. (C.) thienemanni Schafer, 1945 and C. (C.) kimberleyi Karanovic & Marmonier, 2002. Because all these species are subterranean inhabitants, some of the setal reductions may well be stygomorphies. A clade containing Candonopsis (C.) scourfieldi Brady, 1910; C. (C.) boui Danielopol, 1978, C. (C.) unnilae Gupta, 1988; and C. (C.) mareza Karanovic & Petkovski, 1999 is supported with the symplesiomorphic state of the character number 7, i.e. a short claw G2 on the female A2. This claw is long in all other representatives of the tribe. The smallest clade in Candonopsis includes Australian C. (C.) murchisoni Karanovic & Marmonier, 2002 and C. (C.) dani Karanovic & Marmonier, 2002. Although, the connecting character (asymmetry of the valves in the lateral view) is homoplastic in the tribe, and it is found in all Caribecandona species, and in C. (C.) unnilae Gupta, 1988. the two Australian species are indeed more closely related to each other than either is to any other representative of Candonopsis (Karanovic & Marmonier, 2002). The second branch in Candonopsini is supported with 2 synapomorphic characters: 2 extensions of the lobe ‘a’ on the hemipenis, and almost symmetrical prehensile palps. Also, all species have a small terminal segment of the Md-palp. Pioneercandonopsis is a clade defined by 2 autapomorphies: 4-segmented A1 and only 1 seta externally on the second segment of the Md-palp. The reduction of A1 segments more closely relates Pioneercandonopsis to Caribe- candona than to Cubacandona. However, in the former genus, A 1 is 6-segmented, the posterior furcal claw is reduced and the valves are asymmetrical in the lateral view. The new genus and two genera from South and Central America occur as a monophyletic clade in Fig. 6, but in Fig. 7, the clade does not include Cubacandona cubensis (Mehes, 1913) or C. anisitsi (Daday, 1905). According to Mehes’ (191 3) drawings, C. cubensis has 2 strong claws on the terminal segment of the Md-palp, which gives it an isolated position in the tribe. Cubacandona anisitsi stands apart because it has both subterminal sclerified structures well developed (Daday, 105). Cubacandona columbiensis and C. anisitsi , together with about a dozen other species of the Candonopsini, are insufficiently described. Because of this, many characters are unknown (and coded *-’) in the second data matrix (Table 2) and in the cladistic analysis treated as ambiguous. Many ambiguous characters and a great number of homoplasies in the tribe Candonopsini produced confusing cladograms. Therefore, characters from the second data matrix (Table 2) were weighted in accordance to their taxonomic importance. The resulting cladogram (Fig. 7) is very similar to the one obtained after the analysis of the first data set (Fig. 6). The difference is that some newly introduced characters (chaetotaxy of the Al, A2 and Tl) better resolve phylogenetical relationships in the Candonopsis clades. This suggests that a better understanding of the morphology of the insufficiently described taxa would certainly contribute to the cladogram resolution. However, both trees correlate with the present systematics of the tribe Candonopsini with the exception of the genus Cubacandona , which seems to be a polyphyletic genus. ACKNOWLEDGEMENTS The paper was written during tenure of an Australian Biological Resources Study grant to work on the revision of the Candoninae. LITERATURE CITED BALTANAS, A., NAMITKO, T. & DANIELOPOL, D.L. 2000. Biogeography and disparity within the genus Ciyptocandona (Crustacea, Ostracoda). Vie et Milieu 50: 297-310. BROODBAKKFR, N.W. 1983. The subfamily Candoninae (Crustacea, Ostracod) in the West Indies. Bijdragen tot de Dierkunde 53: 287-326. BROODBAKKFR, N.W. & DANIELOPOL, D.L. 1982. The chaetotaxy of Cypridacea (Crustacea, Ostracoda) limbs: proposal for a descriptive model. Bijdragen tot de Dierkunde 52: 103-120. DADAY, J. 1905. Untcrsuchungen iiber die Siisswasser-Mikrofauna Paraguays. Zoologica 44: 1-374. 1910. Die Siisswasser-Mikrofauna Deutsch-Ost- Afrikas. Zoologica 59: 1-314. 1913. Cladoceren und Ostracoden aus Sud- und Sudwcstafrika. Denkschriften der Medizinisch- Nnaturwissenschaftlichen Gescllschaft zu Jena 17: 92-102. DANIELOPOL, D.L. 1969. Rccherches sur la morphologie de Porgane copulateur male chez quelques ostracodes du genre Candona Baird (fam. Cyprididae Baird). Pp. 136-153. In Neale, 316 MEMOIRS OF THE QUEENSLAND MUSEUM 3 8 14 13 18 24 4 6 7 9 10 13 19 20 1111111 l 21 0 - 0 - 1 1 9 IO 13 18 O O O 1 Crypt ocandona dudichi Pioneer . hancocki Carib. trapezoidea Carib. aiiricularia Carib. ansa Cub. falclandica Cub. cubensis Cub. an i sit si Cub. col umb i ens i s C. (C.) anteroarcuata C. (C.) ki ngslei i C (C.) trichota C. (C.) boui C. (C.) mareza C. (C.) scourfieldi C. (C.) u rrn i l a e C. (C.) westaustraliensis C. (C.) thieneinanni C. (A.) aula C. (A.) indoles C. (A.) w ill i ami C. (C.) africana C. (C.) bujukuensis C. (C.) hummel incki C. (C.) kimberleyi C. (C.) nama C. (C.) navi cu la C (C.) pu teal is C. (C.) sol i ta ri a C. (C.) sumatrana C. (C.) tenuis C. (C.) dani ■ C. (C.) m u rch is on i FIG. 6. Cladogram of the tribe Candonopsini based on 24 characters. Number above the hash-marks correspond to the character, numbers below are character states. Black dots are apomorpies, white dots are homoplasies. 0 18 1 1 A NEW OSTRACOD GENUS FROM QUEENSLAND 317 O 3 4 5 I *2-1 21129 Cr. dud i chi Cub . columbiensis Cub. anisitsi Cub. falclandica I ft 20 26 28 32 — Cub. cubensis I I I 2 I 10 II 16 17 IS 1921 24 3032 33 Pioneer. hancocki — 0-00-0- 112 0 28 29 Car. trapezoidea Car. auricularia Car. ansa C. (C.) anteroarcuata C. (C.) trichota 0 0 -ooo— C. (C.) wes tan st ra liens i s i i i C. (C.) kingsleii 9 16 18 192021 31 3233 I I I I C. (C.) thienemanni C. (A.) aula C. (A.) indoles C. (A.) williami ff— C. (C.) mareza C. ( C. ) scourfieldi — o — C. (C.) boui ~^>~C. (C.) urmilae C. (C.) africana 1921 27 28 2931 32 C. (C. ) kimberleyi 22 ° — C. ( C. ) nama C. ( C. ) bujukuensis 22 31 C. (C.) hummel incki C. (C.) navicula C. (C.) putealis C. (C.) sol it aria C. (C.) s uniat rana C. (C.) tenuis C. (C.) dani C. (C.) murchisoni FIG 7. Cladogram of the tribe Candonopsini based on 33 weighted characters. Number above the hash-marks correspond to the character, numbers below' are character states. Black dots are apomorpies, white dots are homoplasies. 318 MEMOIRS OF THE QUEENSLAND MUSEUM J.W. (ed.) The taxonomy, morphology and ecology of recent Ostracoda. (Oliver & Boyd Ltd: Edinbourgh). 1978. Ober Herkunft und Morphologie der Siisswasser-hypogaischen Candoninae (Crustacea, Ostracoda). Sitzungsberichten der Osterreichischen Akademie dcr Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Abteilung I 187: 1-162. 1 980. Deux especes hypogees du genre Candonopsis (Ostracoda, Candoninae) du Sud de la France et de Cuba. Vie et Milieu 30: 315-323. DE DECKKER, P. 1 976. Trigonocypris a new ostracod genus from Queensland. Australian Journal of Zoology 24: 145-157. 1978. Comparative morphology and review of Mytilocyprinid Ostracods (Family Cypridae). Australian Journal of Zoology, Suppl. Series 58: 1-61. 1981. Taxonomy and ecological notes of some ostracods from Australian inland waters. Transactions of the Royal Society of South Australia 105: 91-138. 1982. On Bcnnelongia tunta De Deckker sp. nov. Stereo- Atlas of Ostracod Shells 9: 117-124. 1983. Terrestrial ostracods in Australia. Pp. 87-100. In Lowry, J.K. (ed.) Paper from the conference of the biology and evolution of Crustacea. Australian Museum Memoir 1 80. DE DECKKER, P. & McKENZIE, K.G. 1981. Bcnnelongia. a new Cypridid ostracod genus from Australasia. Transactions of the Royal Society of South Australia 105: 53-58. GOLOBOFF, P. 1999. NONA (NO NAME) version 2. (Published by the author: Tucuman, Argentina). GUPTA, L.P. 1 988. Candonopsis urmilae a new species of subterranean Crustacea (Ostracoda: Candonidae) from India. Records of the Zoological Survey of India 85: 419-427. HARDING J.P. 1962. Mungava munda and four other new species of Ostracod Crustaceans from fish stomachs. The Natural History of Rcnnell Island, British Solomon Islands 4: 51-62. KARANOVIC, I. 2003a. Towards a revision of Candoninae (Crustacea: Ostracoda): Description of two new genera from Australian groundwaters. Species Diversity 8: 352-383. 2003b. A new genus of Candoninae (Crustacea, Ostracoda, Candonidae) from the subterranean waters of southwestern Western Australia. Records of the Western Australian Museum 2 1 : 315-332. In press a. Towards a revision of Candoninae (Crustace, Ostracoda): Australian representatives of the subfamily, with descriptions of three new genera and seven new species. New Zealand Journal of Marine and Freshwater Research. In press b. Towards a revision of Candoninae (Crustacea, Ostracoda): On the genus Candonopsis Vavra, with descriptions of new taxa. Subterranean Biology 2. In press c. Comparative morphology of the Candoninae antennula, with remarks on the ancestral state in ostracods (Crustacea, Ostracoda) and proposed new terminology. Crustaceana. KARANOVIC. I. & MARMONIER, P. 2002. On the genus Candanopis (Crustacea: Ostracoda: Candoninae) in Australia, with a key to the world recent representatives. Annales de Limnologie 38: 199-240. 2003. Three new genera and nine new species of the subfamily Candoninae (Crustacea, Ostracoda, Podocopida) from the Pilbara Region (Western Australia). Beaufortia 53: 1-51. KARANOVIC, I. & PETKOVSKI, T. 1999. Two new species of the subfamily Candoninae (Ostracoda) from Montenegro (SE Europe). Crustaceana 72: 603-616. KITCHING I. J., FOREY, P. L., HUMPHRIES, C. J. & WILLIAMS, D. M. 1998. Cladistics, the theory and practice of parsimony analysis. Second edition. (Oxford University Press: Oxford). KLIE, W. 1930. Ostracoden aus dem paraguay ischen Teile des Gran-Chaco. Archiv fur Hydrobiologie 22: 221-258. 1932. Die Ostracoden der Deutschen Limnologischen Sunda-Expedition. Archiv fur Hydrobiologie, Suppl. 9: Tropische Binnengewasser 3: 447-502. 1935. Ostracoda aus dem tropischen Westafrika. Archiv fur Hydrobiologie 27: 35-68. 1 936. Ostracoden aus Kamerun. Revue de Zoologie et de Botanique Africaines 27: 287-309. 1944. Ostracoda. Exploration du Parc National Albert, Mission H. Damas. Institut des Parcs Nationaux du Congo Beige 12: 1-62. LOFFLER, H. 1968. Die Crustaceenfauna der Binnengewasser Ostafrikanischer Hochbcrge. Hochgebirgsforschung 1: 107-170. MARTENS, K. 1984. On the freshwater ostracods (Crustacea, Ostracoda) of the Sudan, with special reference to the Red Sea Hills, including a description of a new species. Hydrobiologia 110: 137-161. 1987. Homology and functional morphology of the sexual dimorphism in the antenna of Sclerocypris Sars, 1924 (Crustacea, Ostracoda, Megalocypridinae). Bijdragen tot de Dierkunde 57: 183-190. 1992. On Namibcypris costata n. gen., n. sp. (Crustacea. Ostracoda, Candoninae) from a spring in Northern Namibia, with the description of a new tribe and a discussion on the classification of the Podocopina. Stygologia 7: 27-42. A NEW OSTRACOD GENUS FROM QUEENSLAND 319 1998. General morphology of non-marine Ostracods. Pp. 57-75. In Martens, K. (cd.) Sex and Partehnogenesis: evolutionary ecology of reproductive modes in non-marine ostracods. ( Backhuys Publishers: Leiden, The Netherlands). MEHES, G 1 9 1 3. Susswasser-Ostracoden aus Columbien und Argentinien. Memoires de la Societc neuchateloise des Sciences naturelles 5: 639-663. MEISCH, C. 1996. Contirbution to the taxonomy of Pseudocandona and four related genera, with the description of Schellencandona nov. gen., a list of the Candoninae genera, and the key to the European genera of the subfamily (Crustacea, Ostracoda). Bulletin dc la Societe des Natural istes luxembourgeois 97: 211-237. NAMIOTKO, T., MEISCH. C., GIDO, Z. & DANIELOPOL, D. L. 2001. Redescription, taxonomy and ecology of Cryptocandona dudichi (Klie, 1930) (Crustacea, Ostracoda). Bulletin de la Societe des Naturalistes luxembourgeois 102: 109-130. NIXON, K.C. 2002. WinClada version 1.00.08. (Published by the author: Ithaca, New York). PETKOVSKI, T.K. 1977. Ostracoden fauna des Mindelsees (S. W. Deutschland). Acta Musei Macedonici Scientarum Naturalium 15: 49-94. PETKOVSKI, T.K. & MEISCH, C. 1995. Interesting freshwater Ostracoda (Crustacea) from Macedonia. Bulletin de la Societc des Naturalistes luxembourgeois 96: 167-183. ROME, R. 1962. Ostracodes. Exploration Hydro- biologique du Lac Tanganika (1946-1947). Resultats Scicntifiques 3: 1-305. SARS, GO. 1 896. On fresh-water Entomostraca from the neighbourhood of Sydney, partly raised from dried mud. Archiv for Mathematik og Naturvidenskab 18: 1-81. 1889. On some freshwater Ostracoda and Copepoda, raised from dried Australian mud. Christiania Videnskabs-Selskabs Forhandlinger 8: 1-79. SCHAFER, H.W. 1945. Grundwasser-Ostracodcn aus Griechenland. Archiv fur Hydrobiologie 40: 847-866. VA VRA, V. 1 895. Die von Dr F. Stuhlmann gesammelten. Susswasser-Ostracoden Zanzibar's. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten 12: 1-23. 1898. Susswasser-Ostracoden. Hamburger Magalhaensische Sammclreise: 1-25. VICTOR, R. & FERNANDO, C.H. 1978. Systematics and ecological notes on Ostracoda from container habitats of some South Pacific Islands. Canadian Journal of Zoology 56: 414-422. WALKER-SMITH, GK. & POORE, GC.B. 2001. A phylogeny of the Leptostraca (Crustacea) with keys to families and genera. Memoirs of Museum Victoria 58: 383-410. NEW BIOGEOGRAPHICAL INFORMATION ON QUEENSLAND CHONDRICHTHYANS P.M. KYNE, J.W. JOHNSON, A. J. COURTNEY AND M.B. BENNETT Kyne, P.M., Johnson, J.W., Courtney, A.J. & Bennett, M.B. 2005 01 10: New biogeographical information on Queensland chondrichthyans. Memoirs of the Queensland Museum 50(2): 321-327. Brisbane. ISSN 0079-8835. New information is presented on chondrichtbyan fishes. Mobula japanica (Muller & Henle, 1841) is reported for the first time from Australian waters from two specimens, one from Lake Macquarie, New South Wales and one from Fraser Island, Queensland. Squaliolus aliae Teng, 1959 and Asvmholus analis (Ogilby, 1885) are reported tor the first time from waters off Queensland, and aspects of the biology of A. analis are discussed. Significant southern range extensions are reported for Aetomylaeus nichofii (Schneider, 1801) and Dasyatis leylandi Last, 1987. Southern range extensions are also reported for Carcharhinus amboinensis (Muller & Henle, 1839) and Carcharhinus cautus (Whitley, 1945) and these species are recorded for the first time from Moreton Bay. Northern range extensions arc reported for Hypnos monopterygius (Shaw & Nodder, 1795) and Orectolobus maculatus (Bonnaterre, 1788). Comments are also made on the distribution of Dipturus australis (Macleay, 1884) and Mvliobatis hamlyni Ogilby, 1911. □ Biogeography, Chondrichthyes, Queensland waters, Elasmobranchii, Squaliolus, Orectolobus, Asymbolus, Dasyatis, Aetomylaeus, Mvliobatis. Mobula . P.M. Kyne and M.B. Bennett, School of Biomedical Sciences, Department of Anatomy and Developmental Biology, University of Queensland, St Lucia, 4072; J.W. Johnson, Queensland Museum, PO Box 3300 South Brisbane 4101 (e-mail: JeJfJ@qm.qld.gov.au); A.J. Courtney, Southern Fisheries Centre, Queensland Department of Primary Industries, PO Box 76, Deception Bay, 4508, Australia; 30 May 2004. Australian waters support a diverse chondrichthyan fauna, with at least 296 known species (Last & Stevens, 1994), while additional species are still being recorded, for example, Isurus paucus Guitart Manday, 1966 (Stevens, 1995) and Isistius plutodus Garrick & Springer, 1964 (McGrouther, 2001). Over half of these species have been recorded from Queensland waters or the Australian Exclusive Economic Zone adjacent to Queensland (Last & Stevens, 1 994). Despite the diversity of fauna in this state, there exists a paucity of data concerning the basic biology of most Queensland chondrichthyans. Limited published research is available on the life history of tropical elasmobranch species from the Gulf of Carpentaria and other northern regions (e.g. Stevens & McLoughlin. 1991 ; Salini et al„ 1 992), inshore species from the Townsville region (e.g. Simpfcndorfer, 1998), Hemiscyllium ocellatum (Bonnaterre, 1788) from the southern Great Barrier Reef (e.g. Hcupel et ah, 1999) and Apty f chotrema rostrata (Shaw & Nodder, 1794) from Moreton Bay (e.g. Kyne & Bennett, 2002). Similarly with life history data, obtaining information on a species’ geographical and bathymetric distribution is essential for understanding ecology, for management purposes, and in defining levels of interactions with fisheries. Information concerning the biogeography of many species in Queensland waters continues to expand and to be refined with the collection and identification of new' material. This paper presents information on new chondrichthyan material deposited in the Queensland Museum, including that taken by Queensland Department of Primary Industries (QDPI) trawl surveys. A number of range extensions are documented as well as a new record for Australia. MATERIALS AND METHODS A. analis , D. australis , D. leylandi and A. nichofii were captured by demersal otter trawl as part of a QDPI project on the bycatch of the Queensland East Coast Trawl Fishery. D. leylandi and A. nichofii were collected by a 13.6m commercial trawler fitted with twin 4-seam Florida Flyer nets; net body mesh size of 5.0cm; codend mesh size of 4.4cm; headrope length was 7.3m. Specimens of A. analis and D. australis were taken by a 17.1m commercial trawler fitted with three 2-seam Florida Flyer nets; net body mesh size of 5.0cm; codend mesh size of 4.4cm; headrope length was 12.8m or 322 MEMOIRS OF THE QUEENSLAND MUSEUM 21.9m (depending on nets fitted at the time). Capture details for specimens collected by other methods are given with the specimen information. Specimens were identified using Last & Stevens (1994) and Last (1999). Measurements and mass were recorded and maturity assessed (following Bass et al., 1973) in the laboratory. The following abbreviations are used for institutions: AMS, Australian Museum, Sydney: CSIRO, CSIRO Marine Laboratories, Hobart: QM, Queensland Museum, Brisbane. Methods and abbreviations for measurements of sharks follow Coinpagno (2001). Additional measure- ments for batoids are DW (disc width) and DL (disc length). Those for Mobula japanica follow Notarbartolo-Di-Sciara (1987). SPECIES RECORDS AND DISCUSSION Squaliolus aliae Teng, 1959 MATERIAL. QMI32529; <3, 134mm TL (Fig. 1A) was captured by trawl 30-40 miles N of Cape Moreton. Qld. 26°2 1 -3 1 'S, 1 53°28'E, at 1 83-20 1 m depth on 23/04/2000. AMS 143064-001: <3, 130mm TL was captured by plankton tow in the Coral Sea. 1 2°44-50'S, 1 53°52-55'E, at 120-1 50m depth on 16-17/05/1995. REMARKS. Squaliolus aliae is probably the smallest known living shark species (Last & Stevens, 1994). It was known on the Australian east coast by AMS 1205 1 5-00 1,9, 98mm TL, E of Sydney, NSW. 33°44'S, 151°52'E. demersal trawl, 457m, 24/1 1/1977; and AMSI3041 1-001, 9, 213mm TL, E of Kiama, NSW, 34°42'S, 151°16'E, demersal trawl, 474m, 21/11/1978. This species is also reported from NW WA (Last & Stevens, 1994). Regionally, it is reported from Japan, Taiwan and the Philippines (Last & Stevens, 1994). A close relative, Squaliolus laticaudus Smith & Radcliffe, 1912, has a nearly circumtropical distribution, but has not yet been confirmed from Australian waters. S. laticaudus has a larger eye (eye length, EYL, 73.4-85.6% of interorbital space, INO, 61.0%-81.7% of preorbital length, POB) than S. aliae (EYL 46.3-69.9% INO, 42.6%-66.4% POB) (Sasaki & Uyeno, 1987). QMI32529 has an EYL 58.4% INO and 51.0% POB and AMSI43064-001 an EYL 48.6% and 52.3% POB). Sasaki & Uyeno ( 1 987) provided 2 further characters to separate these species. Firstly, in S. aliae the upper margin of the orbit is chevron-shaped, while in S. laticaudus it is nearly straight. The chevron-shaped upper margin is clear in QMI32529 and AMS 143064-001. Secondly, a pair of papillae is present on the lip of the upper jaw in S. aliae and absent in 5. laticaudus . Due to damage to the mouth of QMI32529 and the effects of fixation this feature was not useful in identifying these specimens. However, given the size and shape of the upper margin of the eye, both specimens are S. aliae , as arc the 2 NSW specimens (AMS 120515-001, EYL 63.7% INO: AMS 13041 1-001, EYL 56.8% INO). Orectolobus niaculatus (Bonnaterre, 1788) MATERIAL. Collected by demersal trawl by P. Duncan off north Qld: QMI31417, immature <3, 845mm TL, off Hardline Reefs, 20°46'S, 151°20'E, 218m, 09/08/1999; QMI31438, immature <3, 445mm TL, off Caldcr Island 20°46'S, 151°28'E, 215m, 10/08/1999. REMARKS. Orectolobus niaculatus was known from the S coast of Australia between SE Qld and SW WA, excluding Tasmania. Further records from Japan and the South China Sea require confirmation (Last & Stevens, 1994). The records presented here significantly increase the N distribution , recording it for the first time from the Great Barrier Reef region. Furthermore, the known bathymetric range is increased from a depth of 1 10m to 218m. Asymbolus analis (Ogilby, 1885) MATERIAL. Captured by demersal otter trawl from S Qld waters in depths of 85- 159m. QMI32876, mature <3, 458mm TL, 27°02'S, 153°37'E, 137m, 1 3/03/200 1 ; QMI33353, immature 9, 329mm TL, mass llOg, 27°10'S, 153°38'E, 135m, 29/07/2002; and QMI33354 (Fig. IB), immature c 3, 300mm TL, mass 81g, 27°47'S, 153°49'E, 86m, 24/07/2002. 10 specimens were dissected to obtain life history data (see details below). OTHER MATERIAL. AMS IB8255, 126023-001, 126023-001 134747-001, 135418-001. 135423-001, 135423-002, 140476-001. REMARKS. These specimens are consistent with many of the diagnostic features outlined in Last & Stevens (1994) and Last (1999). They have a greyish background colouration, overlain FIG. 1 . A, Squaliolus aliae , QM 132529, 1 34mm TL, lateral view. B, Asymbolus analis , QM 133354, 300mm TL, lateral view. C, Dasyatis leylandi , QM 133352, 1 30mm DW. dorsal view. D, Aetomylaeus nichofii , QM 133854, 343mm DW, dorsal view. E, Mobula japanica , QM 133855, 1088mm DW, dorsal view. F, Hypnos monopteiygius , photographed on the south side of Heron Island Reef, Great Barrier Reef, November 1995 (Photo: M. Prekker). QUEENSLAND CHONDRICHTHYAN BIOGEOGRAPHY 323 324 MEMOIRS OF THE QUEENSLAND MUSEUM with saddle-like blotches, dark brownish spots and whitish flecks, and densely packed denticles, giving the skin a rough appearance. These specimens may be distinguished from the sympatric Asymbolus rubiginosus Last, Gomon & Gledhill, 1999, another dark-spotted Asymbolus species, by a shorter caudal fin (range of CDM, the dorsal margin of the caudal fin, of the three specimens 19.0-20.6% TL vs 23.7- 26.6% TL in A. rubiginosus ), and a larger, broadly semicircular, posterior nasal flap. The Queensland specimens, however, differ from some characters outlined in Last (1999). The range of the head length (HDL) of the 3 QM specimens (17.3-18.0% TL) falls within the range of HDL given for A. rubiginosus (16.7-18.0% TL) and outside that for A. analis (18.1-19.0% TL). Furthermore, the interdorsal space (IDS) (12.1-13.1% TL) falls within the ranges given for both A. rubiginosus (IDS 12.9-15.1% TL) and A. analis (IDS 10.7-12.5% TL). Interdorsal space for A. analis in Last (1999) is given as ‘much less than 1 .5 times total length of first dorsal fin’. Relative proportions of this feature are used to separate A. analis from Asymbolus pallidus Last, Gomon & Gledhill, 1999 and Asymbolus occiduus Last, Gomon & Gledhill, 1999, 2 other dark-spotted Asymbolus species. However, the IDS of the QM specimens range from 1.46-1.71 times D1L. It appears that interdorsal space and head length proportions are more variable than previously thought, and may not be useful for Asymbolus species ident- ification. Additional AMS material examined (n=8) supported this suggestion, with the range of IDS 1.36-1.66 D1L and the range of HDL 16.7- 19.6% TL for specimens from NSW. Asymbolus analis is endemic to the east coast of Australia, and was previously thought to be confined to NSW and Vic. waters from Port Macquarie S to Lakes Entrance. These records provide the first account of A. analis from Queensland waters, and examination of additional NSW material extends its bathymetric distribution from 79m reported in Last & Stevens (1994) to 199m. Asymbolus includes eight species restricted to Australian waters (Last, 1999) and an undescribed species from New Caledonia (Seret, 1994). Last Sl Stevens (1994) reported that 8 A. analis mature at 520mm TL, but provide no information on 9 size at maturity. Of the 3 8 QM specimens , one was immature at 300mm TL and the others were mature at 458 and 460mm TL, lowering the previous maturity estimate by at TABLE 1. Maximum ovum diameter (MOD) and number of yellow-yolked ova > 5mm diameter for five mature 9 Asymbolus analis. Capture date TL (mm) MOD (mm) Ova number 15/03/2001 455 19 12 1 4/09/200 1 460 17 8 16/09/2001 470 24 7 13/10/2001 460 20 7 14/10/2001 470 23 15 least 60mm. Five $ of 278-329mm TL were all immature; while 5 9 of455-470mm TL were all mature. Mature females had a single functional ovary. None of the 5 mature females were carrying eggeases in their oviducts, but all had a number of ripe ovarian ova with maximum ovum diameters ranging from 17-24mm (Table 1). Oviducal glands were well-developed and expanded in all mature females with an average diameter of 27.3mm (range 25-29mm). The large ripe ova during the 3 months in which mature female A. analis were captured (March, September and October) suggests that the species may not have a well-defined reproductive season, similar to the situation reported for some other scyliorhinid sharks (e.g. Horie & Tanaka, 2000). Carcharhinus amboinensis (Muller & Henle, 1839) MATERIAL . QM 133857; 9 , 1 195mm TL, 940mm FL, 840mm PCL, mass 1 3.3kg was captured by handline by R. Pillans 800m ofFShomcliffe Jetty, Moreton Bay, 27°19'S, 1 53 °05'E, at 4m depth cm 2 7/ 12/2002. It is most likely to be immature, as females are known to mature at about 2 1 5cm TL (Last & Stevens, 1994). REMARKS. Carcharhinus amboinensis has been reported from the E North Atlantic and the Indo-West Pacific and locally in tropical and subtropical waters from Carnarvon, WA to Bundaberg, Qld (24°45'S, 152°24'E) (Last & Stevens, 1994). This provides the first record of C. amboinensis S of Bundaberg and the first record from Moreton Bay, not having been previously reported by Johnson (1999). This species is reasonably common within Moreton Bay (R. Pillans, pers. comm.), although not well represented in the QM collection. It is similar to the bull shark C. leucas (Valenciennes, in Muller & Henle, 1 839), possibly leading to confusion in the field with this more abundant species. QUEENSLAND CHONDRICHTHYAN BIOGEOGRAPHY 325 Carcharhinus cautus (Whitley, 1945) MATERIAL. QMI32158; 9, 670mm TL, 535mm FL, 490mm PCL) was collected from the shoreline adjacent to Toondah Harbour, Cleveland, Moreton Bay, 27°32'S, 1 53° 1 7'E, on 24/05/2000. The shark was found dead, but showed no signs of hook or net capture; QMI33856; 9 , 1260mm TL, 1060mm FL, 950mm PCL, mass 12.5kg) was captured by handline by R. Pillans off Green Island, Moreton Bay, 27°26'S, 153°14'E, at 5m depth on 06/03/2002. Given its size, and the fresh mating scars, this specimen was most likely mature, as both sexes are reported to mature at 800-850mm TL (Last & Stevens, 1994). REMARKS. Known only from S New Guinea, the Solomon Islands and tropical Australia between Carnarvon, WA and Bundaberg, Qld (24°45’S) (Last & Stevens, 1994). These are the first records of C. cautus south of Bundaberg and the first records from Moreton Bay, not having been previously reported by Johnson (1999). Dipturus australis (Macleay, 1884) MATERIAL. QMI33853; 9, 358mm TL, 258mm DW, 212mm DL, mass 320g) was captured by demersal otter trawl off Moreton Island, 27°13'S, 153°39'E at 135m depth on 28/07/2002. Other specimens are QMI 1 1 903, from 26°40'S, 153°15'E and II 1904 from 27°44'S, 153°27'E. REMARKS. This species is reported to be ‘the most common skate on the continental shelf of central eastern Australia’ (Last & Stevens, 1 994), being recorded from off Moreton Bay S to Jervis Bay, NSW. Last & Stevens (1994: 347) stated that ‘records of this species from prawn trawl catches from southern Queensland require validation.' This specimen, taken by a commercial prawn trawler, confirms that the Queensland trawl fishery interacts with this species. Despite its apparent common occurrence off NS W, this species appears to be uncommon in Queensland waters. Hypnos monopterygius (Shaw & Nodder, 1795) MATERIAL. An individual (Fig. IF) was observed by MBB otfthe reef edge on the S side of Heron Island, Great Barrier Reef (23°27'S, 15I°55'E) in November 1995. It was initially buried in coral/sand substrate at 4m depth but was disturbed by a boat anchor. It proceeded to swim and settle again amongst Acmpora coral. REMARKS. This species is endemic to tropical and warm temperate Australia, from Broome, WA to St Vincents Gul f, S A and from Eden, N S W to Caloundra, S Qld. The species is absent from the immediate area between SA and NSW (Last & Stevens, 1 994). This observation extends the N distribution of the species on the E coast and provides the first record from the Great Barrier Reef. Dasyatis leylandi Last, 1987 MATERIAL. QMI33352, immature <3, 225mm TL, 130mm DW, 108mm DL, mass 70g, and, immature <3, 315mm TL, 173mm DW, 146mm DL, mass 160g) (Fig. 1C) were captured by demersal otter trawl in Hervey Bay, 25°07'S, 152°39'E, at 1 lm depth on 14/06/2002. REMARKS. Dasyatis leylandi had previously been recorded from N Australia between Monte Bello Islands, WA and Townsville, Qld ( 1 9° 1 5'S, 146°49'E), as well as New Guinea (Last & Stevens, 1994). These records represent a significant range extension for the species on the E coast (~ 1,200km). The species appears to be relatively common on scallop trawling grounds between Hervey Bay and Gladstone (Peter Kyne, unpubl. data). Aetonivlaeus nichofii (Schneider, 1801) MATERIAL. QMI33854; 9, 343mm DW, 530mm TL, mass 450g) (Fig. 1 D) was captured by demersal otter trawl in Hervey Bay, 25°13'S, 152°43'E, at 8m depth on 11/06/2002. QM 112534, immature <3, 245mm DW, 51 4mm TL,olf Brampton Island, 20°48'S, 149°16'E, 22m, 1 3/08/1957; AMS 134389-027, immature <3, 350mm DW, 690mm TL, off Shoalwater Bay, 22°2 1 f S, 150°45'E, 43m, 25/10/1993. REMARKS. Last & Stevens ( 1 994) report that A nichofii has an Indo-West Pacific distribution from S Japan to Australia and W to India. In Australia it occurs in tropical waters from Bonaparte Archipelago, WA to Cairns, Qld (16°54’S, 145°47’E). The above specimens significantly expand its southern range (by ~ 1,600km). Myliobatis hamlyni Ogilby, 1911 MATERIAL. QMI333 1 8; 9 , 440mm DW, 945mm TL) was collected by demersal trawl by P. Duncan off Swain Reefs, 20°40'S, 151°20'E at 210-220m depth on 07/08/1999. REMARKS. Previously known only from the holotype (QMI 1567) caught off Cape Moreton, Qld (approximately 27°02'S, 153°28'E) in January' 1911 (no other collection details are available), and CSIRO H2578-01 from E of Swansea, NSW (33°05'S, 151°58'E) collected at 1 17-121 m on 09/11/1994. CSIRO H3860-01 from SW of Shark Bay, WA(26°47'S, 1 12°35'E) at 346-347m on 03/01/1991 is closely related to 326 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Mobula japonic a , AMSIB802I, whole specimen before head was removed. Note diagnostic characters of species, white tip to dorsal fin, short cephalic lobes, long tail and spiracles situated above insertion of pectoral fins (Photo: Athel D'Ombrain, courtesy of the Australian Museum, Sydney). M. hamlyni . This western form was identified as M. hamlyni in Last & Stevens (1994) but now appears to be distinct from that species (Peter Last. pers. comm.). Myliobatis hamlyni therefore appears to be restricted to E Australia. Last & Stevens ( 1 994) and Compagno &. Last (1999) reported that M. hamlyni is doubtfully distinct from the W Indian Ocean Myliobatis aquila (Linnaeus, 1758) or the W Pacific M. tobijei Bleeker, 1 854. The relationship between - the 3 species needs further assessment, as does the relationship between the east and west coast forms. Mobula japanica (Muller & Henle, 1841) MATERIAL. AMS1B8021 is the head of a specimen collected inshore by net fishennen from Lake Macquarie, NSW (32 D 59'S, 151 °35'E) on 04/04/1968. QMI33855; immature 60mm long with adjacent pinnae about 20mm apart; pinnae rachis 1mm wide reducing to 0.5mm distal ly, branching from about 70° to almost perpendicular. Rachis longitudinally striated. Pinnules variable in size and shape, sub-opposite to alternate, rounded triangular, inclined to the pinnae rachis at about 50° to nearly perpendicular, 5-(8)-10mm long and 3-(4.5)-6mm wide, closely spaced. Margins generally entire, slightly crenate in places. Pinnule apices pointed, acute to obtuse. Basiscopic margin strongly convex, decurrent at the base; acroscopic margin slightly concave, constricted at the base. Fertile pinnules smaller, apparently borne on a separate frond, stouter and rounder than the sterile pinnules, with many tending toward semi-circular. All other features agree with the sterile pinnules including venation. Pinnae >75mm long; pinnules distally merging to form a pointed acute apex. Fertile pinnules 1.5-(4)-4mm long, 2-(3.5)-4mm wide. COMPARISON. Despite poor preserv ation these specimens can be identified as Lobifolia dejerseya. Cladophlebis sp. A and Todites sp. C of Anderson & Anderson (1983) probably also belong in this species. Division PTERIDOSPERMOPHYTA Class CORYSTOSPERMALES Family CORYSTOSPERMACEAE Thomas, 1933 Umkomasia Thomas, 1933 TYPE SPECIES. Umkomasia macleanii Thomas, 1933. Umkomasia geminata (Shirley, 1898) Rigby, in Playford, Rigby & Archibald, 1982 (Fig. 3B) Beania geminata Shirley, 1898: 16, pi. 20, fig. la-c. gymnospermous seeds, in Walkom, 1917b: 16-17, pi. 20, figs 1-5 Mcgasporophyll. in Jones & de Jersey, 1947a: 56, text-fig. 52. Umkomasia sp. A, in Holmes, 1982: 17, fig. 7G. Umkomasia geminata Rigby, in Playford et al., 1982: 5, figs 1-3, 7-9. MATERIAL. Locality 2, Ipswich Basin: 14 specimens, QMF42528 - 42531 and their respective counterparts QMF42532 - 42535, 42536 - 42538, 42540 - 42543, 42544 (Fig. 3B), 42583, 42589 and counterpart 42593. DESCRIPTION. Panicle, open, with probably irregular branching. Largest specimen incomplete, 130mm long. Cupules numerous, with some in- complete specimens bearing more than 30 cupules. Rachides up to 5mm wide. Branches, up to 40mm long, 2mm wide, each bearing up to 8 cupules arranged oppositely. Cupules, elliptical, 5mm x 4.5mm to 17mm x 11mm, most swollen, suggesting an enclosed ovule, other details unclear. Pedicels up to 1mm wide. Specimen QMF42583 appears to have a juvenile, unfurling frond crozier. COMPARISON. This collection of specimens includes numerous attached cupules which display much size variation. However, there is little doubt that all these specimens belong to the same species as they are very similar in all other characteristics. Previous descriptions of this species, as listed in the above synonymy are based on fewer specimens and thus it should be expected that more natural variation be found in this larger collection. Some specimens in the present collection are larger than specimens figured previously suggesting previously examined specimens were fragmentary. Anderson & Anderson (2003) figured Umkomasia sp. 1, Umkomasia sp. 3, and Umkomasia sp. 4, all of which appear of similar morphology and size, may also belong in U. geminata. DISCUSSION. This fructification is likely to be associated with the male fructification, Pteruchus dubius (discussed below), and the foliage Dicroid- ium feistmantelii (discussed below), which all occurred together at this locality in significant numbers. The liklihood that our attribution of Dicroidium to Umkomasia follows the discovery in South Victoria Land, Antarctica, of U. uniramia by Axsmith et al. (2000) attached to fronds of Dicroidium odontopteroides (Morris, 1 845) Gothan, 1912. Pteruchus Thomas, 1933, emend. Townrow, 1962b TYPE SPECIES. Pteruchus ajricanus Thomas, 1933. Pteruchus dubius Thomas, 1933, emend. Townrow, 1962b (Fig. 3A) MATERIAL. Locality 2, Ipswich Basin: 22 specimens, QMF42545, 42547 and counterpart 42576, 42548 and counterpart 42550, 42551, 42553 - 42555, 42557 and 334 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 4. Dicroidium feistmantelii (Johnston, 1 894) Gothan, 1912, QM F42594, Ipswich Basin, Loc. 2. Scale in mm. counterpart 42558, 42559 - 42563, 42565 - 42566, 42567 and counterpart 42568 (Fig. 3A), 42569 - 42573, 42574 and counterpart 42575. Locality 3, Ipswich Basin: 1 specimen held in the private collection of A. Thomson of Brisbane, Queensland. DESCRIPTION. Fructifications large pinnatafid forming open panicules with rachis up to 1 50mm long. 4.5mm wide. Pinnules petiolate, with adjacent to sub-opposite branching at 50-80°. Petioles up to 1 3mm long, 0.5- 1 .5mm wide. Each pinnule bore a single terminal sporangial head. Some pinnules bifurcate with each branch bearing a head. Some fructifications bore >30 sporangial heads 5-3 8mm long and 5-8mm wide. Each elliptical to ovate sporangial head bore numerous (>100) pendulous sporangia crowded on the underside of the head. The sporangia are about 2.5mm long, and about 0.5mm wide, but their shape and attachment is unclear. The adaxial surface of each head has a slightly rugose texture and may have been crenate. DISCUSSION. Specimens of the ovuliferous fructification Umkomasia geminata and the foliage Dicroidium feistmantelii , which are described herein from Locality 2, are preserved along with P. dubius in significant numbers. All of these almost certainly belonged to the same plant. COMPARISON. These specimens are consistent with Pteruchus dubius as described and figured by Thomas (1933) and Townrow (1962b). A specimen identi tied as P cf. africanus by Jones & de Jersey (1947a: 55) which was discovered near our Locality 2 was referred to P. dubius by Townrow (1962b). Some of the studied specimens are larger than those described by Townrow (1962b). The wide range of sporophyll sizes is probably due to a larger number of specimens available and reflects different growth stages, as all other features are similar. The obscure specimen described as Calamostachys australis by Shirley (1898: 25, pi. 18) occurs on the same slab as Beania geminata (Shirley, 1 898: 16-17, pi. 20) which has since been recombined in Umkomasia geminata (Rigby, in Playford et al. 1982). This suggests that C. australis may be a poorly preserved Pteruchus sp. Dicroidium Gothan, 1912. emend. Townrow, 1957 TYPE SPECIES. Dicroidium odontopteroides (Morris, 1 845) Gothan, 1912. DISCUSSION. The species of this genus are highly variable in form, as illustrated by Retallack ( 1 977) and Anderson & Anderson ( 1 983). These authors have in different ways, developed nomenclatural systems that attempt to cater for this variability. However, several of the names proposed by these authors are invalid with respect to the International Code of Botanical Nomenclature. Given this, for the purposes of this work, the nomenclature of these authors is not used. This is not to suggest that the general MESOZOIC FRUCTIFICATIONS AND FOLIAGE 335 thrust of the various arguments presented by these authors is of no value. Anderson & Anderson (1983) suggested Dicroidium was a plant with a reticulate evolutionary history, which freely hybridised across Gondwanaland. They illustrated aberrant specimens of Dicroidium as evidence of hybridisation between two populations at one site. Birds River, South Africa. This appears to be the only site in Gondwanaland currently known with a strong suggestion of hybridisation. Aberrant fronds are rare elsewhere and are generally attributed to deformities because of physical damage. The issue of variability in Dicroidium species and the reasons for this requires further examination in the light of the work by Jacob & Jacob ( 1 950) who showed that preserv ed cuticle of Dicroidium fronds from the Ipswich area implied the presence of more species than is suggested by megascopic frond morphology. Our specimens have considerable morphological variation. They do not have preserved cuticle, which has been demonstrated as important con- sideration (Jacob & Jacob, 1950) when examining boundaries or continua between species. Dicroidium feistmantelii (Johnston, 1894) Gothan, 1912 (Fig. 4) MATERIAL. Location 2, Ipswich Basin - 16 specimens, QMF42539, 42546, 42549, 42552, 42556, 42564, 42577 - 42582, 42585 and counterpart 42586, 42588 and counterpart 42592, 42590, 42594 (Fig. 4). DISCUSSION. Our specimens are conspecific with D. feistmantelii as described and/or figured by Gothan (1912), Antevs (191 4), Jacob & Jacob (1950), Hill et al. (1965), Jain & Dclevoryas (1967) and Rigby (1977). They are also indis- tinguishable from Zuberia feistmantelii (Johnston, 1894) Frenguelli, 1944 as figured by Frenguelli (1944), and Thinnfeldia feistmantelii Johnston, 1 894, by Johnston ( 1 894, 1 896) and by Walkom (1917a, pL 2, fig. 2; 1924, pi. 17, fig. IF). Our specimens are preserved together with Umkomasia geminata and Pteruchus dubius , which are likely to be the ovuliferous and microsporangiate fructifications respectively, associated with this foliage. Xylopteris Frenguelli, 1943 TYPE SPECIES. Xylopteris elongata Frenguelli, 1943. Xylopteris argentina (Kurtz, 1921) Frenguelli, 1943 (Fig. 5B) MATERIAL. Location 3, Ipswich Basin: 35 specimens, QMF42405 - 424 1 3, 42420 - 42423, 42426 and counterpart 42510, 42494 - 42500, 42502 - 42505, 42511 - 42514, 42518 (Fig. 5B), 42519 - 42523. DESCRIPTION. Frond pinnate, with a leaf-like rachis, 1.5-2mm wide, which bifurcates once, occasionally twice, more or less symmetrically, at <25°. Pinnules apparently borne only on the ultimate segments of the rachis, narrowly linear, margins entire, apex acute, pointed, up to 70mm long, 0.5- 1 .5mm wide, with a faint median vein. DISCUSSION. The strap-like rachides are similar to Sphenobaiera pontifolia Anderson & Anderson, 1989, but are distinguished by their distally borne pinnules. Townrow (1962a) described 2 specimens of X. elongata (Carruthers, 1872) Frenguelli, 1943, from the Ipswich Coal Measures, which had smaller and more numerous branches positioned over the entire length of their rachides. No carbonaceous material was preserv ed in any of the specimens examined herein. Our specimens are consistent with A", argentina of Retallack (1977), and Dicroidium elongation var. argentina (Kurtz, 1921) Anderson & Anderson, 1983. The later combination may be untenable given the likely male fructification associated with X. argentina foliage, Paraxylopteris queenslandensis gen. et sp. nov. (described below), which, if attached, would preclude the combination of Xylopteris in Dicroidium. Xylopteris spinifolia (Tenison-Woods, 1883) Frenguelli, 1943 (Fig. 5A) MATERIAL. Location 2, Ipswich Basin: 3 specimens, QMF42584 and counterpart 42587. 42591 (Fig. 5A). DESCRIPTION. Frond bipinnate, up to 130mm long, with bifurcating rachides up to 3mm wide. Pinnae, up to 40mm long, branching at about 50°. Opposite to sub-opposite pinnules, up to 13mm long, branching at about 30°, with a distinct median vein. Pinnae have three or fewer pinnules per side of the pinnae midrib. COMPARISON. These specimens compare well with specimens of this species described and figured by Frenguelli (1943), Hill et al. (1965) and Retallack (1977). 336 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. A .Xylopteris spinifolia (Tenison- Woods, 1883) Frenguelli, 1943, QMF42591, Ipswich Basin, Loc. 2. B, Xylopteris argentina (Kurtz, 1921) Frenguelli, 1943, QMF42518, Ipswich Basin, Loc. 3. C, Antevsia extans (Frenguellli, 1944) Townrow, 1960, Ipswich Basin, Loc. 3. All scales in mm. Paraxvlopteris queenslandensis gen. et sp. nov. (Fig. 6) MATERIAL. Locality 3, Ipswich Basin: 1 specimen (holotype), QMF42425. ETYMOLOGY. The generic name refers to the specimen's similarity with Xylopteris , while the specific name refers to Queensland, the Australian State in which it was discovered. DIAGNOSIS. Genus monotypic. Frond pinnate consisting of a long strap-like rachis and strap- like pinnules. Sterile pinnules are grouped basally and branch probably sub-alternately. Strobulus terminal on rachis, elongate and slightly ovate. Within the strobilus, and forming the basic structure of the strobilus, the rachis thrice bifurcates (or branches) and gives rise to numerous small, probably bilateral branches, each of which bore several very small pendulous(?) sporangia. DESCRIPTION. Pinnate frond segment, 70mm long, consisting of a leaf-like rachis, 1.5mm wide, with four bilaterally branched pinnules, grouped about 40mm from the likely terminal strobilus. Slightly ovate strobilus, 21mm long and 7-9mm wide. The strap-like rachis bifurcates or branches at a very acute angle three times within the fertile section of the pinnae. Sporangia are preserved in a very crowded and confused MESOZOIC FRUCTIFICATIONS AND FOLIAGE 337 FIG 6. Paraxylopteris queens landens is gen. et sp. nov. Holotype: A, part, QMF42425; B, counterpart, QMF42415, Ipswich Basin, Loc. 3. Scales in mm. manner but appear to consist of numerous very small branches up to 9mm long, branching probably bilaterally from the strap-like rachis at about 30°. Details of these small branches are unclear but they appear to have been delicate bearing several very fine, hair-like pollen sacs probably pendulously. COMPARISON. The general form of these specimens is remarkably similar to the foliage Xylopteris argentina (Kurtz, 1921) Frenguelli, 1943, discussed above, which is abundant at this site. It seems very likely that these specimens are the male fructifications of this species, given their very similar morphology of the sterile pinnae and that they occur at the same locality. This specimen varies considerably from Ptemchus as diagnosed by Townrow ( 1 962b). It had a con- siderably less robust structure than Ptemchus , with the stobilus probably forming part of an otherwise vegetative frond. Unlike Pteruchus , which had a well developed sporophyll head (or bract) beneath which sporangial material was attached, the rachis in this specimen appears to have formed the sporophyll head. Given this apparently less well developed fructification it seems possible that this specimen represents an evolutionary point between more developed pterido- sperms and the pteridophytes. If these fructifications were attached to Xylopteris foliage then this precludes the combination of Xylopteris into Dicroidium by Anderson & Anderson (1983). PELTASPERMACEAE Thomas, 1933 Antevsia Harris, 1937 TYPE SPECIES. Antevsia zeilleri (Nathorst, 1908) Harris, 1937. Antevsia extans (Frenguelli, 1 944) Townrow, 1 960 (Fig. 5C) MATERIAL. Locality 3, Ipswich Basin: 12 specimens, QMF42402 - 42404, 42424 (Fig. 5C), 42493, 42501, 42506 - 42509, 42526 - 42527. DESCRIPTION. Sporangia symmetrical, four-lobed, peltate borne terminally on a short pedicel about 7mm long and about 0.5mm wide. The complete fructification probably bore around 1 2 such sporangia branched from a stem in an unclear arrangement. The pedicel was attached centrally to the structure with four lobes radiating from this central point, lobes elongate and slightly obovate, 6-(8)-10mm long and 2.5-3mm wide, with pointed bases and apices. COMPARISON. These specimens are almost certainly the same as those described and figured by Walkom (1915: 31, pi. 3, figs 3-4) as equi- setaceous tubers. This author believed the pedicel to be a root, however these specimens clearly show sporangia mounted terminally on a pedicel branching from a stem. Townrow (1960) combines these ‘equisetaceous tubers' in A. extans and his description and figures of this species compare well with the specimens described herein. Antevsia sp. A of Anderson & Anderson (1983, pi. 23, figs 3-4) from the Molteno Formation of the Karoo Basin, South Africa, probably also belongs in A. extans. 338 MEMOIRS OF THE QUEENSLAND MUSEUM DISCUSSION. The shape of the sporangial material, which probably had an almost circular cross-section before compression, is more indicative of a dehiscing pollen sac. There is also the possibility that these specimens were cupules which encapsulated a large seed. Only one large seed was identified with these specimens, Pteridosperm seed sp. A (Pattemore, 1998: 98, QMF42492), but its size and shape renders this interpretation most unlikely. No seeds of this size were identified in any of the studied material, nor have they been described in the literature concerned with the Carnian sediments of southeast Queensland. Townrow ( 1 960) refered A. extans , the foliage, Lepidopteris stonnhergensis (Seward, 1903) Townrow, 1956, and the ovuliferous fructification, Peltaspenmun thomasii Harris, 1937 to the same plant. Some fragmentary foliage, which were possibly referable L. stonnhergensis, were also found with the studied A. extans specimens (Pattemore, 1998: 95, QMF42427 and counterpart 42517). These fructifications are numerous and are found along with many specimens of Xylopteris argentina and a specimen of Paraxylopteris queenslandensis gen. et sp. nov., but there is no observed physical connection between these specimens. CYCADOPHYTA Genera of Uncertain Family Nilssonia Brongniart, 1825 TYPE SPECIES. Nilssonia brevis Brongniart, 1825. Nilssonia eskensis Walkom, 1928 DISCUSSION. Anderson & Anderson (1989) elected not to employ Nilssonia and questionably combined N. eskensis from the Esk Trough into Halleyoctenis multilineata (Shirley, 1898) Anderson & Anderson, 1989, which includes specimens with pinnule attachment ranging from lateral to slightly dorsal. The specimen as described and figured by Walkom (1928) has pinnules attached to the upper surface of the rachis (strongly dorsally). We consider the removal of this species to another genus to be inappropriate. Pterophyllum Brongniart, 1828 TYPE SPECIES. Pterophyllum longifolium Brongniart, 1828. Pterophyllum multilineatuni Shirley, 1898 DISCUSSION. This species known from the Ipswich Basin, with laterally attached pinnae typically found in modern cycads, was combined into Halleyoctenis multilineata (Shirley, 1898) Anderson & Anderson, 1989 and grouped with Nilssonia specimens which had dorsally attached pinnae as in Nilssonia eskensis. This combination is questioned on the grounds that lateral and dorsal pinnae attachment is of at least generic significance consistent with the description and figures of P multilineatuni of Shirley (1898) and Walkom (1917b). Yabeiella Oishi, 1931 TYPE SPECIES. Yabeiella brackebuschiana (Kurtz 1921) Oishi, 1931 Yabeiella mareyesiaca (Geinitz, 1876) Oishi, 1931 (Fig. 7A,E) MATERIAL. Locality. 3, Ipswich Basin: 1 specimen QMF42525 (Fig. 7A, E). DESCRIPTION. Leaf, narrowly elliptical, margin entire, 160mm long, up to 2omm wide with a prominent midrib, 2.5mm wide basally and tapering to 1 .5mm wide distally, lamina attached laterally to the midrib. Veins curve arcuately from the midrib then extend across the lamina at 70° without curvature, bifurcating, coalescing and rarely anastomosing, curving into a marginal vein distally. Most bifurcation and coalescence of veins occurs near the midrib or marginal vein. Venation density. 18 veins per 10mm. Marginal vein prominent about 0.8mm wide. DISCUSSION. Our specimen is identical with specimens figured by Geinitz (1876) as Taeniopteris mareyesiaca , Walkom (1917a) as Taeniopteris dunstanii , both Jones & de Jersey (1947a) and Anderson & Anderson (1989) as Y. mareyesiaca. Webb ( 1 980) identified this species from the Esk Trough. Linguifolium Arber, 1917, emend. Retallack, 1980 TYPE SPECIES. Linguifolium lillieanum Arber, 1917 Linguifolium tenison-woodsii (Etheridge, in Jack & Etheridge, 1892) Retallack, 1980 (Fig. 7C,D) Angiopteridium ensis Oldham, in Tenison-Woods, 1 883: 1 19. Angiopteridium tenison-woodsii Etheridge, in Jack & Etheridge, 1892:375. Taeniopteris tenison-woodsii: Shirley, 1898: 23, pi. 9, fig. 2. Taeniopteris tenison-woodsii: Walkom, 1917b: 32-34. Linguifolium tenison-woodsii: Retallack, 1980: 50-51, fig. 7F-H. MESOZOIC FRUCTIFICATIONS AND FOLIAGE 339 FIG. 7. A, E, Yabeiella sp. cf. sp. cf. Y. mareyesiaca (Geinitz, 1 896) Oishi, 1931, QMF42425, Ipswich Basin, Loc. 3. (E, enlarged to show venation). B, ? conifer sp. b. QMF39275, Nambour Basin, Loc. 4. C, D, Linguifolium tenison-woodsii (Etheridge. In Jack & Etheridge, 1892) Retallack, 1980, QMF42487, QMF424 1 5, Ipswich Basin, Loc. 3. Scales in mm; scale on A applies to C, D. The synonymy lists only citations necessary for the identification of the species. The Indian species described as Stengerites crisis Oldham, in Oldham & Morris, 1 863 was later used, but not recombined as Taeniopteris crisis by Feistmantel (1876). Fcistmantel (1877) recombined Oldham's specimens into Angiopteridium crisis. Tcnison-Woods (1883) identified a specimen as Angiopteridium crisis which Etheridge (in Jack & Etheridge, 1892) recognised as differing from the Indian species, and named it Angiopteridium tenison-woodsii . Shirley ( 1 898) recombined, described and figured for the first time, the Australian species as Taeniopteris tenison- woodsii Arber (1913) subsequently erected the genus Linguifolium , which was used by Retallack (1980). MATERIAL. Locality 3, Ipswich Basin: 8 specimens, QMF42415 (Fig. 7D), 42616 - 42418, 42487 (Fig. 7C), 42488, 42489, 42524. DESCRIPTION. Leaf narrowly linear, 7mm wide, with the largest incomplete specimen 65mm long, margin entire, midrib prominent, longitudinally striated, about 0.8mm wide, with lamina laterally attached. Neither leaf apices nor bases are preserved. Veins branch from the midrib at 20°-40°, simple or once forked, recurving slightly to terminate at the margin at a slightly more acute to slightly less acute angle than that made with the midrib, in places the angle between the venation and the margin is so acute as to appear to be almost forming a marginal vein. Venation density varies con- siderably between specimens from 6- 1 6 veins per 10mm. DISCUSSION. Specimens vary considerably both in terms of venation density and geometry. Most specimens are identical with: Taeniopteris tenison-woodsii (Etheridge, in Jack and Etheridge, 1892) Walkom, 191 7a as described by Walkom (1917a, 1928); Doratophyllum tenison-woodsii (Etheridge, in Jack & Etheridge 340 MEMOIRS OF THE QUEENSLAND MUSEUM 1 892) Jones & de Jersey 1 947a; and Linguifoliunt tenison-woodsii as figured by Retallack ( 1 980) also Anderson & Anderson ( 1 989). Venation in some specimens ranges to identical with that of L gracile Anderson & Anderson, 1989. Given the venation density and geometrical variation in the specimens examined herein, L. gracile is a junior synonym of L. tenison-woodsii. Zamites Brongniart, 1828 TYPE SPECIES. Zamites gigas (Lindlcy & Hutton, 1 834) Moms, 1843. Zamites queenslandi (Walkom, 1917b) Webb, ex Anderson & Anderson, 1989 (Fig. 8A) MATERIAL. Locality 1, Esk Trough: 8 specimens, QMF42338, 42339 and its counterpart 42340 (Fig. 8B). 42381 -42383,42386-42388. DESCRIPTION. Frond pinnate, >85mm long, >20mm wide, rachis 1-1. 8mm wide. Pinnae slightly oblanceolate to elliptical, 6- 13mm long, 3. 5 -4mm wide distally, 2.5-3.5mm wide near base, attached dorsally, extending from the rachis at 70-90°, closely spaced but not overlapping; most pinnae broaden distally and have slightly bulbous bases. Venation dense, approximately 5 veins per mm, dichotomising, slightly divergent. DISCUSSION. Webb (1980) referred Otozamites queenslandi Walkom, 1917b to Z. queenslandi on the ground that pinnules had poorly developed auriculate bases. This view of the difference between Zamites and Otozamites is consistent with Seward ( 1 969), and Taylor & Taylor ( 1 993 ). Anderson & Anderson (1989) formalised the combination with Zamites. Division CONIFEROPHYTA Order CONIFERALES Family VOLTZIACEAE Heidiphvllum Retallack, 1981 TYPE SPECIES. Heidiphyllum elongatum (Morris, 1845) Retallack, 1981. Heidiphyllum sp. cf. H. elongatum (Morris, 1845) Retallack, 1981 MATERIAL. Locality 1, Esk Trough: 5 specimens. QMF42332 - 42335, 42337. Locality 3, Ipswich Basin: 1 specimen, QMF42419 (?). DESCRIPTION. Leaves linear oblanceolate, some slightly falcate, length 30-1 00mm, all specimens incomplete, width up to 10mm, constricting abruptly distally to form a rounded acute apex, narrowing to about 2mm at their sessile base, margins entire, veins dichotomous and parallel for most of their course but converge slightly toward the apex terminating at the apical margin. Veins evenly spaced with a density of 1-2 per mm. Some veins may merge in the apical region close to the apical margin. DISCUSSION. Specimens are not well preserved but they conform with the description of Anderson & Anderson (1989). Specimens of Phoenicopsis elongatus Morris, 1 845, as described by Walkom (1917b, 1924) and Hill et al. (1965), probably belong here. Our specimens are compared with the species as they are fragmentary and it is possible that some of these fragments could be ginkgoalean foliage, but the very few visible apices and bases of our, and Pattemore’s (1998) specimens are indicative of Heidiphyllum. One fragmentary specimen from Locality 3, Ipswich Basin, is tentatively referred here. Family PODOCARPACEAE Rissikia Townrow, 1967 TYPE SPECIES. Rissikia media (Tenison- Woods, 1883) Townrow, 1967. Rissikia sp. cf. R. apiculata Townrow, 1967 (Fig. 8D) MATERIAL. Locality 4. Nambour Basin: QMF39268, 39270. DESCRIPTION. Foliage fragments poorly preserved with stem thickness up to 1mm, shoot length up to 50mm. Specimen QMF39268 has branched shoots apparently incomplete, 10mm long, shoot branching arrangement unclear. Leaves linear, inserted spirally 5-8mm long, < 1 mm wide, 1.5-2mm between leaves, bases slightly decurrent, no or minimal leaf-width contraction basally. Leaves covering the entire length of each fragment, branched acutely, recurved away from the stem. Cross-sectional detail unclear, leaf thickness probably 700m) rainforest at Bellenden Ker Range and Mt Bartle Frere, NE Qld. Megateg bartholoniai sp. nov. (Figs 4, 9; Table 2) ETYMOLOGY. For Dr Alan Bartholomai, Director, of the Queensland Museum from 1969 to 1999. MATERIAL. HOLOTYPE. d, Upper Cow C’k, 1.5km NE of Mt Spurgeon, 16°26'S I45°13 , E, NEQLD, 15-21 Oct 1991, GMonteith. H.Janctzki, D.Cook, L. Roberts, QM S3 1 109. PARATYPES: Allotype, 9 , as for holotype, QM S3 1 1 10. 1 6 , as for holotype, QM S3 1 1 1 1 ; 4 d d, Mossman Bluff Track, 5- 10km W Mossman, Site 8, 16°28'S 145°22’E, flight intercept trap, 1-17 Jan 1989, GMonteith, GTLompson ANZSES Expedition, QM S3 1 1 45, S3 1 1 52; 1 d , same data but 20 Dec 1 989- 1 5 Jan 1990, QM SI 6548; 2 dd, same data but 16°25’S 145°20'E, 20-24 Dec 1989, QM S31147, 31151; 1 d, same data but Site 7, 1 6°28'S 145°22'E, 16-30 Dec 1988, QM S3 11 50; 1 9 , same data but site 10, 16°39'S 145°34’E, llight trap, 17-31 Dec 1988, QM S3U33; 2 9 9 , Mt Demi, summit, 16°30’S 145°19’E, pitfall, 17 Dec 1995-25 Jan 1996. G Monteith, GThompson, Ford, QM S41358; I d, Mt Lewis, 16°35*S 145°17'E, sieved litter, 120ct 1980, GMonteith. QMS31143; I 9.MtLcwisRd, 22km from highway (Site 3), 16°35'S 1 45° 1 T E, pitfall, 18 Dec 1989-13 Jan 1990, GMonteith. GThompson, ANZSES Expedition, QM S3 11 92; 2 9 9 , Mt Lewis, 2.5km N, 16°34’S 145°16’E, sieved litter, 3 Nov 1983, D.Yeates, GThompson, QM S3 1191: 1 9 , Mt Lewis, 5.5km N, 16°34 , S 145°16 , E, sieved litter, 8 Sep 1981, GMonteith, D.Cook, QM S3 1 153; 4 d d, Mt Spurgeon, 2k SE, 16°27'S 145°I2 , E, NEQLD, 20 Dec 1988-4 Jan 1989, GMonteith, GThompson, ANZSES Expedition, QM S3 1 146, S3 1 156, S3 1 144; 1 d, Pauls Luck, Carbine Tableland, 16°27’S 145°16’E, pitfall, 28-30 Nov 1990, GMonteith, H.Janetzki, D.Cook, QM S3 11 54; 1 9 , REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 363 FIG. 6. Mcgateg , 8 palpal tibia, left. A, B. M. rcimboldi , sp. nov., dorsal (A) and ventral (B) views. C, M. covacevichae , sp. nov., dorsal view. D, F, M. Spurgeon , sp. nov., ventral (D) and dorsal (F) views. E, M paulstumkati , sp. nov., ventral view. G, H, A/, elegans , sp. nov., ventral view, paratype. Upper Boulder Creek (G); H, QM S39045. Scale line 1mm (B-G), H = 0.8mm. Platypus Ck, Pauls Luck Track, 13km W Mossman, 16°27’S 145°16‘E, pitfall, 1-16 Jan 1990, ANZSES expedition, QM S3 1193. All in NEQld and rainforest, except as noted. DIAGNOSIS. Males are unique in the large triangular thorn on the basal embolus and the large scooped tibial apophysis. Females differ from those of A/, paulstumkati in the full transverse copulatory groove. DESCRIPTION. Holotype <5 QM S3 11 09. Carapace 4.20 long, 3.32 wide. Abdomen 3.00, 2.56 wide. Total length, 7.4. Colour : carapace yellow brown with narrow dark submarginal band, darker on striae. Abdomen dorsally mostly yellow brown with dark ‘shoulders’, two pairs dark ‘eyes’, and mottled black tip above spinnerets; ventrally yellow TABLE 2. Leg measurements of Megateg bartholomai, holotype male and allotype female. Male I II III IV Palp Femur 3.01 3.01 2.92 3.46 1.46 Patella 1.15 1.15 1.15 1.31 0.69 Tibia 4.00 3.61 2.38 3.15 1.08 Metatarsus 3.92 3.00 2.77 4.08 1.23 Tarsus 1.92 1.46 1.31 1.77 Total 14.00 12.23 10.53 13.77 4.46 Female I 11 III IV Palp Femur 2.54 2.69 2.23 2.92 1.31 Patella 1.31 1.38 1.15 1.31 0.85 Tibia 2.46 2.31 1.85 2.61 0.69 Metatarsus 2.00 1.92 1.69 2.85 0.92 Tarsus 0.77 0.69 1.00 1.38 Total 9.08 8.99 7.92 11.07 3.77 364 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. Megateg ramboldi , sp. nov., 6 palpal bulb, scanning electron micrograph, ventral view. A, bulb; B, distal bulb. brown with transverse black flecks. Legs yellow brown with dark ring apically on all leg femora and 2 dark rings ventrally on femur IV and dark bands on distal tibia III, IV. Eyes : AME:ALE:PME:PLE, 10:14:8:13. Eye group front width: back width: length, 55:71:36. Interspaces: AME-AME, 0.7; AME-ALE, 0.3; PME-PME, 1.7; PME-PLE, 0.7. Chelicerae : p=3, r=3-4 teeth. Spines: I: fe pvl strong, p2d3r4; pa rl; ti p2d3r3v2.2.2.2; me p2r2v2.2.2. II: fepvl weak, p3d3r4; pa rl; ti p2d3r3v2.2.2.2; me p4r3v2.2.2. Ill: fe p4:d3r4; pa rl ; ti p2d2r2v2.2.2; me p4r4 v 2.2.2. Distal III & IV met with close paired laterals. IV: fe p4d3r3; pa rl; ti p2d2r2 v.2.2.2; me p4r5v7 unpaired. Palp fe d3rl. Legs : scopula absent or at most very thin on tarsi I. Tibial fracture I-IV distinct pro- and retro- laterally. Trochanteral notches shallow, deeper in back of notch to front, twice as wide as deep. Palp (Fig. 9A-D): tibia long, concave for length retrolaterally: basally, tibia with raised mound of 20-30 long, thick, dark, curved setae in cluster; scoop set wide, tibia distally incrassate. Tibial apophysis broad, converging slightly apically with thicker apex. Tibia with sclerotised collar opposite tibial apophysis tip and two large sclerotised collar-like processes, one distal, one retroventrally against base of cymbium. Cymbium : scopula extends over distal half; basodorsal process small and triangular, arising from discontinuously excavate surface; another triangular process retrolaterally and a small conical mound ventral of that; latter two flank a tibial collar. Paracymbial discontinuity a distinct, triangular, glabrous mound. Bulb: median apophysis small, roughly rectangular with apical hook, opposed by thin translucent short, scooped tegular vane, base irregular, large, cordate; embolus a wide, Hat flange with one of two short conical processes prolateral of median apophysis. Translucent unsclerotised process between median apophysis and embolus (in line between) and one prolateral off base of median apophysis. Tegulum extensive, a broad collar occupying ca. 300° of bulb. Allotype 9 QMS31110. As for male except as follows. Carapace 4.56 long, 3.72 wide. Abdomen 5.56, 4.52 wide. Total length, 10.4. Colour, carapace brown with irregular darker areas centrally & on margins. Abdomen like male with light flecking across abdomen. Legs extensively banded (amaurobiid basic pattern); distal and Ventral femora, lateral patellae, distal tibiae and metatarsi. Eyes : AME:ALE:PME:PLE, 11:16:7:13. Eye group front width: back width: length, 64:91:41. Interspaces: AME-AME, 0.9; AME-ALE, 0.3; PME-PME, 2.2; PME-PLE, 1.0. Chelicerae: p=3, r=3. Spines : I: fepvl strong, pld2rl; paO; ti v2. 2.2.2; me v2.2.2. II: fe pld3rl; pa 0; ti v2.2.2.2; me REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 365 FIG 8. Megateg ramboldi , sp. nov., 9 . A, epigyne. B, C, vulva, axial view from front (B), ventral view (C). v2.2.2. Ill: fe p3d3r3; pa 0; ti p2d2r2v2.2.2. me p4r4v2.2.2. IV: fe p2d2rl ; pa rl ; ti p2d3r2v5; me p5r6v6. Palp: fe pld2; pa 0; ti p2; ta p3. Legs : scopula on tarsi I, II weak. Claws short with 3-4 teeth. Tarsal rod long, in apical 1/3. Epigyne (Fig. 9F,G): a cordate plate with two narrow curved grooves; vulva a pair of spheres. DISTRIBUTION AND HABITAT. High altitude (>700m) rainforest at Mt Spurgeon, Mt Demi, Mt Lewis, Mossman Bluff, and Pauls Luck Track, west of Mossman, NE Qld. Megateg covaccvichae sp. nov. (Figs 4, 5B, 6C, 10, 11; Table 3) ETYMOLOGY. For Jeanette Covacevich. Senior Curator, Reptiles, Queensland Museum, 1966-2002. MATERIAL. HOLOTYPE: d, Mt Windsor Tbld, Whypala SF, \6°\5'S 145°02'E, notophyll vine forest, pitfall. Summer 92/93, S. Burnett, QM S24541. PARATYPES: allotype, 9 , as for holotype, QM S24549; 3 9 9 , as for holotype, QM S33 1 40, S33 1 46, S33 1 56; 1 9 , Windsor Tableland, 5.7 km past barracks, 16°14’S 145°00'E, NEQLD, rainforest, sieved litter, 23 Nov 1997, G Monteith, QM S43024. 1 9 , Mt Lewis Rd, old Barracks area, 16°35’S I45°17’E, 13 Jan 1990, ANZSES expedition, QM S3 1 1 94. OTHER MATERIAL. 5 juvs., taken with holotype, QM S32949. DIAGNOSIS. Differs from M . paulstumkati and \1. bcirtholomai in males having a much longer palpal tibia and the tip of the tibial apophysis is broadly rounded rather than a pointed taper; females differ in that the median septum ridges are clearly closer distally than proximally. DESCRIPTION. Holotype 6. Carapace 4.16 long, 3.20 wide. Abdomen 3.00, 2.56 wide. Total length, 7.5. Colour, carapace yellow brown with brown around fovea, along strial ridges and submarginally; dark vee in front of fovea, along caput edge and in diagonal line lateral of PLE. Legs with dark bands on distal femora to metatarsi and 2 extra below femora. Sternum, maxillae and labium yellow brown. Abdomen entirely darkly mottled. Eyes: AME:ALE:PME:PLE, 8:15:9:15. Eye group front width: back width: length, 53:74:35. Interspaces: AME-AME, 0.8; AME-ALE, 0.3; PME-PME, 1.5; PME-PLE, 0.7. Chelicerae : p=3, r=3. Spines : I: fe pvlp2d3r2; pa 0; ti p3d3r3v2.2.2.2; me v2.2.2. II: fe pv 1 strong, p3d3r3; pa rl; ti p3d3r3v2,2.2.2; me p 1 r2r2v2.2.2. Ill: fe pvlp3d3r4; pa rl; ti p2d2r2v2.2.2; me p4r3v2.2.2. IV: fe p4d3r3; pa r 1 ; ti p2d2r2v.2.2.2; me p5r5v7 unpaired. Palp: fe pvlpldl.2. Tibiae & metatarsi I, II with spines overlapping. Metatarsus I long, bowed. Legs: long; scopula absent. Tibial fracture I-IV distinct prolaterally. Trochanteral notches shallow, I, II & on IV, deeper on III. Tarsal rod at 3/8 from base. Tufts small, entire. 3-4 teeth on claws. RCH not evident. Palp: tibia much longer than in Af. paulstumkati ; bowed, concave, prolaterally; basal mound low, setose. Tibial apophysis a broad, blunt, Hat blade. Cymbium: scopula extent=2/3; basal cymbium prodorsally indented asymmetrically opposite more dorsal lobe on tibia, forms small, square, rounded process on retroventral comer; margin indented retrobasally, wide for most of basal half; margin open apical ly. Bulb: median apophysis a rectanguloid groove with one comer apically hooked, with irregular, extensive, rectangular base margin; embolus narrow, tapered flange; 366 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 3. Leg measurements of Megateg covacevichae , holotype male and allotype female. Male I u ni IV Palp Femur 4.08 3.54 3.15 4.00 2.07 Patella 1.31 1.38 1.08 1.15 0.92 Tibia — — 5.00 3.77 2.46 3.38 1.54 : Metatarsus 5.07 3.92 3.08 4.38 1.00 Tarsus 2.07 1.61 1.38 1.85 Total 17.53 14.22 11.15 14.76 5.53 ] Female i II III IV Palp Femur 1.92 2.31 2.15 2.54 0.92 Patella 1.23 1.31 1.00 1.15 0.61 Tibia 2.07 1.77 1.54 2.23 0.77 Metatarsus 1.77 1.38 1.77 2.69 0.85 Tarsus 0.77 0.77 1.00 1.23 Total 7.76 7.54 7.46 9.84 3.15 FIG. 9. Megateg bartholomai, sp. nov. 6 palp, A-D; 9 , E, F. A, C. D, palpal tibia, cymbium and bulb (B), ventral (A, B, D) and retrolateral view (C); E, anterior shield, abdomen, showing attachment discs (arrows). F, epigyne; G, vulva. translucent vane set just behind embolus; large, u-shaped tegulum. Allotype 9. As for male except as follows. Carapace 4.00 long, 3.20 wide. Abdomen 4.24, 3.36 wide. Total length, 9. As for male except: shorter-legged. No posterior sternum extension but post-sternal cuticle sliver is free. Legs more strongly marked (but vary to less marked in other specimens). Two dark stripes down each chelicerae. Eyes: AME:ALE:PME:PLE, 8:11:9:12. Eye group front width: back width: length, 51:85:37. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 367 FIG. 10. Megateg covacevichae , sp. nov., 6 palpal tibia (D), cymbium and bulb (B). A-C, ventral view. C, 9, epigyne; D, palpal tibia, retrolateral view. Interspaces: AME-AME, 1.3; AME-ALE, 0.6; PME-PME, 2.5; PME-PLE, 1.2. Spines'. I: fe pv 1 p 1 d2r 1 ; pa 0; ti v2.2.2.2; me v2.2.2. II: fe p2d2rl ; pa 0; ti v2.2.2.2; me v2.2.2. Ill: fe p3d3r3; pa rl; ti p2d2r2v2.2.2; me p4r4v2.2.2. IV: fe p2d3rl; pa rl; ti p2d2r2v6; me p4r3v6. Palp: fe pOdl.2; pa 0; ti p2; ta p3. Legs : scopula absent; 2-3 large teeth on claws. Rod at basal 1/3. Epigyne: ovoid with two convergent shallow grooves in V-shape; vulva simple. DISTRIBUTION AND HABITAT. High altitude (>700m) rainforest at Mt Windsor Tableland and Mt Lewis, norteastern Queensland. Megateg elegans sp. nov. (Figs 3F, 4, 5 A, 6G,H. 12, 33E,F; Table 4) MATERIAL. HOLOTYPE: 1 d t Cape Tribulation, 5km W (Site 10), 780m, 16°05 , S 145°26'E, stick brushing, 29-30 Sep 1982, GMonteith, D.Yeates, GThompson, QM S31113. PARATYPES: Allotype, 9, as above, QM S3 1 1 14. 2 9 9 , as above, QM S3 1115; 1 d, Davies Ck Rd, 17°03’S MS^’E, sieved litter, 17 Dec 1989, GMonteith, 368 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 1 1 . Megateg covacevichae , sp. nov., 9 . A. cephalothorax and abdomen, dorsal view. B, epigyne; C, vulva; D, abdomen, ventral view. GThompson, QM S3 1 134; 1 2 , Hugh Nelson Ra (GS3), 17°27'S 145°29’E, pitfall, 6 Mar-4 Apr 1995. P. Zborowski, QM S39079; 1 9. Isley Hills, 17°03 S 145°42’E, sieved litter & moss, 1 Dec 1993, GMonteith, H.Janetzki,QM S39082. Lambs Head, 17°02’S 145°39’E: 3 9 9, sieved litter (. Agathis ), 10 Nov 1981. Earthwatch, Qld Museum, QM S3 1 1 79; 1 9 . pitfall, 1 0 Dec 1 989-8 Jan 1990, GMonteith, GThompson, H.Janetzki, QM S3 1 177. Longlands Gap (BS1), 17°28’S 145°29’E; 2 9 9, flight intercept trap, 30 Nov 1995-3 Jan 1996, L. Umback. QM S39080, 39087; 1 <5, flight intercept trap. 5-27 Feb 1996, L. Umback, QM S39083. Massey Ck (BS3), 17°37’S 145°34’E, L. Umback: 2 d d +1 d. pitfall, 31 Jan-27 Feb 1996, QM S39084. S39088: 1 9, flight intercept trap, 31 Jan-27 Feb 1996, QM S391 15; 9, pitfall. 30 Nov 1995-3 Jan 1 996, QM S39095; 1 9 , pitfall, 2 Aug4 Sep 1 995, QM S39104; 9, pitfall, 4 Jul-2 Aug 1995. P. Zborowski. QM S3 9090; 2 9 9, pitfall, 6 Mar-5 Apr 1995, P. Zborowski, QM S39091, S39108; 2 9 9, pitfall. 1 Dec 1994-3 Jan 1995, P. Zborowski, QM S39121, S39122: 1 <3, 2 9 2, pitfall, 34 Feb 1995. P. Zborowski, QM S39089, S39045. 1 d, Massey Ck, 12km SW Millaa Millaa, 17°36’S 145°33 , E, pitfall, 1 Dec 1993-25 Feb 1994, J. Hasenpusch, QM S3 1 1 35; 1 9 , same data but, sieved litter, 4 May 1 983, GMonteith, D.Yeatcs, QM S3 11 22. 2 9 9, Massey Ra, 4km W, center of Bellenden Ker, 17°16’S 145°49'E, NEQLD, 9-11 Oct 1991, GMonteith, HJanetzki, D.Cook, QM S3 11 78; 1 9, Massey Ra, 17°16’S 145°49’E, sieved litter, 2 May 1983, GMonteith, D.Cook, QM S3 11 59. Mt Bartle-Frere, W Base, 17°23’S 145°46’E: 2 dd, 1 9 , fliglit intercept trap & pitfall trap, 25 Nov 1994-10 Jan 1995, G. Monteith, J. Hasenpusch, QM S3 1 137, S3 1158; 3 dd, same data but pitfall, 10 Jan-31 Mar 1995, QM S3 11 36. Mt Edith (GS2), 17°06’S 145°37’E, flight intercept trap, P. Zborowski: 1 d, 31 May-30 Jun 1995, QM S39120; 1 d, 3 Jan4 Feb 1995, QM S39078; 1 9, pitfall, 1 Dec 1994-3 Jan 1995, QM S39123. Mt Fisher (BS2), 17°34’S 145°34’E, pitfall & flight intercept trap, L. Umback: 2 dd. pitfall, 30 Nov 1995-3 Jan 1996,QM S39086, S39046; 2 d d, same locality but 1 Dec 1994-3 Jan 1995, P. Zborowski, QM S39126; 1 9, 2 Aug4 Sep 1995, QM S39094; 1 d, 2-30 Nov 1 995, QM S39 1 24; 2 9 9 , same data but litter, 27 Apr 1982, GMonteith, D.Ycates, D.Cook, QM S31125; 1 d, 1 9, Mt Fisher, Kjellberg Rd, 17°32’S 145°33’E, pitfall, 1 Dec 1993-25 Feb 1994, J. Hasenpusch, QM S31117; d, 2 9 9, Mt Fisher, Whiteing Rd, 17°33'S 145°34’E, sieved litter, 5 May 1983, GMonteith, D.Yeates, QM S3 1119, S3 1 138; 2 9 9, Mt Formartine South, 1 0km N. Kuranda, 16°43’S 145°37’E, pitfall, 23-24 Nov 1990, GMonteith, GThompson, QM S3 1 176; 1 d, Mt Haig (GS1 ), 17°06’S 145°36'E, flight intercept trap. 31 May-30 Jun 1995, P. Zborowski, QM S39125; 1 d, same data but 29 Sep-31 Oct 1995, L. Umback, QM S39092; 1 9 . Palmerston NP (NQ 11), 17°35'S 145°42'E, NEQLD, pitfall, 30 Oct 1991-24 Jul 1992, R. Raven, P.Lawless, M.Shaw, QM S24725; 1 d.Ravenshoe, \T°W S 145°29'E, 15 Jul 1976, P. Filewood, QM S31142; 1 d. Vine Ck Rd, 17°4rS 145°32’E, sieved litter, 24 Nov 1994, G Monteith, QM S3 11 39; 1 d, Malaan Ra, 2km S Palmerston Hway, 17°36'S 145°24’E, pitfall, 10 Jan-7 Mar 1995, G Monteith, J. Hasenpusch, QM S3 11 20; 1 d, Marecba, 22km SE, 17°07'S 145°36*E, sieved litter, 4 Nov 1983, D.Yeates GThompson, QM S3 1 124. All in NEQ. OTHER MATERIAL. QM S32694, QM S3 1141, QM S3 11 40, QM S20775, QM S3 1 1 2 1 , QM S3 1 1 1 8, QM S3 1 083, QM S41115. DIAGNOSIS. Males differ from those of M ramboldi in the more slender median apophysis, spine-like embolus and cluster of bristles retrobasally on palpal tibia; females differ from REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 369 FIG. 12. Megaleg elegans, sp. nov. A-F, 700m) rainforest, Kamak to Devils Thumb. 8- 12km NW of Mossman and Mt Spurgeon, NE Qld. Megateg spurgeon sp. nov. (Figs 4, 6D,F, 18, 19; Table 8) ETYMOLOGY. From the type locality. MATERIAL. HOLOTYPE: d, Mt Spurgeon, 2.5km S, 16°28’S 145°12’E, open forest, pitfall, 13-21 Oct 1991, G 376 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 18. Megateg spurgeon, sp. nov., S, palpal tibia, cymbium and bulb; retrolateral (A), ventral (B), dorsal (C) views. Monteith, H. Janetzki, QM S3 1 148. PARATYPES: 3 66 1 9. Black Mt, 16°39’S 145°29*E, 29-30 Apr 1982. G Monteith, D. Yeates. D. Cook, QM S3 1 1 55, 3 1 149; 1 6 , as for holotype. S3 11 57; 1 9, 2 700m) rainforest at Mt Finnigan and nearby Mts Hartley and Sampson, NE Qld. Krukt cannoni. sp. nov. (Figs 20, 25-28; Table 10) ETYMOLOGY. For Lester Cannon, Senior Curator of Invertebrates, Queensland Museum, 1976-2002. 382 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 22. Claw tufts, Krukt piligyna, sp. nov., $ , leg I, scanning electron micrographs. A, retrolateral view; B, axial view; C, E, ventral tapering hairs; D, scopula hairs; F, 6 tibia I photomicrograph showing crack, prolateral view. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 383 FIG 23. Krukt piligyna , sp. nov., <5, A-C, 9, D. A, palpal bulb, ventral view; B, palpal tibia and cymbium, retrolateral view; C, tibial apophysis, ventral view; D, epigync, ventral view. MATERIAL. HOLOTYPE: <3, Mt Sorrow summit. Cape Tribulation, 16°06'S 145°26’E, rainforest, sieved litter, 19 Oct 1980, G Monteith, QM S3 1390. PARATYPES. allotype 9, as for holotype, QM S3 1391; 2 9 9, as tor holotype, QM S3 1 392; V , Cape Tribulation, 3km W (Site 6), 16°05\S 145°27’E, rainforest, sieved litter. 19 Sep 1982, G Monteith, D. Yeates, G Thompson, QM S32958; 1 <5, Mt Halcyon, 16°03’S 145°25’E, pitfall, 22-24 Nov 1993, GMonteith, H.Janetzki, D.Cook, L.Roberts, QM S32959; 1 9, Roaring Meg valley, 16°04'S I45°25*E, rainforest, litter, 2 1 Nov 1993, GMonteith H.Janetzki, QM S3 2960. Mt Hemmant, 16°07'S 145°25'E. rainforest: 2 c3<3, pitfall, 25-27 Nov 1993, GMonteith, H.Janetzki, D.Cook, L.Roberts, QM S32955; 19,1 c3, sieved litter, 25 Apr 1983, GMonteith. D.Cook, QM S32954, 32953. Mt Pieter-Botte, 16°04’S 145°24’E, rainforest: 1 <3,2 9 9, pitfall, 21 Nov-8 Dec 1993. GMonteith, H.Janetzki, QM S32950; 1 <3 1 9, pitfall, 2-8 Dec 1993, GMonteith, H.Janetzki, QM S32956; 1 c3, 0.5km E, 16°05’S 145°23’E, sieved litter, 5 Oct 1982, GMonteith, D. Yeates, GThompson, QM S32957. 1 c3, ITiomton Peak, ^lO’S 145°23’E, 24-27 Sep 1984. G & S. Monteith, sieved litter & moss, 20-22 Sep 1981, GMonteith, D.Cook, QM S3 1 1 70. All in NE Queensland. DIAGNOSIS. Differs from most other species in short male palpal tibia and K. ebbenielseni and K. vicoopsae by absence of cymbial cutaway and 384 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 24. Krukt piligyna , sp. nov., 9 . A, epigyne; B, vulva. from K. ebbenielseni in normal tegulum shape and from all others in basodorsal cymbium process being hooked and from K. vicoopsae in lacking a basal tibial apophysis lobe and having the lateral epigyne lobes pointed. DESCRIPTION. Holotype 6. Carapace 3.84 long, 2.92 wide. Abdomen 2.96, 2.08 wide. Total length, 7.0. Colour, carapace yellow brown with darker areas on margins, caput, interstrial ridge posterior lateral of PLE & behind AME. Legs with 3 incomplete rings on femora, one on patellae, two on tibiae, none on metatarsi. Abdomen dorsally mottled orange with irregular ovoid pallid area anteriorly, mottling darker posteriorly. Ventrally pallid with few transverse dark areas. Sternum fawn with dark band medially and on margins; elsewhere pallid. Eyes: AME:ALE:PME:PLE, 8:6:13:11. Eye group front width: back width: length, 50:67:3 1 . Interspaces: AME-AME, 0.7; AME-ALE, 0.3; PME-PME, 1.9; PME-PLE, 1.8. Sternum : narrow, broken, ventral sternal extension. Spines : I: fe pvl strong, pld3r3; pa rl; ti p2d3r3v2.2.2.2.0; me p3r3v2.2.2. II: fe pld3r3; pa rl; ti p2d3r3v2.2.2.2.0; me p3r3v2.2.2. Ill: fe p2d3r3; pa rl; ti p2d2r2v2.2.2; me pl.2.2r2.1.2v2.2.2. IV: fe p2d3rl; pa rl; ti p2d2r2v.2.2.2; me p5r2.2.2v2.2.2. Palp: fe pldl.2, rest 0. Legs : scopula absent; tibial fracture on I-IV prolaterally and retrolaterally distinct; trochanteral notches shallow. Palp (Figs 25A-C, 26A-C): tibia ca. 1 .5 x longer than wide, barrel-like with sclerotised collar (as in all species) around distal edge; tibial apophyis a large but short twisted blunt process, retro ventrally with small separate (not on same lobe) digitiform lobe. Cymbium: scopula extends just over half; basodorsal process viewed from above (back of cymbium) a distinct triangular extension narrowing to small teat clearly hooked to retrolateral comer, below basodorsal process clearly sclerotised. Paracymbial discontinuity retrobasally with small sclerotised comer. Bulb : median apophysis base small, short, rectanguloid, converging quickly to large apical hook; embolus origin large, tapering quickly to narrow scoop. Allotype 2. As for male except as follows: Carapace 4.04 long, 3.20 wide. Abdomen 4.48, 3.16 wide. Total length, 4.8. Colour : carapace brown with dark brown markings, legs strongly banded. Abdomen dorsally dark mottled with paler oval area anteriorly, ventrally darkly mottled. Eyes: AME:ALE:PME:PLE, 9:13:9:13. Eye group front width: back width: length, 59:82:38. Interspaces: AME-AME, 1.0; AME-ALE, 0.3; PME-PME, 1.7; PME-PLE, 1.2. Spines: I: fepvl strong, pi dlrl; pa 0; ti v2. 2.2,2; me v2.2.2. II: fe p2d3rl; pa 0; ti v2.2.2.2; me v2.2.2. Ill: fe p3d3r2; pa rl; ti p2d2r2v2.2.2; me pl.2.2r2.1.2v2.2.2. IV: fe p2d2rl; pa rl; ti p2d2r2vl.2.2; me p5r2.2.2v6 paired. Palp: fe dl.2; pa 0; ti p2rl; ta p3. Legs: scopula absent; claws with 3-4 teeth; tufts united; tarsal rod at basal 2/5. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 385 FIG. 25. Krukt cannoni , sp. nov., d, palpal tibia, cymbium and bulb; dorsal (A), retrolateral (B), and ventral (C) views. Epigyne (Figs 26D, 27C-E): a broad domed central scape widening at mid-basal area and lateral grooves adjacent to diagonal ridge. TABLE 10. Leg meaurements of Krukt cannoni , holotype male and allotype female. Male I 11 III IV Palp Femur 2.31 2.23 2.31 2.61 1.08 Patella 1.00 1.08 1.00 1.08 0.61 Tibia 2.46 2.07 2.07 2.69 0.54 Metatarsus 2.31 2.15 1.77 3.38 1.00 Tarsus 1.00 0.92 1.00 1.38 Total 9.08 8.45 8.15 11.14 3.23 Female I II III IV Palp I 1 Femur 2.23 2.38 2.00 2.31 1.23 Patella 1.31 1.23 1.15 1.23 0.69 Tibia 2.15 1.92 1.54 2.38 0.77 | Metatarsus 1.85 1.77 1.92 3.08 1. 00 I Tarsus 0.92 0.85 1.00 1.15 | Total 8.46 8.15 7.61 10.15 3.69 1 DISTRIBUTION AND HABITAT.High altitude (>700m) rainforest at Mt Sorrow, Roaring Meg Valley, Mt Hemmant, Mt Pieter-Botte, Mt Halcyon, west of Cape Tribulation, and Thornton Peak, NE Qld. Krukt ebbenielseni sp. nov. (Figs 20, 28; Table 11) ETYMOLOGY. For the late Dr Ebbe Nielsen. MATERIAL. HOLOTYPE: d, Thornton Peak, 16°10’S 145°23’E, NEQLD, 24-27 Sep 1984, G & S. Monteith, QM S3 11 69. PARATYPES: 8, Thornton Peak, 16°10’S 145°22’E, 955m, Nov 1975, M. Gray, AM KS9163. DIAGNOSIS. Males are unique in the genus in the posteriorly produced but ventrally extensively unsclerotised tegulum. DESCRIPTION. Holotype 8. Carapace 3.68 long, 2.80 wide. Abdomen 2.80, 1 .92 wide. Total length, 6.8. Eyes: AME:ALE:PME:PLE, 8:11:7:12. Eye group front width: back width: length, 48:70:34. 386 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 26. Krukt cannoni, sp. nov., d, A-C, 9 D. A, palpal bulb, ventral view; B, palpal tibia and cymbium, retro lateral view; C, tibial apophysis, retrolateral view; D, epigyne, ventral view. Interspaces: AME-AME, 0.7; AME-ALE, 0.3; PME-PME, 1.9; PME-PLE, 1.2. Spines : I: fe pvl strong, pld3r2; pa rl; ti p2dlr3v2.2.2.2; me plrlv2.2.2. II: fe p2d3r3; pa rl; ti p2d3r2v2.2.2.2; me plr3v2.2.2. Ill: le p3d3r3; pa rl; ti p2d2r2v2.2.2; me p4r5v2.2.2. IV: fe p3d3rl: pa rl; ti p2d2r2v.2.2.2; me p5r6v2.2.2.2. Palp: fe pld2rl. Legs : tibial fracture prolaterally and retrolaterally distinct on I & II. Trochanteral notches: shallow, I, II deeper in back of notch to front; III, IV symmetrical. Palp (Fig. 28A-E): tibia stout with rounded distal heels prolaterally and prodorsally; tibial apophysis large, scooped with ventral corner folded. Cymbium : scopula extends to distal half; retrobasally indented: basodorsal process broad, rounded; paracymbial discontinuity absent. Bulb : median apophysis large, wide with broad, apical hook, base roughly crescentic; small then becomes extensive; embolus arises medially off prolateral side. Female : unknown. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 387 FIG 27. Krukt cannoni , sp. nov. A, B, 6 palpal tibia and cymbium base, retrolateral (A) and retrodorsal (B) views. C-E, 9 ; C, D, epigyne, photomicrograph (C); E, vulva. DISTRIBUTION AND HABITAT. High altitude Krukt megma sp. nov. (>700m) rainforest at Thornton Peak, NE Qld. (Figs 20, 29-3 1 ; Table 12) TABLE 1 1 . Leg measurements of Krukt ebbenielseni, holotype male. p - I II III IV Palp ( Femur 2.69 2.54 2.61 3.31 1.54 Patella 1.23 1.23 1.15 1.23 0.61 Tibia 2.92 2.54 2.00 2.92 0.69 Metatarsus 2.69 2.31 2.31 3.46 1.38 Tarsus 1.08 1.15 1.00 1.38 Total 10.61 9.77 9.07 12.30 4.22 ETYMOLOGY. An arbitary combination of letters. MATERIAL. HOLOTYPE: 6, Mossman Bluff Track, 5- 10km W Mossman (Site 5), 16°28'S 145°22’E, NEQLD, rainforest, pitfall, 16-30 Dec 1988, GMonteith, GThompson, ANZSES Expedition, QM SI 6650. PARATYPES. Allotype 9 , as for holotype, QM S5822 1 ; 1 6, Mossman Bluff Track, 5- 10km W Mossman (Site 1), 16°28’S 145°22'E, 250m, flight intercept trap, 1-16 Jan 1989, GMonteith, GThompson. ANZSES Expedition, QM S3 1129: I 9, same data but (Site 4). 16°25’S 145 a 20’E, 800- 1 000m, pitfall, 20 Dec 1989-15 Jan 1990, QM S32882; 3 66 1 9, same data but, site 5, 16°39’S 145°34’E, 760m, pitfall, 20 Dec 1989-15 Jan 1990, QM S3 11 32, S3 1131. 388 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 28. Krukt ebbenielseni, sp. nov., 8 palpal tibia, cymbium and bulb; dorsal (A), ventral (B, C), retrolateral views (D), tibial apophysis, retrodorsal view (E). DIAGNOSIS. Males are easily separated from those of other congeners by the very large central median apophysis; females are also easily recognised by the short wide parallel-sided scape in the epigyne. DESCRIPTION. Holotype 8. Carapace 3.52 long, 2.80 wide. Abdomen 5.00 long, 3.88 wide. Colour in alcohol. Carapace orange brown with darker margins in posterior half; centrally with reticulate dark areas forming pallid hemispheres along margin. Eye region not dark. Chelicerae yellow brown with 2 dark stripes. Abdomen dorsally yellowish with dark anterior shoulders and more mottling in posterior half with large almost entirely pallid anterior area. Carapace. AME on common tubercle over- hanging clypeus. Spines . I: fe pvlpld3r2; pa 0; ti p2dlr3v2.2.2.2; me p2r2v2.2.2. II: fe p2d3r2; paO; ti p3d2r3v2.2.2.2; me p2r2v2.2.2. Ill: fe p2d3r2; paO; ti p2d2r2v2.2.2; me pl.2.2r2. 1 .2v2.2.2. IV: p3d3rl; pa rl; ti p2d2r2v2.2.2; me p 1 p 1 p 1 p2r 1 . 1 .2.2v2.2.2. Palp: fe p 1 d 1 .2; rest, 0. Palp (Figs 29A-C, 30A-C). Tibia short, barrel-shaped with large blade-like RTA at half-length; tibia distally with collar and single dorsal lobe; collar absent from retroventral edge. Cymbium with narrow dorsal scopula; very narrow base/junction with tibia; in posterior half, cymbium narrows strongly to basodorsal overhanging process. Tegulum with small marginal basal component, distally large plate. Embolus with very large wide base, tapers quickly to narrow tip; a small triangular lamella at base of embolus; embolus entirely mobile. Median apophysis a large, curved hook narrowly attached to tegulum and mobile. Allotype 9. Colour in alcohol. Carapace like male but orange brown with more extensive darker areas. Abdomen dorsally with darker shoulders, anterior shields distinct. Legs red brown; dark bands on distal femora, tibiae and metatarsi; coxae ventrally dark distally. Bipartite dark shadow centrally on sternum with dark spots marginally opposing coxae. Eyes. AME:ALE:PME:PLE, 7:9:5:8. Eye group front width: back width: length, 41:57:25. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 389 FIG. 29. Krukt ntegma, sp. nov., scanning electron micrographs. A-C, 6 palp; A, bulb, ventral view; B, C, patella, tibia (C), cymbium and bulb (B), ventral view. D, 9 epigyne. Interspaces: AME-AME, 0.7; AME-ALE, 0.4; PME-PLE, 1.9; PME-PME, 1.6. Legs. Scopula absent. Tarsal rod at basal third. Spines. Strong proventral femoral spine on 1. 1: fe pv 1 p 1 d2r 1 ; paO; ti v2. 2.2.2; me v2.2.2. II, as I but fe p2d3r2. Ill: fe p2d3r2; paO; ti p2d2r2v2.2.2; me pl.2.2r2.1.2v2.2.2. IV: fe p2d3rl; pa rl; ti p2d2r2v2.2.2; me pl.l.l.2.r2.2.2v2.2.2. Palp: fe dl.2; paO; ti p2dl; ta p2.1. 390 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 30. Krukt megma , sp. nov., A-C, 8 palpal tibia and cymbium. A, dorsal view; B, retrolateral view; C, inclined dorsal view. Claws. Paired claws with 2-3 teeth. Palpal claw long, 5 teeth, shortest basally. Spinnerets. All on protuberant base. PMS with 1 line of 3-4 spigots dorsally and 4-6 large spigots apically. TABLE 12. Leg measurements of Krukt megma sp. nov. allotype female. I 11 m IV Palp Femur 2.24 1.92 2.08 2.52 1.16 Patella 1.20 1.16 0.96 1.00 0.44 Tibia 1.92 1.76 1.32 2.00 0.64 Metatarsus 1.56 1.48 1.64 2.72 Tarsus 0.76 0.60 1.12 1.12 0.80 Total 7.68 6.92 6.84 9.36 3.04 Epigyne (Figs 29D, 31 A-C). A low flattened plate with long biconvex grooves and small lateral cleats. Vulva similar to K. piligyna. DISTRIBUTION AND HABITAT. Mossman Bluff Track, 5- 1 0km W Mossman, in rainforest at 250- 1000m altitude, NE Qld. Krukt vicoopsae sp. nov. (Figs 20, 32A-D,F-G, 33A-D; Table 13) ETYMOLOGY. For Victoria Coops, Library Technician, Queensland Museum, 1981- 2002. MATERIAL. HOLOTYPE: <3, Mt Boolbun Sth, 15°57’S 145°08’E, rainforest, litter, 6 Nov 1995. G Monteith, QM S3 11 26. PARATYPES: allotype 9, Mt Boolbun Sth, 15°57'S 145°08'E, rainforest, 4-6 Nov 1995, G Monteith, D. Cook, L. Roberts, QM S3 1 1 28; 1 9 , same data but litter, 6 Nov 1995, G Monteith, QM S38 158; 1 9 , same data but dung, pitfalls, &. intercepts, 4-6 Nov 1995, G Monteith, QM S3 1127; 1 700m) rainforest at Mt Boolbun South, NE Qld. Birrana gen. nov. TYPE SPECIES. Birrana bulburin sp. nov. ETYMOLOGY. Aboriginal birrana, throwing stick alluding to the tarsal rod, the gender is feminine. DIAGNOSIS. Differs from Kilyana in the presence of a tarsal rod and from Megateg and Krukt in the shorter rod; males differ from those of Megateg in the short male palpal tibia and small RTA and of Krukt in the small RTA and extensive tcgulum; females differ from those of Megateg in the presence of a median scape, from those of Krukt in the absence of basolateral cleats, and from those of Huntia in having claw tufts and lacking lateral teeth. DESCRIPTION. As for species. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 395 FIG. 36. Birranci bu l burin, sp. nov., A-E, 6 . A, B, palpal tibia, cymbium & bulb; ventral (A) and retrolateral view (B); C, palpal tibia, retrolateral view; D, tarsus I showing claws & claw tufts with ventrodistal hairs. E, 9 epigyne and vulva (inset). REMARKS. Birrana is somatically very similar to Megateg but the male palpal bulb shows strong similarities to Kilyana hendersoni , sp. nov. INCLUDED SPECIES. Birrana bulburin sp. nov. Birrana bulburin sp. nov. (Figs 35-38; Table 14) ETYMOLOGY. Aboriginal word for the type locality. MATERIAL. HOLOTYPE: d, Bulburin SF, 24°3(TS 151°35’E, SE.Q, rainforest, pitfall, 1 Jun-5 Oct 1974, G& S. Monteith, QM S3 1408. PARATYPE. Allotype, 9, as for holotypc, QM S3 1409. 396 MEMOIRS OF THE QUEENSLAND MUSEUM DIAGNOSIS. As for genus. DESCRIPTION. Holotype 6. Carapace 3.48 long, 2.80 wide. Abdomen 2.68, 2.40 wide. Colour, carapace yellow brown with dark margins on undulating inner edge; central region darker with black margins laterally and posteriorly, dark margins near caput edge and diagonal from PLE. Abdomen dorsally fawn with dark shoulders, darker areas on abdomen light, mottled as pattern evident. Legs with double bands on distal femora, distal patellae, tibiae and metatarsi but bolder on III, IV. Abdomen ventrally with irregular dark flecks centrally. Sternum yellow-brown with slight radial tip shadows. Black stripes down chelicerae; reddish brown dagger mark anterior on abdomen. Eyes', almost in 3 rows, 2 4 2. AM E: ALE: PME:PLE, 6:6:8:8. Front of ALE cut back edges of AME; front edge of PLE behind back edge of PME; eyes of back row largest. AM-AM=6, AM-AL=6, PM-PM=8, PM-PL=9, AL-PL= 5. Group front width: backwidth: length, 37:47:21. ALE closer to PLE than AME. Chelicerae : r= 3 small. Spines. I : fe pvl pi d3rl ; pa rl ; ti p2r3pv5rv4; me p2r2v2.2.2. II fe, p2d3r2; pa rl; ti p2r3pv5rv4; me p3r3v2.2.2. Ill: fe p4d3r2; pa rl : ti p2d2r2v2.2.2; me pl.2.2rl.L2v2.2.2. IV: fe p2d3rl; pa rl; ti p2d2r2v.2.2.2; me pi . 1. 1 . 1 . 1 r 1 . 1 .1 .2v2.2.2. Palp: fe p Id 1.2. Legs: scopula absent. Tarsal rod at basal 1 /5th, low on I, II; raised, distinct on III, IV. Tibial crack I -I V prolateral and retrolateral distinct. Trochanteral notches shallow, symmetrical, 3 x wider than deep but becoming shallower from IV to almost indistinct on I. Claws: with 2 long and 1 short tooth on all. Abdomen: anterior face with pair of concave ‘scutes 1 . Palp (Figs 36A,B, 37A,B): tibia with only small conical mound retrolaterally, most distinct dorsally. Cymbium : asymmetrically folded to form short shallow groove on retro-apical comer; margin wide, distally narrow elsewhere with thin darkly sclerotised retromargin; probasally with distinct rounded lobe dorsal scopula for distal 1/3. Tegulum mirrored C-shape, deep basally with short thorn opposite base of median apophysis. Median apophysis a rectanguloid scoop with a small twisted pair of hooks, twisted in opposed planes with (bivalve) shell-like translucent shield at its retrobase; median apophysis free, surrounded by tegular ring distal FIG 37. Birrana bulburin , sp. nov.. A, B, 6 palpal tibia, cymbium & bulb; A, with patella, ventral view; B. retrolateral view; C, 9 vulva. of tegulum is weakly sclerotised. Subtegular tongue narrow, transverse with long sclerotised groove behind embolus. Embolus originates proapically in gradual curve to retro-corner opposite cymbial groove. Allotype 9. Carapace 4.00 long, 3.20 wide. Abdomen 4.40, 3.68 wide. Colour. As male but legs more boldly banded, most evident mottling on ventral femora. Deep Y-shaped dark mark on sternum, inner comers and edges of coxae dark. Chelicerae : 3p, 3r. Eyes: AME: ALE: PME: PLE, 4:4:6:6. Front of ALE well behind back of AME. Front edge of PLE is behind back of PME. Interspaces: AM-AM=1 .3; AM-AL=1.8; PM-PM=2.8; PM-PL=2.5; AL-PL=2.8. Group front width: back width: length, 34:44:18. Legs: scopula weak to absent on tarsi I, II. Tarsal rod low on I, a distinct lobe on IV. Claw tufts strong, similar on all. Spines: I: fe pvlpldl; pa 0; ti pv5rv4; me v2.2.2. II: as for I but fe pldl. Ill: fe p2d2rl; pa rl; ti p2dlr2v2.2.2; me p2.1.2rl.2.2v2.2.2. IV: fe p2d2r 1 ; pa rl; ti p2d2r2v5; me p 1 . 1 . 1 .2r 1 . 1 .2.2v7. Palp: fe d 1 .2; pa 0; ti p2d 1 ; ta p3dlrl. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 397 FIG. 38. Birrana bulburin , sp. nov., <3. A, tarsus I showing claws & tarsal rod (B), retrolateral view. Claws: with 2-3 short teeth on palp & legs. Epigyne (Fig. 36E): broad, ovoid with wide, transverse recurved ridges posteriorly, lateral ovoid depression and short broad posterior median ridge; internally, a short broad lobe folding back on itself. DISTRIBUTION AND HABITAT. Rainforest at Bulburin State Forest, SE Qld. CLADISTICS. Birrana is considered the sister group of Megateg and Krukt with which it shares the tarsal rod albeit clearly shorter. Huntia murrindal also possesses a tarsal rod but without males the homology of the rod cannot be established. Baehr (2003) found a similar overall pattern in Tropasteron with unresolved relationships of the Wet Tropics species having a sister group in the Eungella region. BIOGEOGRAPHY. For some spider groups, the Bulburin forests are where northern taxa reach their most southern and disjunct distribution and the northern limit of some southern taxa. Baehr (2003) found that in the Zodariidae, that point was at more northern at Eungella, west of Mackay. TABLE 14. Leg measurements of Birrana bulburin , holotype male and allotype female. Male I II III IV Palp 1 Femur 2.00 2.07 2.00 2.23 Patella 1.00 1.08 1.00 1.00 0.54 Tibia 2.23 1.77 1.46 2.00 0.54 Metatarsus 1.85 1.61 1.61 2.61 0.92 i Tarsus 0.85 0.77 0.69 1.08 Total 7.93 7.30 6.76 8.92 3.15 Female I II III IV Palp Femur 2.07 2.00 2.00 2.38 1.08 Patella 1.23 1.08 1.00 1.31 0.61 Tibia 1.92 1.69 1.31 2.00 0.69 Metatarsus 1.61 1.31 1.46 2.46 0.92 Tarsus 0.77 0.92 0.77 1.00 Total 7.60 7.00 6.54 9.15 3.30 Kilyana gen. nov. TYPE SPECIES. Kilyana bender soni, sp. nov. ETYMOLOGY. A random combination of letters; the gender is masculine. DIAGNOSIS. Differs from Krukt , Megateg , and Birrana in the absence of a tarsal rod and from Huntia Gray & Thompson, 2001 in the presence of claw tufts and only tw o claws. DESCRIPTION. As for Megateg but: Legs. Scopula present and usually distinct on tarsi I-IV of females, but only weak on metatarsi I, II. Males have scopula on palpal cymbium dorsally and in some species also tarsi. All pedal tibiae basally cracked. 2 claws; strong separate claw tufts; with additional cluster of finely fimbriate hairs in diamond-shaped area below claws. Tarsal organ set at distal quarter of tarsus, low with ovoid aperture. Bothria with 6 transverse ridges; trichobothria in single irregular line on tarsi. Spines. Females, legs I, II: tibia proventrally 5, retro ventral ly 4 thick spines on raised based; metatarsi with 3 pairs of strong spines ventrally. Male Palp. Tibia smaller than patella; tibial apophysis weak to absent, single to tripartite, sometimes simply a long deep groove, apophyses retrolateral to rctrodorsal in position. Cymbium with dorsal scopula, apically truncate and asymmetrical and forming a channel retrodistally in which embolus lies. Tegulum large, roughly mirrored L-shape and ventral. Median apophysis large, free and sometimes with conducting groove along distal edge; in some species a 398 MEMOIRS OF THE QUEENSLAND MUSEUM weakly sclerotised spine-like process arises retrobasally beside median apophysis. Embolus originates probasally as flattened cordate plate and quickly tapers to grooved whip traversing bulb but without conductor; a subtegular tongue-like conducting groove lies distal and parallel to embolus. In females, the enlarged base of the embolus can be found broken off ectally in copulatory groove. In Kilyana hendersoni , an additional sclerite, also mapping the embolus, has long filiform lateral hairs. Epigyne : basically a flattened plate with transverse copulatory groove; vulva simple C-shaped or S-shaped. Spinnerets : PMS of females dorsally with long row of spigots. Colulus broad triangular fleshy and hirsute. DISTRIBUTION AND HABITAT. Rainforests of SE Qld and N NSW. INCLUDED SPECIES (All new). K. bicarinatus ; K. campbelli ; K. corbeni ; K. dougcooki ; K. eungella ; K. hendersoni ; K. ingrami; K. krooni bit; K. lorne ; K. obrieni. CLADISTICS. Two groups are readily evident in Kilyana. The conformation of the male palpal bulbs and tibial apophyses in K. corbeni and K. ingrami are very similar: synapomorphies are the large single, scooped, sail-like median apophysis (e.g., Fig. 49A) and tripartite tibial apophysis (e.g., Fig. 49F). The second group includes K. bicarinatus, K. hendersoni , K. kroombit , and possibly K. lorne. Their synapomorphy is that the tibial apophysis is simply a long retrolateral groove. To some extent, the tibial apophyses of K. obrieni and, to a lesser extent, K. campbelli are similar in that the processes form a broad open valley which could be considered homologous with the groove. That latter wider group shares the presence of a bipartite median apophysis with the second lobe flexibly joined to the base of the main lobe. The presence of long groove on the distal edge of the median apophysis (Figs 43B, 49C,D) of K bicarinatus and K. ingrami in which the embolus lies is considered a conductor analogue and homoplasious within the group. To maintain otherwise would require many homoplasies in K. corbeni and K. ingrami which differ primarily in the presence of the groove. The tibial apophysis of Birrana is very subtle and may be taken to be a reduced form of the groove. However, a parallelism would be required to explain the tarsal rod in Birrana (albeit shorter) and Megateg plus Krukt. The form of the male palpal bulb of Birrana also shares the sausage-shaped transverse tegulum and the elongate transverse embolus. At present, these are considered parallelisms. Hence, the cladogram of Kilyana is: (corbeni-ingrami)(dougcoofd((campbelli-lome-obrieni) ( bicarinatus-hendersoni-kmombit ))). KEY TO THE SPECIES OF KILYANA Males (using palp; males of Kilyana eungella unknown) 1. Retrolateral tibial apophysis weak or a longitudinal groove (Figs 4 1C,43C,47E) 2 Retrolateral tibial apophysis with 3 strong processes, one with or without large apical spine ( Figs 45E,46C,49E) 6 2. Retrolateral tibial apophysis a groove for length of tibia (Figs41C,43C,51D) 3 Retrolateral tibial apophysis not a groove but a pair of short convergent spines separated by depression (Fig. 47E) K. dougcooki 3. Median apophysis masssive, dominant and apically bifid (Fig. 43 A) K. bicarinatus Median apophysis small, much smaller than tegulum (Figs 41 A, 51 A, 53 A) 4 4. Basal half of embolus cradled by long filamentous process (Fig. 4 IB) K. hendersoni Basal half of embolus without juxtaposed long filamentous process (Fig. 53A) 5 5. Retrolateral tibial apophysis with distal spinose process adjacent to cymbial groove (Fig. 53 D, E). . . . K.lorne Retrolateral tibial apophysis distally with truncate aspinose process (Fig. 5 1C) K. kroombit 6. Median apophysis a large scooped plate ( Figs 46A, 49A) Median apophysis not large and scooped (Figs 45 D, 54A) 8 7. Median apophysis with distal edge deeply grooved (Fig. 49C) . . K. ingrami Median apophysis without groove on distal edge (Fig. 46B) K. corbenii 8. Median apophysis a large central dominant complex process (Fig. 54A-C) K. obrieni Median apophysis a small retrolateral hook (Fig. 45A,D) K. campbelli Females (using epigyne; females of K. campbelli unknown) 1. Medial copulatory ridge wide, distinct (Figs 42 A, 44B ). 2 Medial copulatory ridge short or indistinct (Figs 52A, 55B) 7 2. Medial copulatory ridges form Vee (Fig. 44B) K. bicarinatus Medial copulatory ridge straight, recurved, or paired lateral procurved ridges 3 3. Medial copulatory ridge straight with large lateral lumens K. hendersoni Medial copulatory ridge recurved, or paired lateral procurved ridges 4 4. Medial copulatory ridge single and recurved 5 Copulatory ridges paired lateral and procurved .... 6 REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 399 5. Medial copulatory ridge deeply recurved (Fig. 46D) K.corbeni Medial copulatory ridge not so recurved (Fig. 50A,B,D) . . * K. ingrami 6. Copulatory ridges deep, form semicircles (Fig. 47F); vulva ducts convoluted (Fig. 47G) .... K.dougcootd Copulatory ridges less deep not so recurved (Fig. 48C.D): vulva ducts simply form overlapping circle (Fig. 48A.B) ...... . K.eungella 7. Medial copulatory ridge very short, a recurved circle (Fig. 52C) K.kroombit Medial copulatory ridge short, straight (Fig. 55B) K. obrieni kilyana hendersoni sp. nov. (Figs 1, 35, 39-42; Table 15) ETYMOLOGY. The specific epithet is a patronym in honour of Dr Ian Henderson, who kindly sponsored the research of the Queensland Museum. MATERIAL. HOLOTYPE: 8\ Upper Brookfield, 27°30’S 1 52°55'E, SE.QLD, rainforest, litter, 1 Nov 1981, R. Raven, V. Davies, QM S3 1340. PARATYPES: Mt Glorious, 27°20’S 152°46’E. rainforest: 1 9, sieved litter, 20 Sep 1 979, G Monteith, QM S32984; 1 9 , V.E. Davies, QM S32991; 1 9, tlight intercept trap, Jan-Mar 1982, A. Hiller. QM S32989; 8, barracks, 27 0 18'S 152°45'E. pitfall & intercept traps, 7 Dec 1991-6 Mar 1992, G Monteith, QM S43399; 2 88, 13 Apr-26 May 1983, malaise trap, A. Hiller. 8, Mt Mee, 27°03’S 152°4rE, rainforest, pitfall, 29 Nov 1991-8 Jan 1 992, D.J. Cook, QM S30305: 1 9, Mt Ncbo, 27 C 23’S 152°47'E, ex mud wasp nest, 28 Dec 1979, H. Evans, QM S32732; 1 9, Mt Nebo, 1/2 way down track in Reserve, 27°24'S 152°47’E, Araucaria notophyll vineforest, Dec 1980, A.Rozefelds, QM S39049. Upper Brookfield. 27°30 , S 152°55'E, rainforest, litter: 1 9, 12 Jan 1982,QM S32987; 1 8 1 9,9 Nov 1975-27 Feb 1976, G& S. Monteith, QM S32983; allotype 9.QMS31341; 1 8 , 14 Jul 1981 or 1 Nov 1981, R. Raven, V. Davies, QM S3 1342; 2female, 17-31 Aug 1981, R. Raven, V. Davies, QM S32985. All in SE.Q. OTHER MATERIAL: QM S53413, QM S32986, QM S3 1343, QM S32988. DIAGNOSIS. Males are easily recognised by the deeply grooved tibial apophysis and the filamentous brush paralleling the embolus; females are unusual in the large circular lateral depressions in the epigyne. DESCRIPTION. Holotype 8. Carapace 5.28 long, 3.76 wide. Abdomen 4.56, 3.12 wide. Colour. Carapace orange brown with darker ‘wedges' along striae, most evident posteriorly; hoary white hairs in band from PLE back to caput margin. Abdomen yellow brown speckled with 2 pair darker sigilla anteriorly, becoming darker brown posteriorly; ventral ly yellow brown with black hair and pigmentation medial quadrangle flanked by 6-8 small but distinct black irregular markings. Legs orange brown without darker annulations; sternum, labium and all coxae yellow to orange brown. Eyes : AME:ALE:PME:PLE, 12:12:12:14. Eye group front width: back width: length, 64:89:39. Interspaces: AME-AME, 0.8; AME-ALE, 0.6; PME-PLE, 1 .6; PME-PME, 1.1. Centres of ALE cut back edge of AME. Front edge of PLE along back edge of PME. Chelicerae : p=2-3; r=3. Spines: I: fe pvlp2d3r4; pa rl; ti p3d3r3pv5rv4; me p3r3v2.2.2. II: fe pvlp3d3r4; pa rl; ti p2d3r3pv5rv4; me p3r3v2.2.2. Ill: fe p3d3r5; pa rl; ti p2d2r2v2.2.2; me p2r3v2.2. IV: fe p4d3r2; pa rl; ti p2d2r2v2.2.2; me p3r3v2.2.2. Palp: fe pld2rl. Legs: scopula absent or at most very thin on tarsi I, II. Tibial crack I-IV prolaterally distinct; dark & grooved retrolaterally on I, II; not evident retrolaterally on III, IV. Trochanteral notches shallow, deeper in back of notch to front. Palp (Fig. 41A-C): patella incrassate with distinct prolateral mound. Tibia short with deeply intucked groove for length retrolaterally; retrobasally with scooped process, retrodistally with tapering, slender spur. Cymbium squat, almost rectangular, deep; scopula dorsal ly for distal half; basodorsal process small, triangular. Paracymbial discontinuity absent. Tegulum wide, short: median apophysis a deep, broad, scooped hook with basal fold: leaflike; hyaline process arising basally; median apophysis base large, extensive, dominates bulb. Distal to embolus a tapering process with feathery filaments for its distal length. Embolus arises beside median apophysis & distal tegulum with bulbous origin quickly tapering to long slender tip; elongate triangular tapering subtegular tongue for basal half of embolus. Allotype 9 : as for male except as follows. Carapace : 5.92 long, 4.64 wide. Abdomen 8.48, 6.32 wide. Colour : carapace like male but darker areas less distinct. Abdomen dorsally yellow brown with slightly darker areas posteriorly forming series of diamonds medially. Sternum orange brown, labium & maxillae dark brown, coxae yellow brown. Abdomen ventrally yellow brown with irregular dark medial area. Legs red brown without annulations. Chelicerae: 3p, 3r. Eyes: AME:ALE:PME:PLE, 11:14:13:15. Eye group front width: back width: length, 88:1 16:45. Interspaces: AME-AME, 1.5; AME-ALE, 1.3; PME-PLE, 2.3; PME-PME, 1 .7. Centres of ALE 400 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 39. Kilyana hendersoni , sp. nov., 9 . A, B, cephalothorax and abdomen, dorsal view; C, spinnerets, dorsal view showing PMS with biserial row of spigots dorsally; D, abdomen, ventral. behind back edge of AME. Front edge of PLE is just behind back edge of PME. Spines ; I: fe pvlpld3r2; pa 0; ti pv5rv4; me v2.2.2. II: fe pvlp2d3r3; rest as I. Ill: fe p4d3r4; pa rl; ti p2d2r2v2.2.2; me p5r4v2.2.2. IV: fe p3d3rl; pa rl; ti p2d2r2v5; me p4r4v7. Palp: fe d3; pa rl; ti p2; ta p3d 1 rl . Legs: scopula distinct on tarsi I-IV, distal but distinct on metatarsi I, II; absent elsewhere. Claws : 3 long teeth on palp & legs. Epigyme (Fig. 42A-D,F): wide short, curled hoods laterally with broad medial mound and short transverse ridge. DISTRIBUTION AND HABITAT Rainforest around Brisbane and Mt Glorious. REMARKS. Material from Mt Archer is excluded from the type series because it includes only females and is at the most outlying point. TABLE 15. Leg measurements of Kilyana hendersoni , holotype male and allotype female. Male I II III IV F= — ~ ii Palp Femur 3.77 3.77 3.46 4.08 1.92 Patella 1.77 1.85 1.61 1.54 1.08 ! Tibia 3.85 3.23 2.46 3.23 0.92 Metatarsus 3.31 3.23 3.08 4.31 1.77 Tarsus 1.46 1.38 1.15 1.69 Total 14.16 13.46 11.76 14.85 5.69 Female I 11 III IV Palp Femur 3.23 3.38 3.00 3.92 1.77 Patella 2.07 1.92 1.61 1.77 1.00 Tibia 3.08 2.69 2.15 3.00 1.00 Metatarsus 2.69 2.46 2.54 4.15 1.31 Tarsus 0.92 1.08 0.85 1.31 Total 11.99 11.53 10.15 14.15 5.08 REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 401 FIG 40. Kilyana hendersoni , sp. nov., 9 , tarsus I, scanning electron micrographs. A, B, tip showing claw tufts and ventral scopuliform hairs, lateral (A) and axial (B) views; C, ridged sclerite below paired claws, axial view; D, ventral scopuliform hairs showing smoothly tapered tip. Kilyana bicarinatus sp. nov. (Figs 35, 43, 44A-C; Table 16) ETYMOLOGY. ITie specific epithet alludes to the median apophysis of the male. MATERIAL. HOLOTYPE: c$, Bulburin SF, 24°30 , S 151°35’E, SE.Q, 25-28 Mar 1977, R. Raven, V. Davies, QM S32739. PARATYPES: allotype 9, as for holotype but 1 7-24 Mar 1975, R. Kohout, V.E. Davies, QM S53562; 2 d d, same data but 24°31’S 151°29'E, 580m, M. Gray, C. Horseman, AM KS6793. OTHER MATERIAL: 9 juv., as for holotype, QM S3 1458. DIAGNOSIS. Males resemble those of Kilyana corbeni in the flared form of median apophysis but more angular and the tibia apophysis is simple open groove; females differ in that the epigyne is medially two ridges forming a vee-shape; males and females differ from those of the sympatric Birrana bulburin in lacking a tarsal rod. DESCRIPTION. Holotype 6. Carapace 5.52 long, 4.24 wide. Abdomen 4.88, 2.80 wide. 402 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 41. Kilyana hendersoni , sp. nov., 8 palp, scanning electron micrographs. A, cymbium and bulb, ventral view; B, embolus base with filamentous brush, prolateral view; C, tibia showing groove and basal process, ventral view; D, tarsus IV showing tarsal organ (arrow gives position, inset upper right) and trichobothrial cup (inset lower left); E, prolateral cheliceral face with thickened ‘fang setae’ (inset). Colour : freshly moulted; carapace orange brown with fine dark radiating lines on caput and thorax, narrow black margin of closed semicircles; large dark bands down chelicerae; abdomen dorsally (slightly damaged) yellow brown with larger longitudinal pallid areas anteriorly forming into fine transverse lines posteriorly; venter with narrow black medial V broken by two pallid stripes (inferred from juvenile). Legs with bands, slightly paler than carapace, mottled brown under femora. Sternum yellow brown with 3 pairs dark spots opposite coxae I-III; maxillae and labium orange brown with darker central areas. Eyes : Front edge of PLE just behind back edge of PME. AME:ALE;PME:PLE, 6:6:8:8. Eye group front width: back width: length, 36:49:51. Interspaces: AME-AME. 1.2; AME-ALE, 1.0; PME-PLE, 1.7; PME-PME, 1.3. Spines : I: fe pvlpld3r4; pa rl; ti p2d3r3pv5rv4; me p3r3v2.2.1. II: fe pvlp2d3r4; pa rl; ti p2d3r3pv5rv4; me p3r2v2.2.2.1. III: fe p4d3r3; pa r 1 ; ti P 2d2r2v2.2.2; me p 1 .2.2r 1 . 1 .2v2.2.2. IV: REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 403 FIG 42. Kilyana hendersoni , sp. nov., 9 . A, B, F, epigyne; C, vulva; D, leg I, prolateral view; E, spinnerets, axial view with PLS dorsal. fe p3d3r2; pa rl; ti p2d2r2v.2.2.2; me pl.l.l.2rl.2.2.2v2.2.2. Palp: fe pldl.2. Matt of hairs on dorsal femora. Legs : scopula absent. Claws with 3-4 long, wide teeth almost concealed by tufts. Tibial crack I-I V prolateral, more distal on I, II than III, IV. Trochanteral notches shallow, slightly asymmetrical, twice as wider as deep. Palp (Fig. 43A-C): tibia short, no apophysis but retrodorsally with longitudinal keel and more entally an asymmetrical shallow trough. Cymbium : roughly rectangular with wide retrobasal edge and steep sides; prolateral paracymbial flange width forming retrodistal groove and shallow channel along basal fold; scopula dorsally for distal 1/3. Tegulum reverse L-shape, narrow basally and laterally narrow; long triangular translucent pallid flat plate near but not enclosing embolus basally. Median apophysis a large triangular plate slightly upcurved prolatcrally with sharply reflexed triangular process or retrodistal comer; distally with long deep groove, functionally a conductor. Embolus lies in groove formed by distal edge of 404 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 43. Kilyana bicarinatus , sp. nov., 6 palp A-C. A, B, bulb and cymbium, ventral (A) and prolateral (B) view; C% retrolateral tibial apophysis, retrolateral view showing groove; D. epigyne; E, vulva. median apophysis but reaching paracymbial flange. Allotype $. Carapace 5.70 long, 4.31 wide. Abdomen 5.64 long, 1.06 wide. Like Kilyana obrieni but: Colour, carapace dark orange brown with Fine dark radiating lines on caput; chelicerae dark reddish brown; abdomen dorsally fawn with no pattern evident. Legs orange brown. Eyes : lateral eyes on common tubercle; AME on distinct mound. Legs : scopula on metatarsi I, II in 3 lines; dense, uniform for length of tarsi I-IV. Spinnerets', retracted; PMS with spigots in dorsal band and apically. Epigyne (Figs 43D,E, 44B,C): wider than long with outer edges each defined by long concave ridge between which a broadly V-shaped pair of ridges converge posteriorly; vulva of two relatively large ducts overlying each other. DISTRIBUTION AND HABITAT. Rainforest at Bulburin State Forest, SE Qld. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 405 FIG. 44. Kilyana bicarincitus , sp. nov., 9 , A-C. A, cephalothorax & abdomen, dorsal view; B, epigyne; C, vulva. D, Kilyana kroombit , sp. nov., 9, maxillae and labium, ventral view. Kilvana campbelli, sp. nov. (Figs 35, 45, 46F-G; Table 17) TABLE 16. Leg measurements of Kilyana bicarinatus, holotype male. I II III IV Palp Femur 2.08 1.96 1.84 2.12 M2 Patella 1.04 1.00 0.84 0.96 0.50 Tibia 1.88 1.64 1.36 1.92 0.50 Metatarsus 1.72 1.32 1.56 2.40 - Tarsus 0.80 0.60 0.60 0.92 1.00 Total 7.52 6.52 6.20 8.32 ETYMOLOGY. For Bruce Campbell, Deputy Director, Queensland Museum, 1964-1998. MATERIAL. HOLOTYPE: d, Nimbin, 28°36’S 1 53° 13’E,NE NSW, rainforest 14Jun 1982, A.Rozefelds, D.Sinclair, QM S3 1406. PARATYPES: allotype 9, Terania Ck, near Lismore, NE NSW, 28°34’S 153°19’E, 340m, rainforest April-May 1976, M. Gray, C. Horseman, AM KS 10090; 1 9 [2 juv.], same data, AM KS 10090; 1 <3, Red scrub Flora Reserve, north of Lismore, NE NSW, 28°38’S 153°19’E, 1 Apr 1976, M. Gray, C. Horseman, AM KS 9190. 406 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 45. Kilyana campbelli , sp. nov., cJ palp, scanning electron micrographs. A, D, tibia and bulb, ventral (A) and retro lateral (D) view; B, E, tibia, retrolateral (B) and ventral (E) view. DIAGNOSIS. Resembles K. kroombit in regular outline of the unsclerotised zone around the small median apophysis but the embolus lies transverse and the tibial apophysis is a flange not a longitudinal groove; females differ in that the epigyne is two distinct strongly procurved ridges posteriorly much like Birr ana bulburin from which they differ in lacking a tarsal rod. DESCRIPTION. Holotype 6. Carapace 3.92 long, 2.96 wide. Abdomen 4.08, 2.80 wide. Colour : carapace yellow brown with fine radiating dark lines on caput, wider bands on edges and ectal edges, small dark triangle anterior to fovea. Abdomen fawn with darker areas bounded by two fine pale lines and irregular pallid area anteriorly, dark area almost entire on posterior medial abdomen; shadows ventrally on central abdomen. Legs not banded, pallid. Sternum with darkened radial pattern centrally. Eyes: AME:ALE:PME:PLE, 8:9:9:12. Eye group front width: back width: length, 5 1 :66:32. Interspaces: AME-AME, 1.0; AME-ALE, 0.7; PME-PLE, 1.3; PME-PME, 1.6. Front of ALE cut through back edge of AME. Front edge of PLE along back edge of PME. Spines: I: fe pvlpld3r4; pa rl; ti p2d2r3pv5rv4; me plrlv2.2.2. II: fe, p2d3r4; pa rl; ti p2d2r3pv5rv4; me plrlv2.2.2. Ill: fep4d3r4; pa REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 407 TABLE 17. Leg measurements of Kilyana campbelli, holotype male. I 11 III IV Palp Femur 2.92 2.92 2.61 3.38 1.38 Patella 1.38 1.38 1.31 1.38 0.61 Tibia 3.00 2.69 1.85 2.69 0.77 Metatarsus 2.92 2.61 2.46 3.61 1.46 Tarsus 1.00 1.08 1.00 1.46 Total 11.22 10.68 9.23 12.52 4.22 rl; ti p2d2r2v2.2.2; me p 1.2.2 rl.L2v 2.2.2. IV: fe p3d3rl; pa rl; ti p2d2r2v.2.2.2; me pl.l.l.2r2.2.2 v7. Palp: fe pldl.2; pa 0; ti p2. Legs : scopula absent. Claws with 2 long and 1 short basal tooth. Tibial crack on I-IV prolaterally distinct, less so retrolaterally. Trochanteral notches shallow, asymmetrical. Palp (Fig. 45A-E): patella dorsal apex a sclerotised saddle at tibial juncture. Tibia across venter with low asymmetrical mound; tibia short, incrassate with large RTA twisted ventrally truncate to give concave edge; prodorsal and distal ly a broad concave trough runs diagonally to distal dorsal comer. Cymbial scopula dorsally for distal 1/8. Cymbium almost rectangular, rounded edges with broad anterior fold and wide retrodistal groove. Prolateral paracymbial flange a distinct low triangle basally. Tegulum broad, ovoid, basally; with ovoid retrolateral window with retrolateral small claw-like median apophysis. Embolus wide, Hat, in prodistal origin reflexes back slender and slightly to base near tip of median apophysis then reflexes dorsally to lie near distal cymbial groove. Allotype 2 , like male except: Spinnerets : PMS dorsally with 2 lines each of 20-30 spigots. Epigyne (Fig. 46F,G): roughly ovoid defined with two broad U-shaped ridges converging centrally to form narrow septum which is overlaid by n-shaped ridge. DISTRIBUTION AND HABITAT. Rainforest in the Nimbin area of N NSW. Kilvana corbeni sp. nov. (Figs 35, 46A-E; Table 18) ETYMOLOGY. For Chris Corbcn and his role in the discovery of the gastric brooding habits of the frog Rheobatracus silus Liem, 1973. MATERIAL. HOLOTYPE: d, Booloumba Ck, Conondale Ra, 26°39'S 152°39’E, SE.Q, rainforest, pitfall, 29 Nov 1974-22 Feb 1975, G & S. Monteith, QM S3 1396. PARATYPES: allotype 9, Booloumba Ck, Conondale Ra (low), 26°39 , S 152°39’E, rainforest, pitfall, 29 Nov 1 974-22 Feb 1 975, G & S. Monteith, QM S3 1 397; 1 d, same data, QM S3 1398; 1 d 1 2, Conondale Ra, Sunday Ck. 26°43’S 152°34’E, rainforest, intercept flight trap, 29 Nov 1991-7 Jan 1992, D.J. Cook, QM S25182, QM S25184. All in SE.Q. DIAGNOSIS. Males differs from those of the sympatric K. ingrami in lacking the distal groove on the median apophysis, dorsal tibial spines about 1/2 lateral (cf. equal ) and tegulum has very long longitudinal component; females have the copulatory groove clearly inverted U-shaped and only about twice as wide as long whereas in K. ingrami it is broadly recurved and about 3.5 times wider than long. DESCRIPTION. Holotype d. Carapace 6.24 long, 4.96 wide. Abdomen 5.52, 3.76 wide. Colour, carapace yellow brown with broken dark areas along margins, laterally PLE on caput edge, two bands up posterior slope and triangular areas submarginally on interstriae, fovea red. Abdomen dorsally fawn with dark shoulders and small dark areas in posterior half, ventrally yellow brown with small dark areas. Femora yellow brown with broad ring at ends, tarsus yellow brown; rest reddish brown. Apical maxillae dark. Eyes : AME:ALE:PME:PLE, 12:13:12:13. Eye group front width: back width: length, 79:100:41. Interspaces: AME-AME, 1.1; AME-ALE, 0.9; PME-PLE, 2.2; PME-PME, 1.6. Front of ALE cut along back edge of AME. Front edge of PLE behind back edge of PME. Spines : I: fe pvlpld3r4; pa rl; ti p2d3r3pv5rv4; me p2r2v2.2.2. II: fe pvlp2d3r4; pa rl; ti p2d2r3v2.2.2.2.2; me p3r3v2.2.2. Ill: fep4d3r4; pa rl; ti p2d2r2v2.2.2; me pl.2.2r2.1.2v 2.2.2. IV: fe p4d3rl; pa rl; ti p2d2r2 v.2.2.1; me p4r5vl. 2.2.2. Palp: fe pldl.2. Legs: scopula absent; light pile of yellow brown hairs. Large pallid RCH. Tibial crack on I-IV distinct. Trochanteral notches shallow, sym- metrical. Palp (Fig. 46A-C): patella short, not incrassate with broad sclerotised ledge dorsodistally. Tibia: ridge joins basoventrally with low curved ridge and glabrous shallow area distally, retrolaterally with large basal process bearing large socketed truncate spine; retrodistally with narrow bluntly pointed process; mid-dorsally with bowed process bearing triangular large socketed spine much smaller than retrolateral. Cymbium apically widely folded truncate ovoid; 408 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 46. A-E, Kilyana corbeni , sp. nov., scanning electron micrographs. A-C, 6 palp. A, B, cymbium and bulb, ventral (A) and prolateral (B) view with inset showing process beside median apophysis, retrolatcral view; C, tibia and cymbium, showing tibial apophysis, ventral view. D-E, 9 ; D, epigyne, E, vulva. F-G kilyana campbelli , sp. nov., scanning electron micrographs, $ ; F, epigyne, G, vulva. retrolaterally with wide heavily sclerotised or spoon-shaped process apically twisted, angular ridge distally joining with distal fold to Embolus arises probasally with subtegular shield make short deep groove; scopula dorsally for and tegulum; origin conical, reflexes in S-shape distal 1/4; paracymbial discontinuity a slight from short basal to prolateral and emerging in extension. Tegulum large, reverse C-shaped, but long tapering tip in cymbial fold; as lor all species basal lobe more long than across basally. Median prolateral cymbial edge with shield of long curved apophysis is free of tegulum, a large open scoop setae (in right line) extending into embolus. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 409 TABLE 18. Leg measurements of Kilyana corbeni , holotype male and allotype female. Male I 11 III IV Palp Femur 4.23 4.31 3.92 4.69 2.15 Patella 2.00 2.15 1.85 1.85 1.00 Tibia 4.23 3.69 3.00 3.85 0.54 Metatarsus 4.00 3.69 3.46 4.61 2.31 Tarsus 1.54 1.38 1.31 1.69 Total 16.00 15.22 13.54 16.69 6.00 Female I II III IV Palp Femur 3.31 3.15 2.92 3.69 1.61 Patella 1.61 2.07 1.31 1.77 0.69 Tibia 2.85 2.69 2.00 3.08 0.85 Metatarsus 2.38 2.31 2.31 3.61 1.31 Tarsus 0.77 0.85 1.00 1.15 Total 10.92 11.07 9.54 13.30 4.46 Allotype 9, like male except: Carapace 5.36 long, 4.64 wide. Abdomen 8.80, 7.20 wide. Chelicerae : 3p, 3r. Eyes : AME:ALE:PME:PLE, 12:12:14:15. Eye group front width: back width: length, 80:106:44. Interspaces: AME-AME, 1.3; AME-ALE, 1 .2; PME-PLE, 2.0; PME-PME, 1 .5. Legs: scopula absent. Claws with 3 short teeth on palp & legs. Spines: 1: fe pvlpld3r2; pa 0; ti pv5rv4; me v2.2.2. II: as for I but fe pvlp2d3r3. Ill: fe p4d3r4; pa rl; ti p2d2r2v2.2.2; me pl.l.2r2.1.2v2.2.2. IV: fe p3d3rl; pa rl; ti p2d2r2v5; mepl.l.l.2r2.1.1.2 v7. Palp: fed 1.2; pa 0; ti p2dl; ta p3dlrl. Epigyne (Fig. 46D,E): a broad recurved groove; vulva G-shaped. Spinnerets: PMS each with two lines of spigots dorsal ly. DISTRIBUTION AND HABITAT. Rainforest at Booloumba Ck, Conondale Range, SE Qld, where it is sympatric with Kilyana ingrami. Kilyana dougcooki sp. nov. (Figs 35, 47; Table 19) ETYMOLOGY. For Doug Cook. MATERIAL. HOLOTYPE: <3, Upper Tallebudgera Valley, 28° 1 5 'S 153°16 , E,SE.Q, rainforest, Mar-Jul 1985, D.J. Cook, QM S3 1403. OTHER MATERIAL. QM S25073 DIAGNOSIS. Males differ from those of K. ingrami in pincer-like tibial apophysis and simple, longitudinal, hooked median apophysis. TABLE 19. Leg measurements of Kilyana dougcooki , holotype male. I II III IV Palp Femur 3.46 3.61 3.31 3.92 1.77 Patella 1.77 1.69 1.61 1.46 0.85 Tibia 3.92 3.38 2.46 3.31 0.69 Metatarsus 3.61 3.08 3.08 3.69 1.54 Tarsus 1.38 1.23 1.15 1.54 Total 14.14 12.99 11.61 13.92 4.85 DESCRIPTION. Holotype 6. Carapace 4.88 long, 3.60 wide. Abdomen 3.68, 2.64 wide. Colour: carapace orange brown with dark hairs along strial edge. Legs without bands. Abdomen fawn brown, anteriorly pallid, posterior central area with irregular dark mottling; ventral abdomen pallid with dark areas. Sternum pallid with dark radial bands. Eyes: AME:ALE:PME:PLE, 10:9:11:12. Eye group front width: back width: length, 60:83:35. Interspaces: AME-AME, 1.1; AME-ALE, 1.0; PME-PLE, 1.7; PME-PME, 1.3. Front of ALE well back from back of AME. Front edge of PLE along back edge of PME. Spines: I: fe pvlpld3r4; pa rl; ti p2d3r3pv5rv4; me p3r3v2.2.2. II: fe pvlp3d3r4; pa rl; ti p3d3r3pv5rv4; me p3r3v2.2.2. Ill: fe p4d3r4; pa rl; ti p3d2r2v2.2.2; me pl.2.2r2.2v2.2.2. IV: fe p4d3r2; pa rl; ti p2d2r2v.2.2.2; me pl.l.2r2.2.2v7, unpaired. Palp: fepldl.2. Legs: scopula absent. Claws with 2-3 small teeth; tufts united. Tibial crack I-IV prolaterally distinct, weakly discernible retrolaterally on III, IV. Trochanteral notches shallow, asymmetrical. Palp (Fig. 47A-E): Tibia: incrassate distally with raised ovoid diagonal mound retrodistally; mound with small conical reddish process turned distally toward large curved megaspine (socketed) with mounded base, midventrally with low rounded unsclerotised mound with glabrous area prolaterally. Cymbium: subovoid; retrobasally folded widely narrow distally to form shallow groove; margin wide prodistally; dorsal scopula for distal 2/5. Bulb: tegulum large, scoop-shaped on retrobasal corner; median apophysis small, roughly rectangular with small medial point and larger apical distal triangular tip; embolus originates on prolateral edge tapers quickly into smooth curving tip terminated near cymbial groove. 410 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 47. Kilyana dougcooki , sp. nov. A-C, 6 palp, scanning electron micrographs; A, cymbium and bulb, ventral view; B, tibial apophysis, retrolateral view; C, median apophysis, ventral view. D-E, 6 palp; D, E, tibia, cymbium and bulb, ventral (D) and retrolateral (E) view. F-G, $ ; F, epigyne, G, vulva. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 411 DISTRIBUTION AND HABITAT. Upper Tallebudgera Valley and probably also Mt Tamborine, in rainforest. REMARKS. Because the female and male have not been taken at the same locality and the two localities (Mt Tamborine, Upper Tallebudgera Valley, respectively), the female is not designated a paratype but the epigyne is figured (Fig. 47F,G). Kilvana eungella, sp. nov. (Figs 35, 48; Table 20) ETYMOLOGY. A noun in apposition taken from the type locality. MATERIAL. HOLOTYPE: 9, Broken R, Eungclla NP, 21°irS 148°3rE, MEQLD, rainforest. 4 Sep 1988, R. Raven, J. Gallon, T.Churchill, QM S13870. PARATYPES: 9 , Pease's Lookout, Eungella NP, 2 1°07’S 148°31'E, rainforest, pitfall & intercept traps, 17 Nov 92-mid Apr 93, GMonteith D.Cook, QM S3 1404; 1 9, Eungclla (schoolhouse), 21°08’S 148 0 29'E, rainforest, pitfall, 11-14 Feb 1986, R.Raven, J.Gallon,QM S29310;2 9, Mt William, Dalrymple Heights, 21°()rS 148 0 36’E, 1120m, rainforest, Apr 1975. M. Gray, C. Horseman, AM KS6565. All in MEQ. OTHER MATERIAL. AM KS6383. DIAGNOSIS. The paired broadly procurved copulatory grooves in the female are unique in the genus. DESCRIPTION. Holotype 9. Carapace 5.12 long, 4.00 wide. Abdomen 7. 12, 5.36 wide. Colour, carapace, legs and abdomen yellow brown. Carapace with darker radial pattern. Chelicerae without stripes. Abdomen fawn with slightly darker shoulders. Legs 111, IV with distinct bands and mottling sternum with slightly darker radial lines. Chelicerae: 3p, 3r, all large. Eyes. AME:ALE:PME:PLE, 10:11:13:12. Eye group front width: back width: length, 73:103:36. Interspaces: AME-AME, 1.8; AME-ALE, 1.1; PME-PLE, 2.6; PME-PME, 1 .8. Front of ALE behind back edge of AME. Front edge of PLE is just behind back edge of PME. TABLE 20. Leg measurements of Kilvana eungella , holotype female. I II III IV Palp Femur 2.77 2.61 2.54 3.15 1.38 Patella 1.00 1.54 1.38 1.38 0.69 Tibia 3.31 2.15 1.69 2.54 0.85 Metatarsus 2.23 2.07 2.07 3.31 1.15 Tarsus 0.85 0.85 0.61 1.15 Total 10.16 9.22 8.29 11.53 4.07 Spines : I: fc pvlpldl; pa 0; ti pv5rv4; me v2.2.2. II: as for I butfepvlp2d3r3. Ill: fep3d3r2;pa0;ti P 2d2r2v2.2.2; me pl.2.2r2.1.2v2.2.2. IV: fe p3d3rl; pa rl; ti p2d2r2v5; me pl.l.l.2r 2.2.2v2.2.3. Palp: fe dl.2; pa 0; ti plrl; ta p3dl. Legs : scopula very weak on metatarsi, tarsi I, II. Claws: 2-3 short on palp & legs. Trochanteral notches very shallow. Epigyne (Fig. 48A-D): a broad shallow ovoid plate with 2 distal smoothly curving groove leading to spiralled spermathecae. Spinnerets : PMS each with a long dorsal ridge. Colulus a triangular plate. DISTRIBUTION AND HABITAT. Rainforest on the Eungella Range, west of Mackay, mid E Old. REMARKS. As most of the material has 3 teeth retrolaterally on the chelicerae and only one has 4 teeth (QM S3 1340) but the cpigynes & vulva of both are alike, the quadridentate condition is considered an intraspecific variant. The vulva of QM S3 1 304 are relatively slightly longer than the holotype (Fig. 48B). Kilvana ingranii sp. nov. (Figs 35, 49, 50; Table 21) ETYMOLOGY For Dr Glen Ingram. MATERIAL. HOLOTYPE: d, Conondale Ra, 26°45'S 152°37'E, SE.Q, 1-3 May 1976, R.J. Raven. QM S31393. PARATYPES. d, Booloumba Ck, Conondale Ra, 26°39'S I52°39'E, rainforest, litter. 13-18 May 1976, R.J. Raven, QM S3 1395; Allotype 9, Conondale Ra, 26°45'S 152°37’E, 1-3 May 1976, RJ. Raven. QM S3 1394; 1 d [4 juv.], same data, QM S29345; 1 d. Little Yabba Ck, 26°37'S 1 52°4 1 'E, rainforest, pitfall, 10 Aug-9 Nov 1974, G& S. Monteith, QM S3 1399; 5 d, Mapleton Falls NP, 26°38'S 152°5rE, rainforest, flight intercept trap, 8 Jan-3 Mar 1992, D.J. Cook, QM S39589; 1 d 1 9, Tungi Ck, 26°40’S 152°28'E, rainforest, pitfall, 10 Nov-29 Dec 1 974, G& S. Monteith, QM S3 1 407; 1 9 . same data but 1 8 Dec 1996-20 Jan 1997, G Monteith, QM S39093; 1 9, Cold Ck, SE. QLD, 26°28*S 152 0 4TE, 122m, 16 June-23 Aug 1975. GB. & S.R. Monteith, QM S5341 1; 2 9, same data but 31 Dec 1974-27 Mar 1975, QM S 53410; d, Amamoor Ck, 26°24'S 1 52°36’E, 1 20m, rainforest, pitfall trap, 24 Sep 2001-15 Jan 2002, GB. Monteith, QM S54301; 9, Dingo Ck, via Traveston, 26°20'S 151.52’E, SE.Q, 9 Nov-3 1 Dec 1 974, G& S. Monteith, QM S54302. All in SE.Q. OTHER MATERIAL. QM S25200. DIAGNOSIS. Males differ from those of Kilvana corbeni in having a distinct groove across the distal median apophysis of the palp; females differ in that the copulatory groove is broadly recurved and about 3.5 times wider than long 412 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 48. Kilyana eungella , sp. nov., 9; vulva, QMS 13870 (A), QMS31404 (B); epigyne QMS 13870 (C), QMS3 1404(D). whereas in Kilyana corbeni it is clearly an inverted U and only about twice as wide as long. DESCRIPTION. Holotype 6. Carapace 5.52 long, 3.76 wide. Abdomen 4.72, 3.44 wide. Colour, carapace orange brown fine darker margins and along caput edge. Abdomen dorsally pallid with black rings at base of setae, darkness increases in back half. Legs not banded. Sternum with slightly darker areas opposite intercoxal comers; maxillae and labium anterior laterally dark. Abdomen ventrally is pallid with black transverse flecks. Eyes : front edge of ALE along back edge of AME. Front edge of PLE along back edge of PME. ALE clearly smallest. ALE & PLE on common tubercle. AME:ALE:PME:PLE, 11:10:13:15. Eye group front width: back width: length, 63:87:40. Interspaces: AME-AME, 1.1; AME-ALE, 1.0; ALE-PLE, 0.0; PME-PLE, 1.6; PME-PME, 1.3. Chelicerae : 3p, 3r. Spines : I: fe pvlpld3r4; pa rl; ti p2d3r3pv5rv4; me p5r4rv2.2.2. II: fe vlp2d4r4; pa rl; ti p2d3r2v2.2.2.2; mep5r4v2.2.2. Ill: fep4d4r3;pa r 1 ; ti P 2d2r2v2.2.2; me p 1 . 1 .2r2.2. 1 v2.2.2. IV: fe p4d3rl; pa rl; ti p2d2r2v.2.2.2; me pl.l.l.2r2.2.2. Palp: fe pldl.2, pa 0, ti pi. Legs : scopula absent. I, II laterigrade. Tibial crack on I-IV grooved; 2-3 teeth on claws. Trochanteral notches shallow, (3-4 wider than deep) deeper in back of notch to front. Setation on legs, sternum, maxillae and labium short, sparse. Palp (Fig. 49A-F): tibia stout, retrolaterally con- cave, glabrous with 4 processes: basoventrally a rounded diagonal ridge, retrodistally a flattened hand-shaped process; two very large modified TABLE 21. Leg measurements of Kilyana ingrami , holotype male and allotype female. Male I II HI IV Palp Femur 3.77 3.85 3.46 3.85 1.92 Patella 1.92 1.85 1.61 1.69 0.85 Tibia 3.69 3.23 2.31 3.46 0.85 Metatarsus 3.77 3.00 2.77 4.38 2.07 Tarsus 1.38 1.23 1.23 1.38 - Total 14.53 13.16 11.38 14.76 5.69 Female I 11 III IV Palp Femur 3.15 2.54 2.85 3.31 1.46 Patella 1.92 1.69 1.31 1.85 0.92 Tibia 2.77 2.38 1.69 2.77 0.92 Metatarsus 2.54 2.31 2.23 3.85 1.23 Tarsus 0.92 1.00 0.92 1.23 - Total 11.30 9.92 9.00 13.01 4.53 REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 413 FIG 49. Kilyana ingrami , sp. nov., 6 palp, scanning electron micrographs. A, C, cymbium and bulb, ventral (A) and prolateral (C) view; B, D, median apophysis, with distal groove (arrow), ventral view. E, F, 6 palpal tibia, ventral (E) and axial view looking to base (F). 414 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 50. Kilyana ingrami , sp. nov., 9, A, B, D, epigyne, showing variability in transverse groove; C, vulva. spines retrobasally, dorsal spine short conical, broad; retrolateral a wider spine but diagonally truncate to base giving concave ovoid apex, dorsal surface convex; tibia excavate between mega- spines and cymbium. Cymbium: scopula extent apical 1/3; dorso-basally with very sclerotised collar: dorsally with large basal flattened area; apical ly folded to make broad tip and retrolateral groove apically. Tegulum large basal and retrolateral ‘mirror C’ shaped, subtegular shield arises up beside cymbium on prodorsum. Embolus S-shaped, basally small, probasal with long rectangular flange, broken paraembolic process passes ventralTy then reflexes forward arising near cymbial groove with flared tip. Median apophysis large, sclerotised, triangular with two flanges on each side, all converge apically. Allotype $. As for male except as follows: Carapace 5.68 long, 3.92 wide. Abdomen 5.28, 3.76 wide. Carapace : Markings on lateral cephalothorax darker; rings on distal femora- metatarsi; pilosity like male but hairs darker. Chelicerae : 3p, 3r. Eyes : AME:ALE:PME:PLE, 10:1 1:12:16. Eye group front width: back width: length, 79:104:43. Interspaces: AME-AME, 1.3; AME-ALE, 1 .4; PME-PLE, 2.7; PME-PME, 1 .8. Front edges of ALE behind back edge of AME. Front edge of PLE is behind back edge of PME. Spines : I: fe pvlpld3r3; pa 0; ti pv5rv4; me v2.2.2. II: as for I but fcp3d3r3. Ill: fep4d3r4; pa rl; ti p2d2r2v2.2.2; me pl.2.2r.2.2.2v2.2.2. IV: fe p3d3rl; pa rl; ti p2d2r2v2.2.2; me p 1 . 1 . 1 .2r2.2.2v2.2.2. Palp: fe d 1 .2; pa 1 ; ti p2dl ; ta p3r3. Legs : scopula on tarsi I, II; weak and weak in distal third of metatarsi I. II. Epigy me (Fig. 50A-D): a broad excavate shield- shaped plate, centrally with wide inverted U-shaped ridge with recurved end; vulva Ci-shaped. DISTRIBUTION AND HABITAT.Rainforest in the Conondale Range, SE Qld. REVISIONS OF AUSTRALIAN GROUND-HUNTING SPIDERS 415 Kilyana kroombit sp. nov. (Figs 35, 44D, 5 1 , 52; Table 22) ETYMOLOGY. A noun in apposition, from the type locality. MATERIAL. HOLOTYPE: 1 ?????00000070100?1???001000000011110000000010000000001 0000000010000?000001000?0100?000000100000001111000??00010000??0001 lll^OOlOl 100072 71 10001 101 117777772021200007000001 170? 1011 100770101 11 1700101 10007271 10001 101 117777772021200007000001 1207101 1000770101 001 700001 0000 700000 100 100 100702 7700002 10007000001 100 11 02 1001 770700 101 700 10000010 7700000007000201 777000000100? 700001 07001 01 1000 7 70720 101 700 1000001 0770000000 70002 00001000000 100 770000102 071 01 1000770720 101 1001010001 077000000100100701? 72021201 0077000010201 101 1001770120 11 170010000010770001000700007000000000000077000011007101 1101770102 11 07001 10000070000001007000200000001000000 7000001 1107101 1201 770701 01 17000000000700000010070200702 7 700001 110071 001 71 10070751000? 70701 0100000000001077000000070100717770001001 1001001 01 10010151200770100 111100000000070100001007030170010000000100210011111010151100000101 111 1000000000 701 0000 100700017001 0000000 10021 00111170 10 151100000 101 111 10000010007010000011001 17777770000001002 100111 10011121201000111 11 1 10000010007010001100701 1777777002000100210011 11301 1121201000111 111100010100070100001007010170001002000100210011113011121201000111 11110000010007010001 100704017177700200010021001 11 13011 1212010001 11 11 11000100001 17700000107000070277002000100210001 11 101011 10010001 11 0111000000177777700010070000702770020001002000011100111 11000000111 000000000001 107710000007000210177002000110110010100011111001000020 100 7000000001 1770000000 70002017? 7131 000 100 71 00001 11 01 102 11 00000120 110100000000 11 7 70000000 7000200000131 000 100 71 00001 1701 102 11 00000 120 111 00000100011 7 700000007020200010020000000220000111010011000000720 11 100000000111 77000000070202100100200000002700001 10011011000770720 11 100000000 11 177000000070302100100200000002 700001 17? 11011000 7 70 720 0007000010000700000000070117777771010001101 1000010010011 1010001021 0000000010000700000001110100717770000201000100101 11001 000000000020 000000001 0000 70000 100 11 011 007 17771 0002 11 0001 000001 01 0001011 110 1020 00000000100007000011 01 101 10071 777010001 10001000000010001 021 0101020 000001001 100071 7021001 101 10070177000010100010000000101050200010020 00001 1001 00007 1702 1001 10 110221 7 770000 10 10001 00000001 0001 000 10 10020 0000100010000727121001101 10070177020010101010000010100000001010020 0000100010000727121001 11110070177020010101010000010101000000010020 0000000000001 177000000070002001770210001101 100101 10001 131200000020 0000000100001 1770001000702020017702 10101111 100101 10001 131200770020 00000010 1017? 77770000007010071 777201000100000100000100000000001001 0000000010000700000000070117777770010000000101000001000002 10001021 0000001000000700000000070002001 77002000100000000000011 141101000120 00 1000 100000072 7000001 1101 020 17770020001 00000000 101 01 11 3100 1000020 000000001 00007 11 1201001 10 101 71 77700200000001 0000 11 0001 000000000020 00000000 10000711 000001 100 1007177700200010001 00101 10001000000000020 00000000100007000000001 10 10071 7 77000000 11011001 01 1000 1000200000020; proc / ; TUARANGISAURUS AUSTRALIS SP. NOV. (PLESIOSAURIA: ELASMOSAURIDAE) FROM THE LOWER CRETACEOUS OF NORTHEASTERN QUEENSLAND, WITH ADDITIONAL NOTES ON THE PHYLOGENY OF THE ELASMOSAURIDAE SVEN SACHS SACHS, S. 2005 01 10: Tuarangisaurus australis sp. nov. (Plesiosauria: Elasmosauridae) from the Lower Cretaceous of northeastern Queensland, with additional notes on the phylogeny of the Elasmosauridae. Memoirs of the Queensland Museum 50(2): 425-440. Brisbane. ISSN 0079-8835. The skull and associate cervical vertebrae of an elasmosaur from the Lower Cretaceous of northeastern Queensland are described as a new species of Tuarangisaurus , Tuarangisaurus australis. They represent the oldest record of that genus and the" first secure outside New Zealand. A phylogenetic analysis based on 1 9 characters of the skull and cervical vertebrae is undertaken. One tree is presented, including all valid members of the Elasmosauridae for which sufficient cranial material is known. The results give an example for the interrelationship of the Elasmosauridae and show the probable position of Tuarangisaurus. □ Plesiosauria, Elasmosauridae, Tuarangisaurus, Lower Cretaceous, Albian, Queensland. Sven Sachs, Institutjur Palaontologie, Freie Universitdt Berlin, Malteser Strasse 74- 1 00, Haus D, 12249 Berlin, Germany ; 1 3 July 2004. The plesiosaurian Elasmosauridae is known from the Lower Jurassic to the Upper Cretaceous, during which time it had an almost worldwide distribution. Most taxa referred to this group had an extremely elongated neck and a proportion- ately small head. In Australia, only a small number of elasmosaur remains are known (Cruickshank et al., 1999; Rear, 2003). Among them, there arc 4 rather complete specimens, of which 2 are represented by about 80% of the postcranium. The first of these has been excavated by a field-party of the Queensland Museum at the Walsh River (Doncaster Member, Wallumbilla Formation, Late Aptian) west of Chillagoe (NE Queensland); the second one was found at Grampien Valley (Doncaster Member, Wallumbilla Formation, Late Aptian) near Richmond (N Queensland) (Sachs, 2004). Another specimen from Andamooka in the South Australian Museum, Adelaide is about 50-60% complete and consists of opalised bones (Rear, pers. com. 2001). The fourth more complete specimen is the holo- type of Woolungasaurus glendowerensis Persson ( 1 960). The ty pe material was discovered in 1 89 1 (QMF3567) and 1 893 (QMF3568) at Glendower Station near Hughenden and has been restudied (Sachs, 2004). Another specimen Persson ( 1 982) referred to Woolungasaurus is a skull that is broken into an anterior and posterior part. The posterior portion was discovered first (by T. Noonan) along with a number of vertebrae and was donated to the Geological Survey of Queensland (GSQF10552, F10550 a+b and F 1 055 1 ). Later, in 1 976, a field party comprising R. Molnar, R. Thulbom and M. Wade discovered the anterior part of the skull. This specimen was first catalogued at the Australian Museum in Sydney (old catalogue number AM F60056), but has since been transferred to Brisbane where both parts were donated to the Queensland Museum. Other probable elasmosaur remains were found in the Northern Territory (Bathurst Island Formation, Albian, sensu Murray 1987) and in Western Australia (Colalura Sandstone, Bajocian, sensu Cruickshank & Long, 1998 and Molecap Greensand, Cenomanian-?Santonian, sensu Teichert & Matheson, 1944). A greater number of elasmosaur specimens, including relatively complete skeletons, are known from New Zealand (Hector, 1874; Hiller et al, 1997; Owen, 1861, 1870; Seeley, 1877; Welles & Gregg, 1971; Wiffen & Moisley, 1 986). On the basis of this material Mauisaurus haasti Hector, 1 874 and Tuarangisaurus keyesi Wiffen & Moisley, 1986 were erected. In general the remains from New Zealand are younger than the Australian ones. Mauisaurus is known from the Conway Formation (Maastrichtian), while Tuarangisaurus are from the Tahora Formation (Mid-Campanian). In this paper the skull described by Persson (1982) shall be redescribed and classified. Based on this specimen and its associated postcranial elements, 19 phylogenetically interesting features 426 MEMOIRS OF THE QUEENSLAND MUSEUM are proposed. An analysis of these characters within the Elasmosauridae and Muraenosaurus as outgroup member was undertaken. ABBREVIATIONS. Institutional Abbreviations. AM, Australian Museum, Sydney; ANSP, Academy of Natural Sciences, Philadelphia; GPIM, Geologisch-Palaontologisches Museum, Munster; GSQ, Geological Survey of Queensland, Brisbane; MLP, Museo de La Plata, La Plata; NZGS, New Zealand Geological Survey, Lower Hutt; QM, Queensland Museum, Brisbane; UCMP, University of California Museum of Paleontology, Berkeley; USNM, United States National Museum of Natural History, Washington D.C. Anatomical Abbreviations, aac, atlas-axis- complex; aar, angular-articular complex; aas, angular- articular suture; an, angular; bo, basioccipital; bm, bite mark; d, dentary; cn, external naris; exo, exoccipital; fir, frontal; gf, glenoid fossa; hy, hyoid; ipf, interpterygoid fenestra; j, jugal; mt, mandibular tooth; mx, maxilla; mxp, maxillary process; mxt, maxillary tooth; ns, neural spine; pa, parietal; pb, parietal bridge; pi, palatine; pm, premaxilla; pmr, premaxillary ridge; pmt, premaxillary tooth; po, postorbital; prf, prefrontal; ps, parasphenoid; pt, pterygoid; rp, retroarticular process; q, quadrate; qc, quadrate condyle; sa, surangular-articular; sq, squamosal. SYSTEMATICS SAUROPTERYGIA Owen, 1860 PLESIOSAURIA De Blainville, 1835 Superfamily PLESIOSAUROIDEA Nopcsa, 1928 Family ELASMOSAURIDAE Cope, 1868 Tuarangisaurus Wiffen & Moisley, 1986 TYPE SPECIES. Tuarangisaurus keyesi Wiffen & Moisley, 1986. DIAGNOSIS. See Wiffen & Moisley (1986: 207). Tuarangisaurus australis sp. nov. Woolungasaurus cf. W. glendowerensis Persson. 1982: 647-655. “ Woolungasaurus " Thulbom & Turner, 1993: 489ff. Elasmosaurid: Kear. 2003: 288f. ETYMOLOGY. Latin australis, southern; refers to the southern continent Australia. MATERIAL. Holotype. QMF1 1050, an almost complete, but crushed skull and mandible, with attached atlas-axis complex; QMF12216-19, an anterior cervical vertebra; QMF12217, 2 associated posterior cervical vertebrae. FIG 1 . Map of the wider Julia Creek area, showing the locality of QMF1 1050 at Yambore Creek. TYPE LOCALITY. (Fig. 1) Yambore Creek, N of Julia Creek, N Queensland, Toolebuc Fonnation, Wilgunya Group, Albian (Senior et aL, 1978). DIAGNOSIS. Premaxillae forming narrow dorsal keel along midline bearing 5 teeth each; last premaxillary tooth at premaxillary-maxillary suture; posterior maxillary process following jugal over entire length and terminating at the level of last dentary tooth; jugal long, low; anterior margin of dentary forming a beak; mandibular symphysis short, expanded to level of dentary tooth four; dentary with 18 teeth per ramus; last dentary teeth more prominent than last maxillary teeth; retroarticular process short relative to jaw length; atlas-axis-complex short and high, ventrally bearing prominent sagittal crest: anterior cervical vertebrae shorter than high. DESCRIPTION. As mentioned before, the skull consists of two pieces, and is broken just anterior of the orbita. It is almost complete, but compressed dorsolaterally, so that only the left side is well-preserved (Figs 2, 3, 6). Parts of the skull, such as the skull roof and the occipital region could not be described properly as the specimen is badly crushed. The anterior portion of the skull, as well as most of the right mandibular ramus and the dentition is well visible. As is usual in elasmosaurs the skull is relatively long and low. The external naris is visible as a broad oval opening that is situated close to the orbita. The latter is prominently developed and, due to the lack of the ventral border by the maxilla, nearly circular in shape. About in the middle of the orbita, an elongate NEW ELASMOSAUR FROM QUEENSLAND 427 FIG 2. Skull of Tuarangisaurus australis sp. nov., QMF1 1 050 in dorsal (A) and ventral (B) views. Scale = 10cm. element is visible. Persson (1982) misidentified this bone as the frontals, therefore concluding that the dorsal and the ventral half of the left orbita were the left and the right orbitae. Thulborn & Turner (1993) showed that this structure could be a part of either the pterygoid or the vomer. In the posterior part of the skull, the quadrate and the quadrate condyle are well visible. The right mandibular ramus is almost complete and still articulates with the quadrate. A well preserved hyoid is situated ventral ly, in the posterior part of the specimen (Fig. 4). Posteriorly still in articulation with the skull is the atlas-axis complex (Fig. 5A). The fact that the atlas and axis are co-ossified indicates that the specimen represents an adult individual. As the atlas is still in connection with the condylus occipitalis and also partly covered by the squamosal, its shape could not be described. The axis on the other hand, as well as the ventral section of the atlas-axis complex is well visible. As Thulborn & Turner (1993) showed, there are 3 bite marks of a larger plesiosaur visible on the skull (Fig. 3). A very prominent bite mark is situated in the anterior section of the lower jaw, somewhat behind the symphyseal part on the right ramus. A second relatively large bite mark is situated further posteriorly on the lateral surface of the right mandibular ramus, underneath the coronoid process. The third, relatively small, bite mark is on the left side in the lower part of the squamosal. These bite marks could originate from a large pliosaur, such as Kronosaurus queenslandicus , that has been found in the same area. Premaxilla . The premaxilla is visible on the left side of the skull and is here almost completely preserved. Only the anteriormost part is corroded and therefore missing. In anterodorsal aspect both premaxillae are well ossified and are separated by a barely visible, straight suture. Ventrally the well-preserved suture to the maxilla runs posteromedially at an angle of about 60° to 428 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Skull of Tuarangisaurus australis sp. nov. QMF 11050 in dorsal (A) and ventral (B) views with explanations. the transverse plane to the anterolateral edge of the external naris. With a broad dorsal section the premaxilla forms the slightly concave medial edge of the external naris. Behind the external naris, it touches the dorsal margin of the maxilla with a short suture. The posterior termination of the premaxilla, probably to the frontals, is not preserved. The tooth-bearing part is somewhat more robust and bears three teeth. Although the anteriormost section of the premaxillae is missing, it can be calculated that two more teeth were present, bringing the total to five. The first preserved tooth (pint 3) is relatively short and completely exposed. The next one (pmt 4) is massively developed, but incomplete, as most of the labial and the total apecal part are missing. The third preserved tooth (pmt 5) is badly crushed, but was also massively build. It is situated directly at the premaxillary-maxillary suture and is approximately twice as large as the first preserved maxillary tooth. Between each tooth there is a gap of about l cm for the interlocking of the dentary teeth. Maxilla. The left maxilla is almost completely preserved. Only the part that turns posteriorly to frame the orbita anteroventrally and contact the anterior section of the jugal lacks its dorsal margin. On the right side of the skull only the tooth-bearing part of the maxilla is visible. Generally the maxilla is a massive element. It contacts the premaxilla anteroventrally and here forms the lateral and posterior edge of the external naris. With the projection of the dorsal-most part, it contacts a structure that might represent the prefrontal. As mentioned before the posterior section of the maxilla is incompletely preserved. At the posteriormost part that is formed as a long process, an irregular suture to the jugal is visible. It ventrally follows the jugal to nearly its posterior end and thereby gradually NEW ELASMOSAUR FROM QUEENSLAND 429 thins out dorsoventrally, terminating immediately behind the level of the last dentary tooth. The tooth-bearing part of the maxilla is somewhat more robust and nearly all teeth are still in their alveoli. The second, third and fourth tooth are very prominent, in which the third maxillary tooth appears to be the largest in the maxilla. The teeth become gradually smaller posteriorly. In the most posterior section two extremely small replacement teeth are situated. In the right maxilla 1 2 teeth are visible, in the left one 14 (in contrast to Persson (1982: 649) who counted 10 in the right and 1 1 in the left maxilla), so that probably a total number of 15 or 16 teeth per side was present. Postorbital. The postorbital section of the specimen is heavily crushed. The only more undistorted fragment is a short triangular shaped piece that is situated in about the middle of the posterior part of the enlarged orbita. It contacts the jugal with a well visible, smooth, ventrally convex suture. This fragment appears to represent only the anterior section of the ventral portion of the postorbital so that the bone was probably more expanded posteriorly. Some other fragments, which arc situated underneath the named element and medially of the orbita, could also belong to the postorbital. Frontal A broad, flat element that is situated above the named postorbital fragment is here interpreted as part of the frontals. Its visible dorsal surface is smooth and shows a shallow vault in about its midsection. As the underlying parietal fragment bears a sagittal crest, this vaulting can be the result of post-burial compression. Prefrontal A low, elongate, slightly curved element that is situated laterally, behind the premaxilla, touching the anterodorsal process of the maxilla, is here interpreted as the prefrontal. It forms most of the dorsal edge of the orbita. Parietal Although there are some fragments present in the area of the skull roof where the temporal fenestrae were placed, only one can clearly be determined as part of the parietal. It shows a well developed, sharp, but low crest. On both sides of the crest the dorsal margin is shallowly inclined. This fragment probably formed part of the most anteromedial edges of the temporal fenestrae. Jugal The jugal is well exposed on the left side and, in lateral view, has a rectangular outline. Its original shape cannot be reconstructed with certain, as part of the anterior section that probably contacted the midsection of maxilla and took part of the ventral framework of the orbita is not preserved. Posteriorly the jugal contacts the squamosal with a well visible, somewhat anteriorly oriented, serrated suture. Ventrally the irregular posteroventrally running suture between the jugal and maxilla is visible. Dorsally, in the preserved anterior section, the jugal contacts the postorbital with a slightly concave suture. Squamosal The squamosal is almost completely preserved on the left side, in which only its dorsal margin is missing. Anteriorly it contacts the jugal with a high, serrated suture. In lateral view the ventral surface of the squamosal is shallowly posteroventrally curved. In ventral view it can be seen that the bone was here relatively thin. Posteriorly the squamosal covers the quadrate laterally and is then, with its posterior margin, slightly up- and forwardly arched. The suture between the squamosal and the quadrate is not visible, indicating that this area was well ossified. A triangular shaped fragment that is situated anterior of t he ?exoccipital could be a part of the posterodorsal arch of the squamosal to the parietal. Quadrate. The quadrate is well visible on the left side and is covered by the squamosal laterally. Due to the compression the quadrate has moved medially, so that its condyle has an anteromedial position. The latter does not seem to be depressed; it has a rectangular shape and is expanded medially with the ventral surface well rounded anteroposteriorly. At the lateral side of the ventral margin a shallow depression is visible. The posterior side of the condyle is somewhat ventrally oriented. Interestingly there is a concave excavation on the internal side of the condyle, so that at its dorsal and ventral side a short tip is formed. This concavity seems not to be produced postmortem. It could have serv ed for the articulation of the quadrate process of the pterygoid. A dividing of the articulation surface in two clearly distinguishable condyles (condylus lateralis quadrati and condylus medialis quadrati, after Maisch, 1998) as it can be find e.g. in Muraenosaurus , is not present. At the lateral side the condyle emerges from the prominent shaft of the quadrate in which it is well curved postero- medial ly. As mentioned before a clear suture between the quadrate and the squamosal cannot be seen. Eventually a line that starts at the lateroventral side of the squamosal-quadrate-complex and is upwardly directed could represent this suture, but similar lines are visible in other parts of the skull too and may be quoted as breaking lines rather than sutures. Dorsally, the thin 430 MEMOIRS OF THE QUEENSLAND MUSEUM quadrate process of the pterygoid, is attach to the condyle. Pterygoids. Situated in the middle of the orbita there are 2 elongated, broad fragments, which were interpreted as frontals by Persson (1982). Thulborn & Turner (1993) pointed out that they might represent midline bones of the palate and could be parts of either the vomer or the pterygoids. I agree with this determination. Unfortunately, because of their poor and fragmentary preservation it is impossible to give a clearer statement. Only the position and the visible shape as broadly rounded makes it probable that these elements are parts of the pterygoids. Braincase Elements. Situated at the level of the upper part of the squamosal, inward of the atlas-axis-complex, there is an isolated element. It consists of a main body that has a nearly quadratic outline and two short, broad processes, which are situated at the end of the main body. The later is relatively flat and has a slightly expanded end, opposite of where the processes are situated. The surface of this portion is shown as deeply convex, but it seems that parts of the margin are broken. The short ?lateral edge that is exposed, bears a crest. The named processes, which are probably incompletely preserved, are shown as short and broad. The upper one of these has a nearly horizontal position in the specimen and is broad oval in cross-section. The lower process is more vertically directed. Its basal margin seems to be tighter than the well-expanded ventral margin. It was not possible to make a secure determation of this element, but it shows similarities to the exoccipital-opisthotic (these bones are often fused in plesiosaurs, as seen e.g. in Muraenosaurus , Maisch, 1998, fig. 9, here also called otoccipital). Dentary. The lateral side of the mandible is completely exposed at the left side of the specimen. The symphysis is straight and relatively short (expanded to the level of the fourth dentary tooth). A suture separating the two rami is not visible, which again shows that the skull was well ossified. The anteriomiost part of the dentary is oblique posteroventrally by an angle of about 50° to the vertical plane. Laterally, behind the symphyseal portion the margin is smoothly depressed and bears some small foramina of different sizes. The dorsal side of the dentary is more robust. Towards its midsection the mandibular ramus becomes slightly lower dorsoventrally. Posteriorly, in the coronoid section, the ramus is again high and then inclined FIG. 4. Hyoid of Tuarangisaurus australis sp. nov. QMF1 1050 in ventral view, measuring 10cm. posteroventrally towards the glenoid fossa. The ventral margin of the ramus is well rounded and thickened anteriorly. The suture from the dentary to the surangular-articular-complex and the angular is only barely visible, due to the presence of a bite mark (Fig. 3). The dorsal side of the surangular-articular complex bears a sharp edge and slopes down by an angle of about 40° to the horizontal plane. Altogether the mandible bears 36 alveoli (18 per ramus), in which mostly parts of the teeth are still present. Only the anterior- most teeth are prominent, with mandibular teeth 2-5 developed as large fangs. A ngular-Surangular-A rticular. The angular is only partially visible. Internally, at the left ramus it is particularly detached, but can only be described as very thin, flat element as no characteristics are present. Better preserved is the posterior part, which is well visible at the right mandibular ramus. The retroarticular process is broken off and somewhat displaced. In lateral view it is short and almost quadratic in shape. It shows a slight transversal concavity at its posterior edge. The suture from the angular to the articular-surangular (according to Brown, 1981, these bones appear to be fused in all plesiosaurs, so that this suture should here only be called angular-articular suture) is well visible in the ventral section of the posterior part of the right ramus. The suture first runs straight posteriorly and is then, at the level of the anterior section of the glenoid fossa upwardly and slightly hindwardly curved, from where it runs straight posteriorly. Its termination is unclear as it is last visible directly underneath the glenoid fossa (the section behind is thus called the angular- articular-complex). The latter is still connected with the quadrate so that its shape cannot be described. Hyoid. Both hyoids are preserved (Fig. 4) in which only the right one is well exposed, while the left one is underlying the right, so that only its posterior margin is visible. They are rod-shaped, NEW ELASMOSAUR FROM QUEENSLAND 431 FIG 5. Vertebrae of Tuarangisaurus australis sp. nov. QMF1 1050 in lateral views. A, atlas-axis complex; B, anterior cervical vertebra; C, posterior cervical vertebrae; scale bar units = 1cm. very long, slightly curved and slender. The anterior and posterior end facts are oval in cross-section. The anterior end facet is only weakly concave while the posterior one shows a somewhat stronger concavity. Both facets are slightly broader than the shaft. The later is flattened and has a sharp medial edge, while the lateral edge is well rounded. The anterior half of the right hyoid is somewhat depressed dorsoventrally and therefore slightly broadened lateromedially. Atlas-Axis Complex . The atlas-axis complex is still connected with the skull (Fig. 5A), therefore only the axis is well exposed and can be described. In general the complex has a nearly quadratic shape and is relatively high. In the center of the ventral surface a prominent, sharp sagittal keel is formed. This keel is present upon the entire length of the centrum and anteriorly becomes larger dorsoventrally. Posteriorly the keel is fused with the edge of the articular facet. On both sides of the keel the ventral margin is slightly longitudinally excavated. The rib facets are situated on the lateroventral side of the complex. On the left side also part of the cervical rib is preserved that has a horizontally oval cross-section and is lateromedially relatively thin. The lateral side of the atlas-axis complex is nearly flat and only in the ventral half posteriorly slightly depressed. Dorsally parts of the thin neural arches are preserved, the neural canal was relatively broad, ventrally just weakly concave and occupied about 50% of the breadth of the axis centrum. The articular facet of the latter is about circular and slightly concave. Its dorsal side is almost straight; the lateral sides are weakly rounded. At the ventral side the attachment of the named keel is visible medially. Dorsally, anterior of the atlas-axis complex, there is a thin, well-rounded crest that is mostly covered by sediment. The posterior margin of this crest shows a breaking surface. It most probably is this the neural spine that has been pressed forwards during the compaction of the skull. Cervical Vertebrae. The centrum of the preserved isolated anterior cervical vertebra (QMF 1 22 1 6- 19) has a broad oval outline and is about as wide as long, but clearly lower (5B). Its articular facets are weakly concave, with well-rounded edges. The lateral margin is depressed at one side. In about its mid-section a sharp, medially situated lateral longitudinal crest is formed. Dorsally and ventrally of the crest the margin is longitudinally excavated. Due to the compression, the neural canal is not visible. The rib facets are situated laterally, about in the middle of the ventral surface. They are elongated and ovoid in outline. The ventral surface of the centrum is somewhat depressed. A well developed thin sagittal crest is visible that separates two relatively small nutritive foramina, which are situated in the anterior half of the centrum. On both sides of the crest, the margin is depressed anteroposteriorly. The posterior cervicals (QMF 122 17) are accurately preserved (Fig. 5C). The centra are slightly broader than long and longer than high, so that they have a nearly quadratic outline. Dorsally and ventrally, the centra are weakly excavated medially. The ventral margin of the centra is concavely depressed. Medially, some- what anterior to the midline, two relatively large nutritive foramina are present. A small, flat crest separates them. The rib facets are very prominent, elongated oval in outline and are situated ventrally in the middle of the lateral side. Anterior and posterior to the rib facets the lateral margin is somewhat concave. Each rib facet is oriented l atero ventrally at an angle of about 45° to the vertical plane. In the second vertebra, remains of the cervical ribs are preserved. Laterally, above the parapophyses a weak 432 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. Reconstruction of the skull of Tuarangisaurus australis sp. nov. in lateral view; A, outline drawing (stippled areas are incomplete or not preserved); B, restoration. Scale = 10cm. longitudinal crest is visible. This crest probably represents the lateral longitudinal crest that is well developed in the anterior cervical vertebra. Above the crest the lateral margin is somewhat depressed. The articular facets of the centra are weakly concave, with rounded edges. A dorsal suture between the centrum and the neural arches is not visible. The neural arches are relatively thin and border the broadly triangular neural canal. The margin of the zygapophyses is discernable. In lateral view the pre- zygapophyses rise above the level of the centrum to about half of their length, in which they are dorsally oriented at an angle of about 40° to the horizontal. The postzygapo- physes are higher than the level of the prezygapophyses. They do not rise over the level of the centrum and are dorsally oriented at an angle of about 60° to the horizontal plane. The neural process is present only by its basal margin. It is relatively thin, but prominently broadened anteroposteriorly. COMPARISON OF TUARANGISAURUS KEYESI AND TUARANGISAURUS AUSTRALIS The type material of T. keyesi Wiffen & Moisley, 1986, comprises a skull (NZGSCD425) (Fig. 7) and the 9 anterior-most cervical vertebrae including the atlas-axis complex (NZGSCD426). The skull is fairly complete and, except for the posterior sections of the mandibulae and the occiptal region, well preserved. T. keyesi shares a number of characters with T. australis. In both the premaxillae form a beak with a narrow dorsal keel along their midline. The posterior sections of the premaxillae form the concave medial edge of the external nares. Each premaxilla bears five fang-like teeth; the last of them is very prominent and situated directly at the premaxillary- maxillary suture. The later runs to the antero- lateral margin of the external naris with an angle of about 60° to the horizontal. The maxilla forms the lateral and posterior edge of the external nares. Posteriorly a thin maxillary process is formed that follows the jugal to its posterior section, thereby becoming gradually smaller dorsoventrally. The maxillary tooth count is 15-716 per side. The anterior section of the jugal contacts the maxilla and the postorbital with a large serrated suture. Posteriorly the jugal contacts the squamosal via a distinct serrated suture. The dorsal jugal-postorbital suture is somewhat concave. The ventral surface of the squamosal is shallowly posteroventrally curved. In lateral view the squamosal is only slightly up- and forwardly arched. The anteriormost part of the dentary is posteroventrally oriented by an angle of about 50° to the transverse plane. The anterior portions of the dentary form a beak. The posterior dentary teeth are very prominent and clearly larger than the posterior maxillary teeth. Towards its midsection the dentary becomes slightly narrower dorsoventrally. Posteriorly in the coronoid section, the jaw becomes higher again. The ventral margin of the mandibular ramus is rounded and anteriorly relatively thick. The axis is nearly quadratic in outline, having a NEW ELASMOSAUR FROM QUEENSLAND 433 FIG 7. Skull of Tuarangisaurus keyesi NZGS CD425 in lateral and ventral view with schematic outline drawings (stippled areas are incomplete or not preserved). Scale = 10cm. prominent sagittal keel in the centre of the ventral surface. The cervical rib is lateroventrally directed by an angle of about 45°. The neural canal is relatively broad and occupies about 50% of the breadth of the axis centrum. T. australis can be distinguished from T. keyesi by a number of osteological features, its providence and age. All materials of T. keyesi known so far are from the Mangahouanga Stream site at Hawke’s Bay (North Island, New Zealand). The fossil bearing Maungataniwha Member of the Tahora Formation is interpreted as being approximately mid-Campanian in age (Isaac et al., 1991 ). The Queensland specimen is from the Albian Toolebuc Formation. The osteological differences between T. australis and T. keyesi are the position of the external nares, which in T. keyesi are somewhat more anteriorly situated, about at the level of the third and fourth maxillary teeth. The third premaxillary tooth that in T. keyesi is very large and about as prominent as the 5th premaxillary tooth, while it is rather short in T. australis. The last premaxillary tooth that (calculated from the size of the alveoli), in T. keyesi did not differ much in size from the 1st maxillary tooth, while in T. australis the last premaxillary tooth is about twice as large as the first maxillary tooth. The posterior process of the maxilla, which in T. keyesi terminates at the level of the penultimate dentary tooth, while it terminates directly behind the last dentary tooth in T. australis. The anterior parietal section that in T. keyesi is formed as a thin bridge, while it is rather sharp in T. australis. The jugal, that in T. keyesi is rather trapezoid in outline, while it appears to be more rectangular in T. australis. The jugal-squamosal suture that in T. keyesi is clearly more anteriorly situated than in T. australis. The quadrate condyle, which in T. australis is shorter and more robust. The mandibular symphysis that in T. keyesi only expands to the level of dentary tooth 3, while it expands to the level of dentary tooth 4 in T. australis. The dentary that in T. keyesi bears 21 teeth per ramus and only 18 in T. australis. The dentary teeth, which in T. keyesi are almost equal in size throughout the dentition, while in T. australis the anterior 5 teeth are larger. The retroarticular process that in T. keyesi is about twice as long as broad, while it is about as long as broad in T. australis. The posterior articular facet of the axis, which in T. keyesi is broad oval in shape and deeply concave, while it is more circular and less concave is T. australis. 434 MEMOIRS OF THE QUEENSLAND MUSEUM Based on the listed osteological similarities T. australis is referred to Tuarangisaurus , and because of the identified differences from T. keyesi , it is treated as a separate species. CLADISTIC ANALYSIS The ingroup includes all valid members of the Elasmosauridae of which sufficient cranial material is known. Thus Elasmosaurus , Aphrosaurus , Fresnosaurus and Moraeriosaurus are excluded from the analysis. As outgroup member Muraenosaurus was chosen, which according to O’Keefe (2002) belongs to the Cryptoclididae, although it has long been included in the Elasmosauridae. Recent analyses of elasmo- saurid taxa have been undertaken by Carpenter (1999), Bardet et al. (1999) and Gasparini et al. (2003), and the characters are partly based upon their results. REFERRED TAXA. OUTGROUP: Muraetio- saunis Seeley, 1 874. INGROUP: Occitanosaurus Bardet, Godefroit & Sciau, 1 999, Brancasaurus Wegner, 1914, Libonectes , Carpenter, 1997, Thalassomedon Welles, 1943, Tuarangisaurus Wiffen & Moisley, 1986, Callawayasaurus Carpenter, 1999, Styxosaurus Welles, 1943, Hydralmosaurus Welles, 1943, Aristonectes Cabrera, 1941, Hydrotherosaurus Welles, 1943 and Terminonatator Sato, 2003. CHARACTER DESCRIPTIONS. 1. Orbita - temporal fenestra relative size. 0. equal in size. /. orbita smaller than temporal fenestra, (modified from Gasparini et al., 2003). Outgroup. In Muraenosaurus the orbitae and temporal fenestrae are about equal in size (Andrews, 1910, fig. 46). Ingroup. In most elasmosaurs the temporal ' fenestrae are clearly larger than the orbitae, as present in Occitanosaums (Bardet et al., 1999, fig. 3), Libonectes (Carpenter, 1997, fig. 2), Thalassomedon (Carpenter, 1999, fig. 12), Callawayasaums (Carpenter, 1999, tig. 13), Styxosaurus (Welles & Bump, 1949, fig. 3), Hydralmosaurus (Carpenter, 1999, fig. 7), H\ xlmthemsai uvs (UCMP 33912, Sachs, pers. obs. 2001 ) and Tuarangisau/us. In Brancasaums both are about equal in size (Wegner, 1914, fig. 1 ). In Aristonectes (Gasparini et al., 2003) and Terminonatator (Sato, 2003) the conditions are unknown. 2. Premaxillary tooth count. 0. five. L between 1 and 6. 2. > 6. (modified from Gasparini et al., 2003). Outgroup. Primitively five premaxillary teeth are present in Muraenosaunis (Andrews. 1910). Ingroup. The number of premaxillary teeth is highest in Aristonectes with 10-13 per side (Gasparini et al., 2003), which also is the most increased number of premaxillary teeth in all sauropterygians. Tenninonatator has nine premaxillary teeth per side (Sato, 2003). Eight premaxillary teeth have been estimated for Elasmosaurus (Cope, 1 869, 1 875), but only 6 alveoli arc visible (ANSP 10081, Sachs, pers. obs. 2001). Five premax illary teeth are present in Libonectes (Welles, 1949), Brancasaurus (Wegner, 1914), Hydralmosaurus (Carpenter, 1999), Callawayasaums (Welles, 1962), Styxosaurus (Welles & Bump, 1949), Occitanosaurus (Bardet et al., 1999), Hydrotherosaurus (UCMP 33912, Sachs, pers. obs. 2001, in addition to 4, counted by Welles, 1943) and Tuarangisaurus. The number of premaxillary teeth is lowest in Thalassomedon with only 4 teeth per side (Welles, 1943). 3. Premaxilla-parietal contact. 0. absent. /. present, (modified from Carpenter, 1999). Outgroup. In Muraenosaurus a premaxillary-parietal contact is not established as the frontals intermediate between the 2 elements (Andrews, 1910). Ingroup. A premaxilla-parietal contact is established in Callawayasaurus, Hydralmo- saurus, Libonectes , Styxosaurus , Thalassamedon , Tuarangisaurus keyesi (after Carpenter. 1 999: mb. 2) and Terminonatator (Sato, 2003). The prcmaxillae and parietals are separated by the frontals in Occitanosaurus (Bardet et al., 1999, fig. 3), Brancasaurus (Wegner, 1914, fig. 1 ) and Hydrotherosaurus (Sachs, pers. obs 2001 ). This section of the skull is not well preserved in QMF 1 1 050 and Aristonectes (Gasparini ct al., 2003). 4. Maxilla-jugal contact. 0. maxilla does not follow the jugal posteriorly. /. a posterior process of die maxilla follows the jugal to its posterior section. 2. maxilla terminates posterior of the jugal. Outgroup. In Muraenosaurus only the anteriormost sections of the jugals are in touch with the maxillae (Andrews, 1910, fig. 46). Ingroup. In most elasmosaurs the maxilla follows the jugal to nearly its posterior end, as it is present in Libonectes (Welles, 1949, plate 1), Styxosaurus , Hydralmosaurus (Welles, 1952, fig. 5+6) and Tuarangisaurus, In Brancasaums (Wegner, 1914, fig. 1) and Thalassomedon (Carpenter, 1999, fig. 13) the maxilla is somewhat larger than die jugal and runs further posteriorly. This appears also to be the case in Terminonatator (Sato, 2003, fig. 5a), although the jugal is only fragmentarily preserved. For Callawayasaurus Welles (1962: 18) wrote ‘the maxillary probably ends posteriorly below the middle of the jugal arch in a downcurved slender point'. Only a short contact in the anterior section of the jugal is present in Occitinosaurus (Bardet et al, 1999, fig. 3). The jugal is not known in Aristonectes (Gasparini et al., 2003) and the condition is unclear in Hydrotherosaurus as not clear suture is visible in that part of the skull (UCMP33912, Sachs, pers. obs. 2001 ). 5. Maxillary toodi count. 0. 1 7 or more. /. 1 4- 1 6. 2, 13 or fewer. Outgroup. According to Brown (1981) Muraenosaunis bears 16 teeth per maxilla. Ingroup, rhe number of maxillary teeth is 14-16 in Libonestes (14 after Carpenter, 1999) and Tuarangisaurus (15-16). The number is low er than 14 in Hydralmosaums (13 after Welles, 1952), Terminonatator (13, after Sato, 2003), Occitinosaurus (12 after Bardet et al., 1999), Styxosaurus ( 1 1 after Welles & Bump, 1949) and Callawayasaurus (9 after Welles, 1962). 13 alveols are visible in NEW ELASMOSAUR FROM QUEENSLAND 435 Hydmthemsaunis, in which the posterior ones are not clear (UCMP 33912, Sachs pers. obs. 2001). According to Welles (1943: 158) Thalassomedon only bears 7 teeth per maxilla, while in the drawing of Carpenter ( 1 999. fig. 12) 1 3 teeth are visible. The largest number of maxillary teeth among all sauropterygians is present in Aristonectes with about 51-53 teeth per side (Gasparini et al., 2003). The number is unclear in Brancasaunis , as the maxillae are incompletely preserved (Wegner, 1914). 6. Maxillary termination. 0. maxilla terminates posterior to the level of the hist dentary tooth. /. maxilla terminates close to the level of the last dentary tooth. Outgroup. In Muraenosaurus the maxilla terminates farther behind the level of the last dentary tooth about at the level of the posterior margin of the orbita (Andrews, 1910, fig. 46). Ingroup. The maxilla terminates close to the level of the last dentary tooth in Callawaymaunis (Welles, 1962, fig. 3), Styxosaurus (Welles, 1952, fig. 5), Tenninonatator (Sato, 2003. fig. 5), Tuarangisaunts and probably also in Hydralmosaunis (Welles, 1952, fig. 6) and Aristonectes (Gasparini et al. 2003, fig. 1). The maxilla is further posteriorly expanded in Libonectes (Carpenter, 1997, fig. 5), Thalassomedon (Carpenter, 1999, fig. 12) and Brancasaunis (Wegner, 1914, fig. 1). The situation is unknown in Oecitanosaurus as the lower jaw is missing (Bardet et. al., 1999) and in Hydmthemsaunis (UCMP 339 1 2, Sachs, pers. obs. 200 1 ) as this part of the skull is not well preserved 7. Jugal-squamosal contact. 0. by a short suture or no contact is established. /. by a long suture. Outgroup. In Muraenosaurus only a short suture in the ventral half of the squamosal is developed (Andrews, 1910. fig. 46). Ingroup. A long squamosal-jugal contact is present in most elasmosaurs. It can be found in Styxosaurus , Hydralmosaunis (Welles, 1952, fig. 5+6), Thalassomedon (Carpenter, 1999, fig. 12), Libonectes (Welles, 1949, plate 1) and Tuarangisaunts. In Callawayusaunis this section is somewhat distorted, but a long suture between the squamosal and the jugal is visible (Welles, 1962, fig. 3). A short suture is developed in Occitinosaunis (Bardet et al., 1999, fig. 3) and Brancasaunis (Wegner, 1914. fig. 1) as here the anterior margin of the squamosal also articulates with the postorbital. As mentioned before the jugal is only fragmentarily preserved in Terminonatator (Sato, 2003) and is missing in Aristonectes (Gasparini et al., 2003), so that no statements are possible for these taxa. In Hydmthemsaunis the posterior suture to the squamosal is not preserved (UCMP33912, Sachs, pers. obs. 2001). 8. Jugal-postorbital contact. 0. suture long relative to the length of the jugal. /. suture short relative to the length of the jugal, (modified from Bardet et al., 1999). Outgroup. A long postorbital .suture is present in Muraenosaurus , reaching over most of the dorsal edge of the jugal (Andrews, 1910, fig. 46). Ingroup. The jugal-postorbital suture is comparatively long in Styxosaurus (Welles, 1 952, fig. 5), Hydralmosaunis (Welles, 1952, fig. 6). Callawayasaums (Welles, 1962, fig. 3), Occitinosaurus (Bardet et al., 1999, fig. 3), and Tuarangisaurus. It is short in Libonectes (Welles, 1949, plate 1), Thalassomedon (Carpenter. 1999, fig. 12) and Brancasaunis (Wegner, 1914, fig. 1 ), while this section of the skull is not preserved in Aristonectes (Gasparini et al., 2003) and Tenninonatator (Sato, 2003) and the condition is unclear in Hydmthemsaunis (UCMP 339 1 2, Sachs, pers. obs. 200 1 ). 9. Jugal-orbita contact. 0. Jugal does not take part in the margin of the orbita. /. Jugal takes part in the margin of the orbita. Outgroup. In Muraenosaunts the jugal forms the posterior half of the lateral margin of the orbita (Andrews, 1910, fig. 46b). Ingroup. The jugal is participated in forming the orbita in all referred elasmosaur taxa with the exception of Oecitanosaurus (Bardet et al., 1999, fig. 1) Here the jugal is dorsally covered by the postorbital. In Tenninonatator only a fragment of the right jugal is preserved, but according to its position it appeared to have been participated in forming the orbita too (Sato, 2003, fig. 5). The jugal is unknown in Aristonectes (Gasparini et al., 2003 ). 10. Jugal-temporal fenestra contact. 0. jugal is excluded from the dorsal margin of the temporal fenestra. /. jugal takes part in the dorsal margin of the temporal fenestra, (sensu Carpenter, 1999). Outgroup. In Muraenosaunts the jugal is not participated in the margin of the temporal fenestra, as it is dorsally covered by the postorbital, which forms the antcroventral margin of the fenestra (Andrews, 1910). Ingroup. Tire jugal takes part on the dorsal margin the temporal fenestra in Thalassamedon (Carpenter, 1999, fig. 12), Callaway asaurus (Welles, 1962, fig. 3), Hydralmosaunis (Welles, 1952. fig. 6), Styxosaurus (Welles, 1952, fig. 5), Libonectes (Carpenter, 1997, fig. 5) and Tuarangisaurus. The jugal is dorsally covered by the postorbital and therefore not participated in Occitinosaurus (Bardet et al., 1999, fig. 3) and Brancasaunis (Wegner, 1914, fig. 1 ). The condition is unknown in Terminonatator (Sato, 2003) and Aristonectes (Gasparini, et al. 2003) and unclear in Hydmthemsaunis (UCMP 33912, Sachs, pers. obs. 2001). ' 11. Postfrontal. 0. present. /. absent, (sensu Bardet et al., 1999). Outgroup. The postfrontals arc present in Muraenosaurus (Andrews, 1910). Ingroup. The postfrontal is a primitive element and is present in Oecitanosaurus (Bardet et al., 1999, fig. 3) and Brancasaunis (Wegner, 1 9 1 4, fig. 1 ) and absent in all other referred elasmosaur taxa. The according section of the skull is unknown in Aristonectes (Gasparini et al., 2003). 12. Squamosal-postorbital contact. 0. present /. absent. Outgroup. In Muraenosaunts . a clear contact between the postorbital and the squamosal is present (Andrews, 1910, fig. 46). Ingroup. The squamosal does not touch the postorbital in Styxosaurus (Welles, 1952, fig. 5), Thalassomedon (Carpenter, 1999, fig. 12), Callaway a- saurus (Welles, 1962, fig. 3) and Tuarangisaurus. A contact is present in Oecitanosaurus (Bardet et al., 1999, fig. 3) and Brancasaunis (Wegner, 1914, fig. 1). In Hy dmthemsaunis a zigzagged suture seems to be present, but is not well visible (UCMP 33912, Sachs, pers. obs. 2001). A contact has also been suggested for Hydralmosaunis (Welles, 1952, fig. 6), but this reconstruction appears to be doubtful. The section is not 436 MEMOIRS OF THE QUEENSLAND MUSEUM well preserved in Libonectes (Carpenter, 1997, fig. 2) and is missing in Aristonectes (Gasparini et al., 2003) and Terminonatator (Sato, 2003). 13. Dentary teeth per ramus. 0. 21 or more. /. 18-20. 2. 17 or fewer. Outgroup. In Muraenosaurus the tooth count is 22 per dentary ramus (Brown, 1 98 1 ). Ingroup. The number of dentary teeth is 18-20 in Libonectes (17/18, after Carpenter, 1997), Callawayasaurus (19, after Welles, 1962), Terminonatator (17/18, after Sato, 2003) and QMF 1 1 50 ( 1 8). According to Welles ( 1 943) the mandible of Thalassomedon only bears 16 teeth per ramus. However, following the photo of UNSM 50132 in Carpenter (1999, fig. 12) at least 19 teeth per ramus are present. The number is higher in Tuarangisawvs kewsi (21, NZGS CD425, Sachs pers. obs. 2001) and Brancasaums (21, after Wegner, 1914) and lower in Styxosaurtis (17, after Welles & Bump, 1949). In Hydralmosanrus 17 teeth are present (after Welles, 1952), but Carpenter (1999, p. 156) suggests that the number might be higher. The number is most increased in Aristonectes with about 60-65 teeth per dentary ramus (Gasparini et al., 2003). The condition is unclear in Hydwtherosaurus (UCMP 33912, Sachs, pers. obs. 2001) and unknown in Occitanosaurus (Bardet et al., 1999). 14. Shape of dentary teeth. 0. posterior-most dentary teeth are not markedly more prominent than posterior-most maxillary teeth. /. posterior-most dentary teeth very prominent in relation to the posterior-most maxilla teeth. Outgroup. In Muraenosaunts the posterior dentary teeth are about equal in size with the last maxillary teeth (Andrews, 1910, fig. 46). Ingroup. The posterior dentarv teeth are clearly more prominent than the posterior maxillary teeth in Libonectes (Carpenter, 1997, fig. 5). Styxosaunts (Welles. 1952, fig. 5), Hydralmosaunts (Welles, 1952, fig. 6), Callawayasaurus ( Welles, 1962, fig. 3), Tenninonaiutor (Sato, 2003. fig. 5), Hydrothemsaunis (UCMP339 1 2, Sachs, pers. obs. 2001) and Tuarangisaurus. The posterior dentary teeth are about equal in size with the last maxillary teeth in Thalassomedon (Carpenter, 1999, fig. 12). This section is not well preserved in Brancasaurus , but the teeth seem not to be more prominent than the posterior maxillary teeth (GPIM A36, Sachs, pers. obs. 1999). Rather small posterior dentary teeth have been reconstructed for Aristonectes, but they still appear to be larger than the posterior maxillary teeth (Gasparini et al., 2003). 15. Length of retroarticular process. 0 . long relative to jaw length. /. short relative to jaw length. Outgroup. Muraenosaurus has a long retroarticular process (Andrews, 1910, fig. 46). Ingroup. The retroarticular process is comparatively short in Styxosaunts (Welles, 1952, fig. 5), Hydralmosaunts (Welles, 1952, fig. 6), Callawayasaurus (Welles, 1962, fig. 3), Terminonatator (Sato, 2003, fig. 5), Aristonectes (Gasparini et al., 2003, fig. 1) and Tuarangisaurus (in which the process is somewhat longer in T. keyesi than in QMF 1 1050). The retroarticular process is rather long in Thalassomedon (Carpenter, 1999, fig. 12), Libonectes (Carpenter, 1997, fig. 5), Hydwtherosaurus (Welles, 1943, fig. 4) and Brancasaurus (GPIM A36, Sachs, pers. obs. 1999). 16. Axis shape. 0. centrum short and high. /. centrum long and low. (modified from Carpenter, 1999). Outgroup. The axis is short and high in Muraenosaurus (Andrews, 1910, fig. 49). Ingroup. The axis centrum is short and high in Brancasaurus (Wegner, 1914, fig. 2), Libonectes (Carpenter, 1997, fig. 1), Thalassomedon (Welles, 1943, plate 22), Occitanosaurus (Bardet et al, 1999, plate 2), Aristonectes (Cabrera. 1941, fig. 2). Hydmthemsaunts (Welles, 1943), Callawayasaurus (UCMP38349, Sachs pers. obs., 2001 ) and 7 uarangisaurus. The centrum is long and low in Styxosaurus (Sachs, 2004), Hydralmosaurus (Cope, 1 877) and Elasmosaurus (Cope, 1 869, plate 2). The axis vertebra is unknown Terminonatator (Sato, 2003). 1 7. Shape of the cervical centra. 0. anterior cervical centra shorter than high or the length and height is about equal. /. anterior cervical centra longer than high. Outgroup. In Muraenosaurus the length and height are about equal (Andrews, 1910). Ingroup. The anterior cervical centra are longer than high in Styxosaurus (Sachs, 2004), Hydralmosaurus (Cope, 1 877), Libonectes (Welles, 1 949), Elasmosaurus ( Welles, 1952), Occitanosaurus (Bardet et al., 1999), Callawayasaurus (Welles, 1962), Terminonatator (Sato, 2(X)3), Hydmthewsaurus (Welles, 1 943) and Tuarangisaurus. These measurements are about equal in Thalassomedon (Welles, 1943), Brancasaurus (Wegner, 1914) and Aristonectes (Cabrera, 1 94 1 ). 18. Cervical vertebrae number. 0. fewer than 30. /. 30-40. 2.40-50. 3, 50-60. 4 . 60 or more, (modified from Carpenter, 1999). Outgroup. Muraenosaunts comprises 44 cervicals (Brown, 1981). Ingroup. The number of cervicals is lowest in Brancasaurus (37, Wegner, 1914). Occitanosaurus (43, Bardet et al., 1999), Terminonatator (51. Sato, 2003) and Callawayasaurus (56, Welles. 1962). The number of cervicals is 60 or more in Hydmthemsawvs (60, Welles, 1943), Libonectes (62, Carpenter. 1999), Thalassomedon (62, Welles, 1943), Styxosaunts (62, Carpenter, 1999), Hydralmosaunts (62, Welles, 1999) and Elasmosaurus (72, ANSP 10081. Sachs pers. obs. 2001). The total number is unknown in Aristonectes. but at least 23 cervical vertebrae are present in M LP 40-XI- 1 4-6 (Cabrera, 1941, Gasparini et al., 2003) and Tuarangisaurus. 19. Lateral longitudinal crest on cervical centra. 0. absent. /. present, (sensu Bardet et al.. 1999). Outgroup. A lateral longitudinal crest is present in Muraenosaunts (Andrews, 1910). Ingroup. This character is present in all elasmosaurs except for Brancasaurus (Wegner, 1914), which is a juvenile. EXPLANATION OF THE PHYLOGENETIC ANALYSIS METHODS. The data matrix (Table 1) was analysed using PAUP version 3.1.1 (Swofford & Begle, 1993) with 100 heuristic searches and random addition of sequences. The first goal of the analysis was to show an example for the possible interrelationship of the Elasmosauridae; the second goal was to find the position of Tuarangisaurus within the family. All characters NEW ELASMOSAUR FROM QUEENSLAND 437 were left unordered. With a heuristic search, 3 equally most parsimonious trees were obtained, each with a length of 32 steps, a consistency index (Cl) of 0.719, homoplasy index (HI) of 0.281 and retention index (RI) of 0.763. All trees support the position of Libonecles as sister taxon of Thalassomedon, Terminonatator as sister taxon of Aristonecies and Styxosaurus as sister taxon of Hydrcilmosaurus. In all trees Tuarangi- saurus and Callawayasaurus have a single standing position (Fig. 8). The principal difference of the trees is the position of Callawayasaurus, which is either the sister taxon of group 7 or of group 8. Group 1 includes all elasmosaurs, except for Brancasaurus where character 19, the longitudinal lateral crest in the cervical centra, is not present. According to Brown ( 1 993) the crest is a diagnostic feature for the Elasmosauridae, but is only fully developed in adults. However, as shown in Styxosaurus gtendowerensis (Sachs, 2004), a well developed crest can be present in juveniles too. Thus it remains questionable if Brancasaurus really represents an elasmosaur or would rather belong in the relationship of the Cryptoclididae (supported by two characters only present in Brancasaurus and Muraenosaurus ; 1(0) and 18(2)). Group 2 includes the members of groups 4, 7 and 8 based on characters 6(1) and 15(1). The maxilla is primitively more expanded posteriorly than at the level of the last dentary tooth and the retroarticular process is then usually also rather long. Both characters can therefore be quoted as advanced if present. Group 3 includes elasmosaur taxa showing more plesiomorphic conditions, containing the members of group 5 and 6 as well as Hydrotherosaurus . It is characterised only by character 15(0), an elongate retroarticular process. Group 4 includes Callawayasaurus, Tuarangisaurus and group 7, showing a feature complex of characters 5(1) + 8(0) that is unique for this group. Group 5 comprises the most basal members of the Elasmosauridae, thus showing characters 3(0), 7(0), 10(0) and 11(0) as plesiomorphic conditions that are characteristic for this group. Group 6 that is formed by Thalassomedon and Libonectes is characterised a feature complex of character 6(0) + 18(4) that is unique among the Elasmosauridae. Group 7 shows a number of advanced features, e.g. character 5(2), 1 3(2), 16(1) and 1 8(4) that in this combination is only present in this group. It can be concluded that an elongate but low axis FIG 8. Phylogenetic tree of the Elasmosauridae. A, consensus tree, showing the position of Tuarangisaurus and its interrelationships; B, composition of groups 1-8. centrum is an advanced feature, especially considering the fact that the number of cervical vertebrae is equal with that in Libonectes and Thalassomedon where a more quadratic shaped axis centrum is developed. These characters support the high phylogenetic position of group 7. Considering the fact that an elongate and low axis centrum, together with the most increased number of cervical vertebrae in all sauropterygians is also present in Elasmosaurus platyurus , it can be concluded that this taxon probably belongs to 438 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Data matrix. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 _J 9 | Muraenosaurus 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 2 1 Occitanosaurus 1 0 0 0 2 7 0 0 0 0 0 0 7 7 7 0 1 2 1 Brancasaunis 0 0 0 2 9 0 0 1 1 0 0 0 0 0 0 0 0 1 0 Libonectes 1 0 I 1 1 0 1 t 1 1 1 9 1 1 0 0 1 4 1 Thalassomedon 1 1 1 2 2 0 1 1 1 1 1 1 1 0 0 0 0 4 1 Tuarangisaurus 1 0 1 1 1 1 1 0 1 1 1 1 0+1 1 1 0 1 7 1 1 Callawayasaurus 1 0 1 1 2 1 1 0 1 1 1 1 1 1 1 0 1 3 1 Styxosaurus 1 0 1 1 2 1 1 0 1 1 1 1 2 1 1 1 1 4 1 Hvdralmosaurus 1 0 1 1 2 1 1 0 1 1 1 0 2 1 1 1 1 4 1 Aristonectes 7 2 9 9 0 1 9 9 9 9 9 ? 0 1 1 0 0 7 1 Hydrotherosaurus 1 0 0 ? 2 9 9 9 1 9 1 0 7 1 0 0 1 4 1 Terminonatator 9 2 1 2 2 1 9 9 1 9 1 9 1 1 1 ? 1 3 1 group 7 too. Group 8 is characterised by only one character 2(2), the incensement of premaxillary teeth. DISCUSSION. This analysis indicates that Tuarangisaurus , is more primitive than Styxosaurus and Hydrcilmosaurus from the Western Interior Seaway of North America, Terminonatator from the Maastrichtian of Canada and Aristonectes from the Maastrichtian of Patagonia. Tuarangisaurus shows characters 3(1), 6(1), 12(1), 14(1), 15(1) and 17(1) as advanced features. It is more advanced than group 3 in characters 6(1) and 15(1); more advanced than group 5 in characters 3(1), 6(1), 7(1) and 10-15(1); more advanced than group 6 in characters 6(1) and 15(1) and more advanced than Hydrotherosaurus in characters 3(1), 12(1) and 15(1). Tuarangisaurus is less advanced than Cal la way as a u rus in character 5(2), less advanced than group 7 in characters 5(2), 13(2) and 16(1) and less advanced than group 8 in characters 2(2) and 4(2). Thus Tuarangisaurus belongs in the closer relationship of group 7 and Callawayasaurus , but seems to be less advanced than both. ACKNOWLEDGEMENTS I thank Scott Hocknull (Queensland Museum), who took time to work out the phylogenetic analysis with me, discussed various questions and read and commented upon the manuscript. Alex Cook and Trevor Clifford (Queensland Museum) also read and commented parts of an earlier version of the manuscript. Craig Jones (Lower Hutt) and Raymond Coory (Wellington) allowed me to observe the specimens of Tuarangisaurus keyesi under their care. Axel Hungerbiihler (Tucumari) and Mike Everhart (Hays) sent photocopies of rare literature to me. I wish to thank all these people for their help and support. LITERATURE ANDREWS, C. W. 1 9 1 0. A descriptive catalogue of the marine reptiles of the Oxford Clay. Part 1 . British Museum (Natural History): 1-205. BARDET, N., GODEFROIT, P. & SCIAU, J. 1999. A new elasmosaurid plesiosaur from the Lower Jurassic of southern France. Palaeontology 42(5): 927-952. BROWN, D.S. 1981. The English Upper Jurassic Plesiosauridea (Reptilia) and a review of the phylogcny and classification of the Plesiosauria. Bulletin of the British Museum (Natural History), Geology series 35(4): 253-347. 1993. A taxonomic reappraisal of the families Elasmosauridae and Cryptoclididac (Reptilia: Plesiosauroidea). Revue dc Paleobiologie, Special Volume 7: 9-16. CABRERA, A. 1941. Un plcsiosaurio nuevo del Cretaceo del Chubut. Revista del Museo de La Plata 2: 113-130. CARPENTER, K. 1997. Comparative cranial anatomy of two North American Cretaceous plesiosaurs. Pp. 191-216. In Callaway, J.M. & Nicholls, E.L. (eds) Ancient marine reptiles. (Academic Press: San Diego). 1999. Revision of North American elasmosaurs from the Cretaceous of the Western Interior. Paludicola 2(2): 148-173. COPE, E.D. 1869. Extinct Batrachia, Reptilia and Aves of North America. Transactions of the American Philosophical Society 14: 1-252. 1875. The Vcrtebrata of the Cretaceous formations of the West. F.V. Hayden, Report of the United States Geological Survey of the Territories 2: 1-303. NEW ELASMOSAUR FROM QUEENSLAND 439 TABLE 2. Measurements of the alveols (approximate), skull and postcranial elements. Pmt 1 ?mm Dt 1 1 1mm Length of the skull antcropoteriorly 40cm Pmt 2 ?mm Dt 2 16mm Anteroposterior length of the prcmaxilla along the alveolar-part ca. 7.0cm Pmt 3 ?mm Dt 3 15mm Anteroposterior length of the maxilla ca. 2 1 cm Pmt 4 12mm Dt 4 14mm Height of the maxilla at the anterior edge of the orbita ca. 6.0cm Pmt 5 13mm Dt 5 12mm Length of the posterior process of the maxilla ca. 6.0cm Mxt 1 8mm Dt 6 1 1mm Largest distance between the posterior edge of the external naris and the anterior edge of the orbita 1.7cm Mxt 2 12mm Dt 7 9mm Mxt 3 13mm Dt 8 ?mm Anteroposterior diameter of the external naris 2.3cm Mxt 4 13mm Dt 9 10mm Dorsoventral diameter of the external naris 1.2cm Mxt 5 1 1mm Dt 10 10mm Anteroposterior diamteter of the orbita 8.0cm Mxt 6 8mm Dt 11 9mm Length of the jugal anteroposteriorly ca. 8.0cm Mxt 7 8mm Dt 12 13mm Length of the mandibular symphysis 6.2cm Mxt 8 7mm Dt 13 12mm Length quadrate condyle 3.6cm Mxt 9 7mm Dt 14 1 1mm Breadth quadrate condyle 1.4cm Mxt 10 6mm Dt 15 10mm Height quadrate condyle 1.4cm Mxt 11 ?mm Dt 16 9mm Length of the hyoid anteroposteriorly 10.0cm Mxt 12 ?mm Dt 17 9mm Breatdh of the hyoid posteriorly 1.0cm Mxt 13 ?mm Dt 18 7 mm Height of atlas-axis complex 3.4cm Mxt 14 ?mm Breadth of atlas-axis complex 3.3cm Mxt 15 ?mm Length of anterior cervical vertebra 6.4cm Breadth of anterior cervical vertebra 6.9cm Height of anterior cervical vertebra 5.0cm Length of first posterior cervical vertebra 9.8cm Breadth of first posterior cerv ical vertebra 1 0.4cm Height of first posterior cervical vertebra 8.3cm 1877. Report on the geology of the region of the Judith River, Montana, and on the vertebrate fossils obtained on or near the Missouri River. Part 2. Vertebrata from the Niobrara Cretaceous. United States Geological and Geographical Survey of the Territories, Hayden Survey, Bulletin 3: 565-597. CRUICKSHANK, A.R. & LONG J. 1998. Further records of plesiosaurian reptiles of Jurassic and Cretaceaous age from Western Australia. Records of the Western Australian Museum 19: 47-55 CRUICKSHANK, A.R., FORDYCE, R E. & LONG J.A. 1999. Recent developments in Australasian sauropterygian palaeontology (Reptilia: Sauropterygia). Records of the Western Australian Museum 57: 201-205. GASPARINI. Z.. BARDET, N.. MARTIN, J. & FERNANDEZ, M. 2003. The elasmosaurid plesiosaur Aristoneetes cabrera from the latest Cretaceous of South America and Antarctica. Journal of Vertebrate Paleontology 23 (1): 104-115. HECTOR, J. 1874. On the fossil Reptilia of New Zealand. Transactions of the New Zealand Institute 6: 333-358. HILLER, N., MANNERING A. & JONES, C. 1997. Mauisciurus haasti - a new look at an old species of plesiosaur. Abstracts, Conference on Australasian Vertebrate Evolution, Palaeontology and Systematics: 3 1 . ISAAC, M.J., MOORE, P.R. & JOASS, Y.J. 1991. Tahorc Formation: the basal facies of a Late Cretaceous transgressive sequence, northeastern New Zealand. New Zealand Journal of Geology and Geophysics 34: 227-236. KEAR, B. 2003. Cretaceous marine reptiles of Australia: a review of taxonomy and distribution. Cretaceous Research 24: 277-303. MAISCH, M.W. 1998. Notes on the cranial osteology of Muraenosaurus Seeley, 1874 (Sauropterygia, Jurassic), with special reference to the ncurocranium and its implications for sauropterygian phylogcny. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 207(2): 207-253. 440 MEMOIRS OF THE QUEENSLAND MUSEUM MURRAY, P. F. 1987. Plesiosaurs from Albian aged Bathurst Island Formation siltstones near Darwin, Northern Territory, Australia. The Beagle 4(1): 95-102. O’KEEFE. R.F. 2002. The evolution of plesiosaur and pliosaur morphotypes in the Plesiosauria (Reptilia: Sauropterygia). Paleobiology 28(1): 101 - 112 . OWEN, R. 1861. On the remains of a plesiosaurian reptile ( Plesiosaurus australis ) from the Oolitic Formation in the Middle Island of New Zealand. Geologist 4: 444-445. 1870. Notice of some saurian fossils discovered by J. H. Hood, Esq. at Waipara, Middle Island, New Zealand. Geological Magazine 7: 49-53. PERSSON, O. 1960. Lower Cretaceous Plesiosaurians (Reptilia) from Australia. Lunds Univcrsitets Arsskri ft 56(12): 1-23. 1982. Elasmosaurid skull from the Lower Cretaceous of Queensland (Reptilia: Sauropterygia). Memoirs of the Queensland Museum 20(3): 647-655. SACHS, S. 2004. Rcdescription of Woolungasaurus glendowerensis (Plesiosauria: Elasmosauridac) from the Lower Cretaceous of Northeast Queensland. Memoirs of the Queensland Museum 49 (2): 215-233. SATO, T. 2003. Terminonataior ponteixensis , a new elasmosaur (Reptilia: Sauropterygia) from the Upper Cretaceous of Saskatchewan. Journal of Vertebrate Paleontology 23 (1): 89-103. SEELEY, H.G. 1877. On Mauisaurus gardneri (Seeley), an elasmosaurian from the base of Gault at Folkestone. Quarterly Journal of the Geological Society of London 33: 541-546. SENIOR, B.A., MOND, A. & HARRISON, P.L. 1978. Geology of the Eromanga Basin. Bureau of Mineral Resources, Australia, Bulletin 167. SWOFFORD, D.L. & BEGLE, D.P. 1993. PAUP - Phylogenetic Analysis Using Parsimony, Version 3.1.1. Laboratory of Molecular Systematics, Smithsonian Institution, Washington D.C. TEICHERT, C. & MATHESON, R.S. 1944. Upper Cretaceous ichthyosaurian and plesiosaurian remains from Western Australia. Australian Journal of Science 6: 167-170. THULBORN, T. & TURNER, S. 1993. An elasmosaur bitten by a pliosaur. Modern Geology 18: 489-501.’ WEGNER, T. 1914. Brancasaurus hrancai n.g. n.sp., ein Elasmosauride aus dem Wealden Westfalens. Pp. 235-305. In Schoendorf, F. (ed.) Branca Festschrift. (Gcbriider Bomlraegcr, Berlin). WELLES, S.P. 1943. Elasmosaurid plesiosaurs with description of new material from California and Colorado. Memoirs of the University of California, 13(3): 125-254. 1 949. A new elasmosaur from the Eagle Ford Shale of Texas. Fondren Science Series, Southern Methodist University 1: 1-28. 1952. A review of the North American Cretaceous elasmosaurs. University of California Publications in Geological Sciences 29: 47-144. 1 962. A new species of elasmosaur from the Aptian of Colombia and a review of the Cretaceous plesiosaurs. University of California Publications in Geological Sciences 46: 1-96. WELLES, S.P. & BUMPT J. 1949. Alzadasaurus pembertoni, a new elasmosaur from the Upper Cretaceous of South Dakota. Journal of Paleontology 23: 521-535. WELLES, S.P. & GREGG D.R. 1 97 1 . Late Cretaceous marine reptiles of New Zealand. Records of the Canterbury Museum 9(1): 1-111. WIFFEN, J. & MOISLEY, W. 1986. Late Cretaceous reptiles (Families Elasmosauridac, Pliosauridae) from the Mangahouanga Stream, North Island, New Zealand. New Zealand Journal of Geology and Geophysics 29: 205-252. TWO NEW SPECIES OF BRANCHINELLA (ANOSTRACA: THAMNOCEPHALIDAE) AND A REAPPRAISAL OF THE B. NICHOLLSI GROUP BRIAN V. TIMMS Timms, B. V. 2005 01 10: Two new species of Branchinella ( Anostraca: Thamnocephalidae) and a reappraisal of the B. nichollsi group of the Australian arid-zone. Memoirs of the Queensland Museum 50(2): 441-452. Brisbane. ISSN 0079-8835. Branchinella clandestina sp. nov. is described from the Queensland Paroo and B. mcraei sp.nov. from the Pilbara in Western Australia. Both have a simple frontal appendage consisting of a trunk and two branches, but differ from each other and other similar species such as B. ajftnis Linder, B. longirostris Wolf and B. latzi Geddes by many features. These two species increase the number of described Australian Branchinella to 31 species. The raising of B. nichollsi hattahensis Geddes and B. nichollsi buchananensis Geddes, both present in the Paroo catchment, to species status is confirmed with lectotypes designated and species descriptions provided. Similarities and differences between B. nichollsi, B. hattahensis and B. buchananensis are discussed. □ Branchinella , Anostraca, Thamnocephalidae, new species. Brian V. Timms, School of Environmental and Life Sciences, University of Newcastle, Callaghan 2308 ( e-mail : brian.timms@newcastle.edu.au); 24 June 2004. Until a few years ago, Australia had 18 described species of Branchinella. Belk & Brtek (1995) elevated, with no discussion or justification, the 2 subspecies of B. nichollsi Linder to species rank (thus B. buchananensis Geddes 1981 and B. hattahensis Geddes 1 98 1 ) to swell the Australian list to 20 species.Timms (200 1 , 2002) and Timms & Geddes (2003) added 9 species, mainly from remote areas. Collecting from such areas has yielded 2 further species, described here. The uncritical elevation of B. n. buchananensis , and B. n. hattahensis to species status has not been questioned, or supported, on anatomical grounds, but Remigio et al.(2003), using DNA, concluded that B. buchananensis and B. hattahensis were related at the subspecific level (no material of nichollsi was available). No formal decision was made. SYSTEMAT1CS Class CRUSTACEA Bmnnich, Mil Order ANOSTRACA, Sars, 1867 Family THAMNOCEPHALIDAE Packard, 1883 Branchinella Sayce, 1902 Branchinella clandestina sp. nov. (Figs 1,2) ETYMOLOGY. The name arises from this fact that this species lives in a pool that was seen regularly for many years of a long-term study of the Paroo but not sampled until the 1 3th year. In other words it probably lived many times when the pool was full, right under the collector’s gaze, yet went unnoticed — a clandestine existence. MATERIAL. HOLOTYPE: 8 Quensland Museum (hereafter QM) W26951. ALLOTYPE: 9 QMW27006. PARATYPES: QMW27007 five 8 8. All collected from the type locality by the B.V. Timms, 5 December, 2001. OTHER MATERIAL. Four 8 8 raised in December 2002 from dried mud from the type locality, QMW27008. TYPE LOCALITY. Unnamed flood plain pool near Caiwarro Crossinu of the Paroo River. Curraw'inya National Park, 28°4r55”S, 144°46’40”E, SW Queensland. Collected under permit W1TK00786602 issued by the Queensland Parks and Wildlife Service. DESCRIPTION. Male. Length of mature males 10-11 .2mm; holotype 10.8mm. First antennae slightly longer than proximal segment of second antenna (Fig. 1A); apex bevelled and bearing subapically 2 short setae and 2-3 recurved hair-like setae. Second antennal basal segments fused medially for about one-third of their length proximally. These segments with a raised field of minute denticles along much of their length medially. Distal segment of second antenna slightly longer than length of the proximal segment, flattened, blade-like, curved with apices directed anteriolaterally (Fig. 1A). In life (Fig. IB) the second antennae presents with the proximal segment vertical at right angles to the body axis, and the distal segments curved in three planes, so that the concave medial surface faces anteriorly and the convex lateral surface is aligned ventromedial ly. Curvatures on the medial and lateral margins of the distal segment different so that segment is widest in the central part. Apical 442 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. Branchnella clandestina sp. nov. A, lateral view of head of 6: B, dorsal view of 6 second antenna; C, dorsal view of frontal appendage of 6 ; D, rigid portion of base of penes; E, penis: F, dorsal view of head of 9 ; G, lateral view of brood pouch. Scale bars =lmm. two-thirds of distal segment with transverse ridges on the concave medial margin which appears anteriolaterally placed when viewed dorsally (Fig 1 A). Frontal appendage (Fig. 1C) about half body length and consisting of a long narrow trunk and two simple branches about a fifth of the length of the trunk. Trunk weakly pseudosegmented with small, simple, blunt papillae spaced along each lateral margin. Branches distal two thirds bearing dense papillae on both margins of the branches and continuing on the ventral surface in rows of 3-5 papillae. Rigid basal portion of penes (Fig. 1 D) cylindrical and protruding beyond the genital segments onto the first abdominal segment. Each attended laterally by a small rounded protrusion, smaller than the bases of the penes. Eversible portion of penes (Fig. 1 E) about twice as long as rigid base and margined with a row of triangular denticles laterally and similar denticles apically on medial side, but soon becoming spike-like and crowded on the distal two-thirds of the penis. Fifth thoracopod (Fig. 2A) with endites 1+2 (fused) and 3 broad and with evenly curved margins bearing numerous setae; endite 1+2 about 4 times the size of endite 3. Each endite with a one-sided pectinate anterior setae, the second setae about half as long again as the first setae. Both attended by a small spine proximally. Endites 4-6 small asymmetrical protrusions covered in small spines. Endites 4, 5 and 6 with 2,2, and 1 anterior setae respectively, the distal member of each pair distinctly longer than the proximal member. These antenior setae plumose. Endites 4, 5 and 6 with 3, 2 and 2 posterior setae respectively. Endopodite broadly rounded, almost quadrangular bearing many plumose setae, long on the lateral margin decreasing to short on the medial margin. Bases of these setae unadorned. Exopodite twice as long as endopodite, suboval and bearing numerous plumose setae. Epipodite suboval, widest proximally and longer than the endopodite; margin unadorned. Praeepipodite twice as long as broad; margin with a series of small asymmetrical spines. Telson with cercopods subequal in length to the three posterior-most abdominal segments and bearing plumose setae on both margins. Setae longest midway along each cercopod. NEW SPECIES OF BRANCHINELLA 443 FIG. 2. Fifth thoracopods: A, Branchinella clandestine i sp. nov.; B, Branchinella mcraei sp. nov. Scale bar = 1mm. Female. Length of allotype 12.2mm. No other female lengths available. First antennae (Fig. 1 F) slightly shorter than the eye plus eye stalk, and also shorter than the second antennae. First antenna setae as in male. Second antennae subcylindrical, tapering proximal ly and slightly longer than eye stalk plus eye; apex rounded and bearing a sharp projection Hanked laterally by a small longitudinal notch. No setae on distal margin. Brood pouch (Fig. 1G) extending over three abdominal segments, distal part cylindrical. Thoracopods and cercopods similar to those of male. DIAGNOSIS. Male with frontal appendage of a trunk and 2 simple branches. Each branch about 1/5 length of the trunk. Distal segment of second antenna curved medially and anteriorly and twisted so that lateral surface and medial surfaces appear opposite to their usual position. REMARKS. This species is most similar to B. qffinis , B. latzi , B. longirostris and the other new species described here, B. mcraei . All share a broadly similar frontal appendage; i.e. an appendage consisting of a trunk and two simple branches and with sensory papillae. Of this group, B. longirostris readily separates as it has spines at the base of the branches and a lateral protuberance proximal to these spines (Timms, 2004, fig. 70). The relative length of the trunk and branches separate B. clandestina from B. qffinis and B. mcraei — in B. clandestina the branches are about one fifth of the total length of the frontal appendage (Fig. 1C), whereas in B. qffinis they are about half (see Timms, 2004, fig 69) and in B. mcraei and B. latzi about one quarter to one third (Fig. 4B and ibid, fig 71). B. clandestina has the distal segment of the second antenna curved in three planes, so that it is curved medially and anteriorly (Fig. IB), whereas in the other four species this segment is curved only medially (Figs 3, 4A and ibid, figs 69, 70, 7 1 ). Furthermore there is a difference in the number of sensory papillae in the four species — in B. latzi both trunk and branches are covered in papillae (ibid, fig 71), compared to papillae only on the distal half of the frontal appendage (i.e. the branches plus some of the trunk) in B. mcraei (Fig. 4B), only on the distal two thirds of the branches in B. clandestina (Fig. 1C) and not at all in B. qffinis (ibid, fig. 69). Other less diagnostic differences between the 5 species concern the basal segment of the second antenna and the base of the penes. Considering the second antenna, only the two new species have a pad of papillae medially on the unfused distal part of the basal segment. B. qffinis usually has a small area of papillae medioapically on the 444 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Lateral view of 6 Branchinella mcraei sp. nov. Scale bar - 1mm. basal segment, while B. longirostris and B. latzi lack such papillae. Four of the fi ve species have lateral processes on the base of the penes, the exception being B. latzi. Not surprisingly for Branchinella (Geddes, 1981; Timms, 2004) the females cannot be distinguished apart, except for B. clandestine and B. longirostris which have bulbous second antenna. These two can be separated by the prominent apical point being centrally placed in B. longirostris , while B. clandestine! has a small apical point medially displaced and a notch lateral to this (cf. fig. 89 in Timms, 2004 with Fig. IF). Branchinella mcraei sp. nov. (Figs 2-4) ETYMOLOGY. For Jane McRae, Western Australian Dept of Conservation and Land Management, Wanneroo. who collected the specimens and previously two other new * fairy shrimp from Western Australia (Timms, 2002). MATERIAL. HOLOTYPE: 6 West Australian Museum (hereafter WAM) C34035. ALLOTYPE: 9 WAM C34036. PARATYPES: WAMC34037 five 66 and 5 9 9 . OTHER MATERIAL. 1 3 50), thinner and longer than exopodite setae, but still plumose and with pedestals crowned with little spines (3-6). Epipodite oval three times longer than wide and unadorned. Praeepipodite twice as long as broad and with minor crenulations on lateral margin. Telson with cercopods about as long as last two abdominal segments. Cercopods bear plumose setae of both margins; these setae almost subequal along each cercopod, but definitely shorter proximally and apically. Female. Length of adults 20-33mm, paralectotype 28mm. First antennae (Fig. 6D) filiform and about one third the length of second antenna. Second antenna lamellar with a rounded asymmetrical apex and reaching back to 2nd or 3rd thoracic segments. Brood pouch (Fig. 6E) extending over genital segments and almost four abdominal segments; distal part cylindrical. Thoracopods and cercopods similar to those of male. DIAGNOSIS. Male lacks a frontal appendage, but has an apomedial antennal appendage from basal segment of second antenna. Proximally this appendage is lamellar with numerous long lateral papillae. Base of penis without a transverse protrusion laterally. REMARKS. Material from Gidgee Lake differs slightly from the type material from Lake Buchanan and also from the Hatch Lake sample. The antennal appendage has fewer long papillae than in the Lake Buchanan specimens, with only 7-8 anvil-tip papillae and 4 of the spear-pointed papillae in the September 1 998 sample. The May 2000 sample differs further in that these long NEW SPECIES OF BRANCHINELLA 449 papillae are without specially modified apices and are also fewer in number (Fig. 5F). Also the surface of the claspers is unadorned. Given that these Gidgee specimens (mean length 18.7mm) are smaller than Buchanan specimens (mean length 24.8mm) and the September 1998 Gidgee males (mean length 28mm), they may be undeveloped (meaning that the apices differentiate later in development, and that adornment of the distal segment of the second antenna are also a late-developing feature). On the other hand, all Gidgee specimens had a reduced number of long papillae on the apomedial outgrowth of the second antenna, so this character is not fixed in this species. Branchinella hattahcnsis Geddes, 1981 (Figs 5C,D, 8) Branchinella nichollsi hattahensiss Geddes, 1981: 264, fig 5b. Branchinella hattahcnsis Geddes; Belk & Brtek, 1995: 324. ETYMOLOGY From Hattah Lake. MATERIAL. LECTOTYPE: <3,QMW26946, November 1971. coll. G Arthur, PARALECTOTYPES: four < 36 , one 9, QMW26947 November 1971, coll. G Arthur. OTHER MATERIAL: QM W26948, Lake Numalla, 28°42’S, I44°I9*E, QM W26949, Mid Kaponyee Lake, Currawinya National Park, SW Qld, 28°50'S, 144° 1 9' E, QM W26950, South Kaponyee Lake, Currawinya National Park. TYPE LOCALITY. Hattah Lake, 34°44’S, 142°2rE, NW Victoria. DESCRIPTION. Male. Length of adults 18-44mm; lectotype 44mm. First antennae subequal in length to proximal segment of the second antenna (Fig. 8A); apex bevelled and bearing subapically a tuft of 2-3 subequal short setae. Second antennae proximal segments (Fig. 8A) fused at base with approximately half of distal portion free, cylindrical and aligned with body axis. Clypeus with a blunt triangular outgrowth ventromedially. Distal segment of second antennae with a swollen asymmetrical base with long, thin remaining portion, rounded in cross section, slightly curved medially but near apex curvative reversed so that apex curved laterally. Distal segment about 1.5 times longer than proximal segment. Dorsal surface of distal segment granulated, ventral surface with small raised transverse ridges. Frontal appendage absent. Antennal appendage present on medial surface of proximal segment of second antenna (Fig. 8A). Antennal appendage tubular, narrowing only a little along its length and almost as long as the distal segment of antenna; numerous short papillae on apical half. Rigid basal portion of penes (Fig. 8B) cylindrical and protruding onto approximately a third of the first abdominal segment. Mid length each penis base with a short transverse protrusion laterally and attended laterally by a pointed soft outgrowth about same length as bases of penes and separated from them basally by a hemispherical cavity. Everted penis about 3mm long and almost the length of two abdominal segments. A narrow row of broadly based asymmetrical spines on lateral surface and a broad row of similar spines on medial surface near apex, but subapically changing to numerous narrow spines. All spines point basally. Fifth thoracopod as in Branchinella huchanan- ensis but with epipodite relatively shorter, i.e. length twice width. Telson and cercopods also as in Branchinella huchananensis. Female. Length of adults 16-45. 5mm, paralectotype 45.5mm. First antennae (Fig. 8C) filiform and about 1/4 length of second antenna. Second antenna lamallar with a markedly asymmetrical apex with a blunt point and reaching back to 2nd or 3rd thoracic segments. Brood pouch (Fig. 8D) extending back over 2 genital segments and 3.5 abdominal segments; distal part cylindrical. Thoracopods and cercopods similar to those of male. DIAGNOSIS, Male lacks a frontal appendage, but has an apomedial antennal appendage from basal segment of second antenna. Antennal appendage tubular with short papillae on apical half. Clypeus with a blunt triangular outgrowth ventromedially. Base of penis with a transverse protrusion laterally. REMARKS. The Queensland material is slightly different from the Victorian specimens in that the antennal appendage is only about 3/4 length of the distal segment of the second antenna and has fewer papillae (Fig. 5D). This difference may not be phenotypic, but could be developmental (cf. remarks on B. huchananensis). DISCUSSION Branchinella clanclestina sp. nov. and B. mcraei sp. nov. have characteristics which place them in Geddes (1981) Group II: both have a frontal appendage consisting of a long trunk and two simple branches, short endopodites, and 2-4 anterior setae on endites 4-6. Geddes (1981) 450 MEMOIRS OF THE QUEENSLAND MUSEUM FIG 7. Fifth thoracopod of Branchinella buchananensis Geddes. Scale bar = 1mm. places the B. nichollsi complex ( B . nichollsi , B. hattahensis , B. buchananensis) among his Group I species, because of their large, robust bodies, long endopodites which are sparsely setulated medially, numerous anterior setae on endites 4-6, and females with long second antennae. While these similarities seem valid morphologically, relationships between species based on mitochondrial DNA did not support these 2 groupings, though it did recognise his Group III (Remigioetal.,2003). Group II species separated on their DNA affinities as 2 groups quite distinct from each other; it is not known to which of the 2 subgroups the new species belong . Group 1 species also comprise 2 well separated groups, with B. hattahensis and B. buchananensis comprising 1 subgroup (and presumedly B. nichollsi also belongs here). The other subgroup contains B. australiensis and B. occidentalis. The 3 species in the B. nichollsi group are closely related, probably more so than other species of Branchinella. Based on mitochondrial DNA evidence, Remigio et al. (2003) claimed they are of subspecies status, but there are many distinct morphological differences between the three. In antennal features, B. buchananensis is quite different with its antennal appendage having many long papillae on basolateral surface and shorter papillae apically, compared to only a few short papillae mainly in the apical half in B. nichollsi and B. hattahensis . In the later pair there is some variation between populations in these papillae (for B. nichollsi compare Fig. 5A with 5B; and for B. hattahensis compare Figs 5C, 8A with 5D). These 2 species are however easily separated on antennal features, since B. hattahensis has a medial process on the clypeus and B. nichollsi does not. The 3 are also easily separable on features of the male genital area. Both B. hattahensis and B. buchananensis have a hemispherical cavity between the penis base and lateral outgrowth, whereas B. nichollsi does not. B. hattahensis is unique in having a transverse ridge on the penis base. The penes themselves are similar in structure in B. buchananensis and B. hattahensis: no data are available for B. nichollsi. Another difference between the species is in the structure of the fifth thoracaopod, with this time B. nichollsi being the most different. It has 7:7:5 posterior setae on endites 4-6, whereas the other two have 6:5:5 respectively. This slightly greater number of posterior setae is reflected elsewhere on the thoracopod, e.g. ca 50 on the first endite in B. nichollsi compared to ca 40 in the other 2 species. These differences are about the same order of magnitude as perceived differences among the B. affinis group (B. afjinis , B. clandestina sp. nov., B. latzi , B. longirostris , B. tncraei sp. nov. — see earlier in remarks about B. clandestina sp. nov.) and also, for example, between B. halsei and B. lyrifera (Timms & Geddes, 2003). These comparisons indicate that division at the species rank is warranted. The females of all 3 forms are inseparable from each other, not unusual for Branchinella (Geddes, 1981: Timms, 2004). However, B. buchananensis, B. hattahensis and B. nichollsi together are distinguishable from other females of Branchinella (Timms, 2004). This is because of the lamellar second antennae (Fig. 6B) being about twice the length of the intereye distance, and so much shorter than the lamallar antenna of B. australiensis and B. occidentalis and much bigger than those of most other species. The brood sac (Fig. 6C) tends to be almost 6 segments long overall and thus is a little longer than in many species of Branchinella where it is about 5 segments long. NEW SPECIES OF BRANCHINELLA 451 FIG. 8. Branchinella hattahensis Geddes. A, dorsal view of 6 head; B, ventral view of 6 genital area with one penis everted; C, dorsal view of head of 9 ; D, lateral view of brood pouch of 9 . Scale bars = 1mm. Ecologically, there are similarities but some differences between the 3 species. Geddes (1981) thought, on limited evidence, that all 3 species were halophilic; certainly B. buchananensis was reported to live in waters from 15.7-42.6g/L. However Timms (2002) showed that B. nichollsi lives in fresh — hyposaline conditions and the highest actual salinity recorded is only 1 .5g/L in Lake Arrow (Chapman & Timms, in press). My unpublished records from the Paroo show B. hattahensis can live, at most, in subsaline water (i.e. < 3g/L) and B. buchananensis lives in hyposaline waters to 1 5. lg/L. So it seems that B. buchananensis is the species with the greatest halotolerance, with the other 2 only slightly more tolerant than most other Branchinella , except B. simplex and B. australiensis (Geddes, 1973). Halotolerance refers to NaCl waters, virtually the only type in Australia (Hart & McKelvie, 1986). Another ecological difference between the 3 species is that B. nichollsi and B. buchananensis live in lakes that are usually have clear water, whereas B. hattahensis lives in turbid freshwater lakes that may increase in salinity as they dry. B. nichollsi is apparently confined to a small area in the eastern Goldfields of WA, while B. hattahensis occurs from N Victoria to SW Qld, with the northern part of this range overlapping with the distribution of B. buchananensis which is now known from NW NSW to NE Qld. The area of overlap is quite small and is restricted to a small area near Hungerford, SW Qld; in this area, as elsewhere, the 2 species inhabit different types of lakes (Timms & Sanders, 2002). While the 3 forms in the B. nichollsi complex have many morphological similarities, some ecological similarities and are very closely placed in the phylogram of Remigio et al. (2003), I believe they are distinct species, as did Belk & Brtek (1995). Despite some variation between populations, some of which is explainable as variation due to stage of development (as in B. buchananensis in Gidgee Lake), each is now known from a number of sites and they are always distinct morphologically. Furthermore, B. buchananensis and B. hattahensis are sympatric but apparently do not interbreed. The 5 species discussed here are of very unequal conservation status. B. buchanenensis has legal status as a vulnerable fish species in NSW (Anon, 2002), but it is unprotected in Queensland. The rationale for this is the limited habitat in NSW is threatened by gypsum mining whereas in Queensland there are no threats to its habitat. The other 4 species are unprotected, and although rarely collected, the habitats of 3 of them are unthreatened. The exception is B. nichollsi whose habitat is some episodic salinas in the eastern goldfields of WA (Timms, 2002). The problem is that the hydrology of some of these is being changed by either mining directly on the lake or using the lake as a receiving basin 452 MEMOIRS OF THE QUEENSLAND MUSEUM for saline groundwater. In that the species apparently hatches and grows only in fresh or slightly saline water, the addition of extra salt in its habitat could threaten its existence (see Timms, in press). If IUCN (2000) Red List criteria were applied to the conservation status of these species, all would be classed as ‘data deficient" because so little is known on their biology, but more research would probably result in a ‘vulnerable’ classification for B. nichollsi and B. buchananensis . ACKNOWLEDGEMENTS I thank Jane McRae for not only collecting the species named in her honour but particularly for drafting Figs 1, 3 and 4. I acknowledge with thanks the cooperation of the Qld National Parks and Wildlife Service for a permit to collect invertebrates in Currawinya National Park and to the present owner of Bloodwood Station, Reg Collins, for permission to collect on his lease. I am also indebited to Michael Geddes for providing original collections of B. buchana- nensis and B. hattahensis , to Buz Wilson for advice and to Christopher Rogers for helpful comments on the manuscript. LITERATURE CITED ANON 2002. NSW Fisheries - Fishnote NSWF 1074. Buchanan's fairy shrimp. BELK, D. & BRTEK., J. 1995. Checklist of the Anostraca. Hvdrobiologia 298: 315-353. CHAPMAN A. & TIMMS, B.V. In Press. Waterfowl usage of Lake Arrow, and arid zone wetland in the eastern Goldfields of Western Australia, following cyclonic rain. Australian Field Ornithology. GEDDES, M.C. 1973. Salinity tolerance and osmotic and ionic regulation in Branchinella austral tens is and B. compacta (Crustacea: Anostraca). Comparative Biochemistry and Physiology 45A: 559-169. 1981. Revision of Australian species of Branchinella (Crustacea: Anostraca). Australian Journal of Marine and Freshwater Research 32: 253-295. HART, B.T. & McKELVIE, l.D. 1986. Chemical limnology in Australia. Pp. 3-3 1 . In Dc Deckker, P. & Williams, W.D. (eds) Limnology in Australia. (CSIRO/Junk: Melbourne/Dordrecht). INTERNATIONAL UNION OF CONSERVATION OF NATURE AND NATURAL RESOURCES, 2000. IUCN Red List of Threatened Species, compiled by Craig Hilton-Taylor. (IUCN - The World Conservation Union Species Survival Commission: Cambridge). LINDER, F. 1941. Contributions to the morphology and the taxonomy of the Branchiopoda Anostraca. Zoologiska Bidrag fran Uppsala 20: 101-302. REMIGIO, E.A., TIMMS, B.V. & HEBERT, P.D.N. 2003. Phylogenetic systematics of the Australian fairy shrimp genus Branchinella based on mitochondrial DNA sequences. Journal of Crustacean Biology 23: 436-442. TIMMS, B.V. 2001. Two new species of fairy shrimp (Crustacea: Anostraca: Thamnocephalidae: Branchinella) from the Paroo, Inland Australia. Records of the Australian Museum 53: 247-254. 2002. The fairy shrimp genus Branchinella (Crustacea: Anostraca: Thamnocephalidae) in Western Australia, including a description of four new species. Hydrobiologia 486: 71-89. 2004. An identification guide to the fairy shrimps (Crustacea: Anostraca) of Australia. Cooperative Research Centre for Freshwater Ecology, Identifiaction and Ecology Guide 47. In Press. Salt lakes in Australia: present problems and prognosis for the future. Hydrobiologia. TIMMS, B.V & GEDDES, M.C. 2003. The fairy shrimp genus Branchinella Sayce 1903 (Crustacea: Anostraca: Thamnocephalidae) in South Australia and the Northern Territory, including descriptions of three new species. Transactions of the Royal Society of South Australia 127: 53-68. TIMMS, B.V. & SANDERS, P.R. 2002. Biogeography and ecology of fairy shrimps (Crustacea: Anostraca) in the middle Paroo catchment of arid-zone Australia. Hydrobiologia 486: 71-89. Volume 50 Memoirs of THE Queensland Museum volume 50 2005 PUBLISHED BY ORDER OF THE BOARD VOLUME 50 IS COMPLETE IN TWO PARTS © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 61 7 3840 7555 Fax 61 7 3846 1226 National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Memoirs of the Queensland Museum may be reproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Director. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site www.qmuseum.qld.gov.au/resources/resourcewelcome.html. A Queensland Government Project Typeset at the Queensland Museum Printed by Watson Ferguson & Co 35 Hamilton Road, Moorooka, Queensland 4105 CONTENTS PART 1 (Issued 16 August 2004) JELL, P.A. The fossil insects of Australia 1 PART 2 (Issued 10 January 2005) AMEY, A.P., KUTT, A.S. & HUTCHINSON, M. A new species of Lerista (Scincidae) from central Queensland 125 BAEHR, M. A revision of the Australian odacanthine ground beetles, including checklists for Australia and the Papuan subregion (Insecta: Coleoptera: Carabidae) 133 DAVIES, V.T. Teeatta , a new spider genus from Tasmania, Australia ( Amaurobioidea: Amphinectidae: Tasmarubriinae) 195 GERSHWIN, L. Spectacularia vanoppenae gen. et sp. nov., a new hydromedusa (Cnidaria: Hydrozoa: Dipleurosomatidae) from the Great Barrier Reef 201 HOLT, T.R., SALISBURY, S.W. & WILLIS, P.M.A. A new species of mekosuchine crocodilian from the middle Palaeogene Rundle Formation, central Queensland 207 HYMAN, I.T. & STANISIC, J. New charopid land snails chiefly from limestone outcrops in eastern New South Wales (Eupulmonata: Charopidae) 219 KARANOVIC, I. A new Candoninae genus (Crustacea: Ostracoda) from subterranean waters of Queensland, with a cladistic analysis of the tribe Candonopsini 303 KYNE, P.M., JOHNSON, J.W., COURTNEY, A.J. & BENNETT, M.B. New biogeographical information on Queensland chondrichthyans 321 PATTEMORE, G.A. & RIGBY, J.F. Fructifications and foliage from the Mesozoic of southeast Queensland 329 RAVEN, R.J. & STUMKAT, K.S. Revisions of Australian ground-hunting spiders: II. Zoropsidae (Lycosoidea: Araneae) 347 SACHS, S. Tuarcingisaurus australis sp. nov. (Plesiosauria: Elasmosauridae) from the Lower Cretaceous of northeastern Queensland, with additional notes on the phylogeny of the Elasmosauridae 425 TIMMS, B.V. Two new species of Branchinclla (Anostraca: Thamnocephalidae) and a reappraisal of the B. nichollsi group of the Australian arid-zone 441 VAN DYCK, S. & GYNTHER, 1. Nesting strategics of the Water Mouse Xeromys myoides in southeast Queensland 453 WALL, M.A. & CASSIS, G. Diabolicoris , a new genus in the tribe Ploiariolini (Hemiptera: Reduviidae: Emesinae) from New Caledonia 481 NOTES BLOME, D. Portmacquaria nom. nov. pro Macquaria Blome, 2002 (Nematoda: Chromadoridae) 132 * ^ 1133 CONTENTS AMEY, A.P., KUTT, A.S. & HUTCHINSON, M. A new species of Lerista (Scincidae) from central Queensland 125 BAEHR, M. A revision of the Australian odacanthine ground beetles, including checklists for Australia and the Papuan subregion (Insecta: Coleoptera: Carabidae) 133 DAVIES, V.T. Teeatta , a new spider genus from Tasmania, Australia (Amaurobioidea: Amphinectidae: Tasmarubriinae) 195 GERSHWIN, L. Spectacularia vanoppenae gen. et sp. nov., a new hydromedusa (Cnidaria: Hydrozoa: Dipleurosomatidae) from the Great Barrier Reef 201 HOLT, T.R., SALISBURY, S.W. & WILLIS, P.M.A. A new species of mekosuchine crocodilian from the middle Palaeogene Rundle Formation, central Queensland 207 HYMAN, I.T. & STANISIC, J. New charopid land snails chiefly from limestone outcrops in eastern New South Wales (Eupulmonata: Charopidae) 219 KARANOVIC, I. A new Candoninae genus (Crustacea: Ostracoda) from subterranean waters of Queensland, with a cladistic analysis of the tribe Candonopsini 303 KYNE, P.M., JOHNSON, J.W., COURTNEY, A.J. & BENNETT, M.B. New biogeographical information on Queensland chondrichthyans 321 PATTEMORE, G.A. & RIGBY, J.F. Fructifications and foliage from the Mesozoic of southeast Queensland 329 RAVEN, RJ. & STUMKAT, K.S. Revisions of Australian ground-hunting spiders: II. Zoropsidae (Lycosoidea: Araneae) 347 SACHS, S. Tuarangisaurus australis sp. nov. (Plesiosauria: Elasmosauridae) from the Lower Cretaceous of northeastern Queensland, with additional notes on the phylogeny of the Elasmosauridae 425 TIMMS, B.V. Two new species of Branchinella (Anostraca: Thamnocephalidae) and a reappraisal of the B. nichollsi group of the Australian arid-zone 441 NOTES BLOME, D. Portmacquaria nom. nov. pro Macquaria Blome, 2002 (Nematoda: Chromadoridae) 132