MEMOIRS QUEENSLAND MUSEUM BRISBANE = ~— ~~ VOLUME 30 CONTENTS Abstract ,..,.-..-. ne cinett. tes sy:2 fate 1 Introduction ............--..4- riatehyet l Abbreviations Used 21.5.2... 0.0.2, 000. 2 Material... 00.022... ee ee eee eee 2 Previous Studies .......-..--2-06-0055 3 Distribution, Habitat, Ecology and Conservation... .. 6... cee ee eee eee 4 Patterns of Variation ,.............-2.. 5 Systematics 00... 00... ce eee eee 14 CHAROPIDAE ........,....-0.---55 14 CHAROPINAE ........,0 00 cue e eens 14 Ngairea Zen. MOV. oc cee cece eee ces 14 N. dorrigoensi§s ... 0c cee ee eee 16 N. levicostata sp. NOV. 2.2.20. eevee 23 N, murphyi oo... eee cee 26 N. canaliculata sp. NOV... eee 27 N. corticicola ..... 006.6. c cp 31 Mussonula . 00.2 eee 36 MLverax oo. 2 ep eee a pees 38 M., fallax sp. NOV. 2.26. e eee ees 39 Hedleyaconcha wi. cece tees 42 FEAR. on A A aig ara a a A as as 43 A. ailaketoae sp. nOv. .........6.-64- 48 Fea ai ta oo Fh doco ny Be a eagle sts 51 Lenwebbia gen. nov... ..-. 2.040200 ». 52 L. protoscrobiculata sp. NOV. «1.6.44. 52 Setomedea . 0... ccc cee cee ens 57 S. seticoStata 6.0... eee eee 59 S. nudicostata 8p. NOV........-..+4--. 64 S, JQMA@ SP. NOV. . 6.6 eee eee cee 69 S. monieithi sp. MOV... eee es 72 Oi Shes ye Guibas eerste rae hh as 77 Gyrocochlea oo... ee eee 77 G. vinitincta ..... pees Cts bay says 79 G. CONVOIMIA oo eee 86 G, paucilamellaia sp.NOV oo... .. 5.5 e. 87 G. CUFUSIAQNA oo eee eae 90 Biomphalopa gen. nov. .... 0.6... 02. ee 94 BL r@cava oo ee eee cere ees 195s tHe 95 BL COMCINNG Coc eee 98 Nautiliropa gen. NOV, 2.0225 eee 101 N. OMICTON 000 cece tees 102 Letomold occ eee eee 109 L, CONIOTIUS Co ete eee 110 Rhophodan .. cc. ccc eect een aes 114 R. Peregrinus 0.0... cece ee 120 R. CONSOBTINUS 2 eee 121 R. kempseyensis Sp. MOV. 0.24... e cee 125 R, elizabethae sp. MOV. 6.6... eee ees 129 R. minutissimus sp. NOV. vo... eee eae 132 R, colmani sp. n0V. 22.622 ee eee 136 R, bairnsdalensis .....0...-2.00..045 , 138 Discocharopa .... 0... cee ieee 143 D, APTA Lecce ccc eee teens 143 Cralopa wo... ccc eevee eens ytatEta 147 C, stroudensis 00... e eee ee 150 C. kaputarensis 8p. NOV. ...-....2644- 153 C. carlesSisp. NOV. oo... eee ee eens 156 EISOtherd oo eee ies 160 EL sericatula .2iiais ct es aia beeeaes 161 E. mautilodea .... 0.0... c cee eee es 164 E. genithecata sp. Nov. .-.. 0.622545 166 Coenocharopa gen. NOV... 6. cee ee es 169 C, sordidus $p. NOV. 61... eee es 172 C, macromphala sp, nov. ...-......-- 176 C. alata Sp. NOV... 66. eee ee 179 C. parvicostata sp. NOV. 0... ... 6265 ee 186 C. yessabahensis sp. NOV. vi... eee e ues 187 C. multiradiata sp. nOV. 0.1.0.6... ees 190) ERilOMeR. oe. ae pbi thie ttid pes gis 193 E. cochlidium . 0.02.00 oc ee ce ee 193 E. globosa sp. NOV... oe eee eee es 197 Omphaloropa gen, NOV. «2... 06s eee 198 O. VariCOS@ SP. NOV... ee ee ees 201 Sinployea ..... TPey see eT SS see ashe) 205 S.UMONSA oo eee eee ees 206 ROTADISCINAE, ,.......-.--....0-. 210 Rotacharopa gen. NOV. ... eee eee ee ees 210 R. aunabelli sp. nOV. .... 0. eee ees 211 R. kessneri Sp. NOV. vec ee eee 213 R. densilamellata sp. nov. ..........-- 219 Extralimital Relationships ,............ 223 Phylogeny ......2. 20... cc ese eeeecaes 226 Biogeography .. 2.2.2... 06252. 0.000e 227 Conclusions. ........ 00. .c cere ee eee 234 Acknowledgements .........-..0000 05 235 Literature Cited .. 2.2.0... eee eee 235 Appendix... 0.25.22 e eee eee eee eee 238 SYSTEMATICS AND BIOGEOGRAPHY OF EASTERN AUSTRALIAN CHAROPIDAE (MOLLUSCA, PULMONATA) FROM SUBTROPICAL RAINFORESTS JOHN STANISIC Stanisic, J. 1990 12 24: Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pulmonata) from subtropical rainforests. Memoirs af the Queensland Museunr, 30(1)1-241. Brisbane. ISSN 0079-8835. Fifty land snail species in 18 genera of Charopidae from eastern coastal Australia are reviewed, Ngairea. Lenwebbia, Omphaloropa, Nautilirapa, Coenocharopa. Biomphalopa and Rotacharopa are new. Exgilodonta Iredale, 1937 is placed in synonymy with Rhophodon Hedley, 1924. Ngatrea levicostata. N. canaliculata, Mussonula fallax. Hedleyuconcha ailaketoae, Lenwebbia protoscrobiculata. Setomedea moantetthi, S. nudivostata, S. janae, Gyrocochiea paucilamellata, Rhophodon minutissimus, R, colmani, R, elizabethae, R. kempseyensis, Cralopa carlessi, C. kaputarensts. Egilomen globosa, Elsothera genithecata. Omphaloropa varicosa, Coenocharopa sardidus, C. macramphala, C. yessabahensis, C. parvicostata, C. multiradiata. C, alata. Rotacharopa annabelli, R. kessnert, R. densilamellata (27 species) ure new, Neoivpes are nominated lor Helix murphyi Cox, 1864, Helix omicron Pfeiffer, 1851 and Rhophodon peregrinus Hedley, 1924, All except 3 species are assigned to the Charopinae: those 3 species. assigned to the Rotadiscinae, are the first record of the subfamily in Australia, Analysis of shell and anatomical patterns is presented: differences in shell, radular, pallial and genital structures are evaluated. Patterns of variation are correlated with three levels of functional and ecological significance - species recognition, exploitation of adaptative zones, and the need to conserve water. Morphological pallerns of Australian species are compared with those of New Zealand and the Pacific Islands and their relationships are discussed. The proposed phylogeny identifies major morphological irends and multiple origins of the fauna. A cladistic analysis (PAUP) utilises 7S shell and anatomical characters, A biogeographic synthesis integrates distributions, ranges, and ecology with aspects of past climatic. geologic and vegetational change. Charopid evolution in eastern Australia is closely linked to climate induced changes in mesic communilies since the Cretaceous. CF Mollusca, Pulmonata, Charapidae, Systematics, Phylogeny, Biageagraphy. John Stanisic, Queensland Museum, PO Box 300, South Brisbane, Queensland 4101, Australia; 19 October 1989. The Charopidae 1s.a family of small to minute, radially ribbed, pulmonate land snails abundant in leaf litter of eastern Australian rainforests. A few species also occur in drier forests throughout the rest of the continent.Burch (1976), based on the earlier checklists of Iredale (1937a, 1937b, 1941a, 1941b), listed 110 species and 29 genera of Charopidae from Australia. The Australian Museum/Queensland Museum cast coast rainforest surveys of the mid-1970's recorded 38 charopid species (28 new) from northern New South Wales, southern and central Queensland (Broadbent and Clark, 1976). Sur- veys by the Queensland and Australian Museums (1981-1983) in subtropical rainforests of northern New South Wales and southern Queensland revealed an even greater number of undescribed species from this regian. Collecting by the author has exposed extensive radiations of charopids in central Queensland mesic forests and on the mountain peaks of the Ingham-Cook- town rainforest massif in tropical Queensland (Stanisic, 1987). Previously, the Charopidae were tegarded as a mainly coal temperate group with greatest diversity in Victorian and Tas- manian forests. In general, eastern Australian charopids dis- play an intimate ecological bond with rain- forests. Most of the 50 species (18 geneta) reviewed have subtropical distributions although it has been necessary to include several temperate and tropical taxa, The aims are to initiate a survey of the subtropical Charopidae, to identify patterns ice) of conchological and anatomical variation, and to place them in biogeographical context. The species reviewed here may represent only 15% of Australian charopids. Therefore phylogenetic conclusions are based on the analysis of a seemingly mosaic pattern of varia- tions. Subtropical eastern Australia is defined as the coastal to subcoastal area between Dorrigo, New South Wales and Rockhampton, Queensland. Floristically, this area encompasses the distribu- tion of dry and moist subtropical closed forests (Fig. 1). ABBREVIATIONS USED INSTITUTIONS AM Australian Museum, Sydney BMNH British Museum (Natural History), London FMNH Field Museum of Natural History, Chicago NMV Museum of Victoria, Melbourne SAM South Australian Museum, Adelaide QM Queensland Museum, Brisbane COLLECTORS/COLLECTION DATA AM/QM-ABRS: Australian Biological Resources Survey funded field work (1981- 1983) involving various collectors from the Australian and Queensland Museum. Earthwatch/QM: American Earthwatch spon- sored fieldwork involving volunteers and various staff of the Queensland Museum. VEGETATION TYPES CNVF, complex notophyll vine forest; NVF, notophyll vine forest; MVF, microphyll vine forest; CMVF, complex mesophyll vine forest; SEVT, semi-evergreen vine thicket; SMVFF, simple microphyll vine fern forest; SMVFT, simple microphyll vine fern thicket. ANATOMICAL TERMINOLOGY DG, prostate; E, epiphallus; EC, epiphallic caecum; EF, epiphallic flagellum; EP, epiphallic pore; EPP, epiphallic pilaster; ERC, epiphallic retractor caecum; G, ovotestis; GD, hermaphroditic duct; GG, albumen gland; GT, talon; H, heart; HG, hindgut; HV, principal pul- monary vein; I, intestine; K, kidney; KD, ureter; KX, ureteric pore; MA, mantle lobe; MC, mantle collar; MG, mantle gland; P, penis, PP, penial pilaster; PPM, main penial pilaster; PPT, prepu- tial tube; PRM, penial retractor muscle; PV, penial verge; S, spermatheca; SP, sper- matophore; SS, spermathecal stalk; UT, uterus, UV free oviduct; UVP, free oviduct pilaster; V, MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. Distribution of selected rainforest floristic regions. provinces and areas in eastern Australia. a, humid tropical northern region; b, subtropial forests of the Eungella area; c, dry to moist subtropical province: d, warm temperate to cool temperate province: e, temperate forests of the Lamington Plateau; f, cool temperate province; g, dry subtropi- cal province. (Adapted from Webb and Tracey, 1981). vagina; VC, vaginal caecum; VD, vas deferens; X, carrefour; Y, atrium; Z, digestive gland. MATERIAL This study is based on 6230 specimens mainly in the Queensland Museum, Brisbane and the Australian Museum, Sydney. Most specimens have been collected since 1980. *Wet’ material available was <10% of the total specimens; 150 specimens (44 species) were dissected. The material contains a high proportion of juvenile and subadult specimens because in the early stages, when species presence was being established, most specimens were obtained by extraction from collected litter. This material consisted of a large number of dead juveniles which reflects the overall higher field mortality among young snails. Later collecting was con- centrated on live adults for dissection; so ’wet’ SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 3 Tewirdlen Cree FIG, 2. Distribution of major rainforestareas in north- ern New South Wales and southern Queensland. material includes a greater proportion of adults. However, because many species are difficult to locate live in the field, individual ‘wet’ lots often contain only single or few specimens. Scasonal climatic effects may be a contribut- ing factor to the numbers of live adults versus live subadults present at any time, but the small amount of wet material and uneveness of callect- ing effort through the year prevent sensible con- clusions: only a few species were abundant enough for seasonal studies, Line drawings of shells and dissections were made by the author using a camera lucida attach- ment. Measurements are recorded using the method of Solem (1976, pp. 11-15). PREVIOUS STUDIES Previous studies of Australian Charopidae have been introductions in broader faunal cheek- lists except for works of Hedley (1924), Solem (1984), Smith and Kershaw (1985) and Stanisic (1987) who dealt specifically with Australian charopids, With the exception of Hedley (1889), who figured the radula and jaw of Hed- levoconcha delta (Pfeiffer, 1857), and Odhner (19L7) who figured the jaw and radula of Pilshrycharopa tumidus (Odhner, 1917), malacologists of the late 19th and early 20th century used conchological details to establish species and geners, Smith and Kershaw (1985) and Stanisic (1987) provided anatomical details for several genera and species from the Tas- mania-Victoria and north Queensland areas, respectively, Iredale (1937a, 1937b) compiled records up to that date. Subsequent works introducing new species are Gabriel (1947), Gabriel and Mac- pherson (1947), Cotton (1939), Iredale (1937c, 1939, 1941) and Stanisic (1987). Mostearly shell-based descriptions are of little value In contemporary systematics. Only Hed- ley (1912, 1924) set an acceptable standard. Solem (1983) set a modern standard with his review of Pacific Island taxa: he showed the importance of soft parts in species diagnoses. CLASSIFICATION Solem (1983, p. 47) presented a full synonymy of the family level units available for the en- dodontoid complex, and in the Charopidae in- cluded Flammulinidae Crosse, 1894, Dipnelicidae Iredale, 1937 and Hedleyo- conchidae Iredale, 1942. These family names had previously been applied to various sub- divisions of the Australian Charopidae by Gabriel (1930) and Iredale (1937a, 1941b, 1942). Salem (1983, p. 70) recognised five sub- families: Otoconchinae Cockerell, 1893, Rotadiscinae Baker. 1927, Charopinae Hulton, 1884. Semperodoninae Solem, 1983, and Truk- churopinae Solem, 1983. The Charopinae was broadly defined and considered amenable to sub- division subsequent to a revision of Australian and New Zealand taxa. The phylogeny developed in this study utilises anatomical features to define subfamilial, generic and species units. Specifically. the structure of the pallial cavity. gross morphology of the reproductive system, and internal charac- teristics of the terminal genitalia are considered important criteria for phylogeny. Conchological detail such as shell shape. ribbing, and coiling pattern may be convergent and often mask quite crucial differences in anatomy, However, some conchological features including apical sculp- ture, microsculpture and primary shell sculpture show more positive correlations with underlying anatomical patterns, Hence it has been possible to produce a practical classificatory system which utilises shell features. Because this intro- ductory study investigates selected species. from 4 MEMOIRS OF THE QUEENSLAND MUSEUM an area of mid-range diversity for the family, phylogenetic gaps between genera are not ‘even’; they reflect the diverse ecological habits of species studied, Genera revicwed and, where possible, redefined using both shell and anatomical fea- tures are:- Cralopa, Discachuropa, E/sothera, Gvrocachlea, Mussonula, Rhephadon, Seto- medea, Hedleyoconcha, Egilomen, Letomola. Exilodonta is synonymised with Riiephodan. DISTRIBUTION, HABITAT, ECOLOGY AND CONSERVATION In eastern Australia, charopids are mainly found in closed forests (= rainforests). Histori- cal development of this association is examined further in the biogeographic discussion. ‘The strength of this association, antiquity of the Charopidae, and long-term historical persistence of mesic forests in eastern Australia, have com- bined to produce a complex pattern of narrow, wide, disjunct and relict distributions. During arid phases of the mid to late Tertiary, rainforests Would have acted both as refugia and cpicentres of charopid evolution. Today all Australian rainforests are refugia (Webb and Tracey, 1981). Charopids flourish in the four major categories of rainforest tefugia of Webb and Tracey (J981):- A, large, relatively wet areas such as the Border Ranges of southern Queensland, the Dor- rigo Plateau, NSW, and mountain summits such as Mt Warning, NSW, and Mt Glorious, SEQ; B, small, relatively dry, topographic isolates which include rocky limestone outcrops such as the Caves, near Rockhampton, MEQ, Mt Biggen- den, SEQ, and the Kempsey Caves complex, NSW - besides providing protection trom fire and a relatively high level of moisture, these areas also offer an abundant supply of calcium; C, small edaphic isolates where soil types are critical in determining rainforest existence - gal- lery forests growing on fiverine alluvia and the deciduous forests of Cooloola, SEQ, are ex- amples, D, mountain tops in lower rainfall areas where topography, soils and climate combine to ensure persistence of closed forests - examples are the Bunya Mountains and areas of the Many Peaks. Range. SEO and Mt Dryander, MEQ. Tn NQ and MEQ chuaropids are muinly restricted to montane regions indicating that past environ- mental changes have been effective in geographically cornering of species. This al- titudinal factor is less noticeable in the subtropi- cal region where both lowland and foothill rainforests have a rich complement of species, Although rainforest provides ample litter con- taining a vanety of living spaces through which snails can crawl, species investigated here donot exploic the litter zone. Heeleyocuncha is at- boreal with //, dedta living on the leaves of trees. and shrubs. Neairea and Sinployea are semiar- horeal preferring to live under the bark of fallen, rotting trees. Lemwebbia has been found sitting in the Open on rotting logs while Setamedea is the only group known to prefer very damp places in or under logs, Living under logs is the preferred habit for Naurliropa, Gvrocechlea, Biamphalopa, Elsotherq, Retacharopa, Cralopa srroudensis, C. kaputarensis, and Egilomen globosa. Species of Rhaphodon, Letomola con- fortus and Coenocharepa yessabahensis, live on tack surfaces and some (c.g. C. vessabahen- sis) show habitat-related radular Specialisation, Coenocharopa alata lives in moss on logs and rocks. while C. macromphala and C. sordidus probably live among the friable earth below the litter, as do Omphalorapa varicesa and Egdomen cochlidaim, Microhabitats of the other species ure unknown. Feeding habits are unknown, Radular mor-: phology is conservative, rarcly (c.g. in Nauuliropaomreran and Ngairea corticicola) is there any structural indication of possible dictary shift. Sympairy has been noted in Rhaphadon, Neairea, Gyrocochlea and Coenocharopa. Microsympatry has been observed between Neatrea dorrigoensis and Neairea corticicola with the two species having been collected from under the same piece of bark. These examples are discussed in more detail in the systematic review under the relevant taxonomic heading. Species are largely restricted to rainforest and in some instances e.g. Rhophodon elizahethae and Seromedea nudicostata, are confined to very small patches. In muny cases, panicularly in northern NSW and southern Queensland, clear- ing for agriculture and other human activity has drastically reduced lowland and upland rain- forest. Whether or not species will survive this synanthropic fragmentacion of habitatremains to be seen, Charopids are useful indicator species of bush quality. Arcas where diversity ts high, or where restricted species. reside have special refugial significance, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE a TABLE 1. Mean shell measurements in 49 species of east Australian Charapidae MINIMUM FIRST ~— MEDIAN THIRD = MAXIMUM QUARTILE QUARTILE WHORL COUNT 31/2- 41/8 4 1/2- 4 W4- 6 1/R+ SHELL HEIGHT (MM) 0.49 14 171 2.94 6.01 H/D RATIO 0.40 0.52 0.56 0,60) 0.81 SHELL DIAMETER (MM) 1.12 2.15 3.08 5.35 7.91 D/U RATIO 232 2.68 3,58 B15 closed RIBSON BODY WHORL’ =. 26.8 su 80.0 118.3 279.7 RIBS/MM' 1.96 55 7.23 11.26 33.34 BODY WHORL WIDTH (MM) 0.23 0.7 0.99 1.79 3.50 ' Fourteen species with reduced and irregular ribbing were omitted from these figures. PATTERNS OF VARIATION SHELL Size and shape — Whorl count (Table 1) is usually (33 species) 4-5; four species have <4 whorls while 12 species have 5 or more whorls. Higher whorl numbers occur in several different lineages. Hedleyoconcha delta has the largest number of whorls with 6 1/8+ average; Rotacharopa annabelli has 5 3/4- whorl average (Figs 23a, 140a); Biomphalapa recava and Setomedea monteithi have mean whorl counts of 5 1/4-: Ngairea murphyi has mean of 5 1/2-, Discocharopa aperta with 3 1/2- whorls has the lowest whort count (Fig. 93a), while Sinplovea intensa with 3 5/8+ whorl average (Fig. 137a), is near the minjmum mean whorl count for Pacific Basin Sinployea (Solem, 1983, p. 83). The highly modified Letamola contartus (Fig. 67a—c) has a mean whorl count of 3 7/8-. Species with low whorl counts tend to be small in size, but the opposite does not apply to species with high whorl counts. Of those species with average whorl count >5, six are <3 mm in diameter. Only Hedleyoconcha delta, H. ailaketoae, and Ngairea murphyi have mean diameters which fall into the upper quartile tange. Significantly these three species have strongly keeled body whorls. Hence whorl ad- dition does not appear to have been a major factor in attaining large size. The largest species is Gyrocochlea vinilincta while Discocharapa aperta is the smallest. Be- tween these extremes of shell diameter a sig- nificant number of species fall into the Ist to 2nd quartile range which covers about 27% of the total range of variation (Table 1). In contrast, the larger species show a much greater size range, suggesting that repeated evolutionary experi- ments in size increase occurred. This conclusion is further supported by the many different shell morphotypes represented among larger species, Nine of the 15 largest species belong to genera which have an incomplete secondary ureter and elevated, keeled shells. Only Lenwebbia protescrobiculata, and Setomedea, which show several secondary conchological and anatomical specialisations, have incomplete secondary ureters and more average-sized shells. Other large species belong to Gyracochlea, Nautiliropa, and Elsothera. Conservatism in H/D ratios (Table 1) em- phasises stability of shell shape. Prominently high spired species, Ngairea levicostata (Fig. 6c). Mussonula verax (Fig, 18c), Hedleyaconcha delta (Fig, 23c) and H. ailaketoae (Fig, 6c) have an above average H/D ratio that is >0.70. High H/D ratio in Egilomen globosa is due to greater spire protrusion, closed umbilicus and sharp deflection of the body whorl (Figs 13Ja-c), whereas high whorl count, narrow umbilicus and more protruded spire produce an above average H/D ratio in Rotacharopa densilamellata (Figs 146a-c). In Neairea and Mussonula, spire protrusion and large body whorl width con- tribute to increased height of shell, while in " MEMOIRS OF THE QUEENSLAND MUSEUM Hedleyocencha delta, above average whorl count is a further mitigating factor. In contrast the height of Gyrocochlea, which has a strongly depressed spire, is due to a combination of large body whorl and dramatic deflection of the body whorl in the last third of its volution Umbilical width varies from closed in some species of Cralopa and Elsothera (Figs !01b, 104b. 106b, 108b) to widely open in Oyn- phaloropa, Discecharopa and Rhophadon (Figs 71b, 74b, 77b, 78b, 81b, 84h, §7b, 93b). Anopen umbilicus is most common: a narrow umbilicus oecurs mainly in groups with altered whorl con- tour. Hedleyoconcha, Mussonula and Neatrea (except N. murphvi) have very small umbilici and angulate to keeled body whorls. NV. aurphyi has secondarily narrowed whorls and more open umbilicus (Fig. 9b). In Egilomen globosa (Fig. 13lu.b), Rotacharopa deasilamellata (Vig. I4ha,b). R. kessneri (Fig. 143a,b), and R. an- nabellt (Fig. 140a,b) with tightly coiled shells, whorls are globose and expanded, constricting ihe umbilicus with their inner margins, Umbili- cul contour varies from narrow L-shaped, through wider V-shaped to wide saucer-shaped in Rhaphodon kempseyensis (Fig. 78h) and Dis- cocharopa aperta (Pig, 93b). Whorl profiles range from laterally com- pressed in Coenecharopa alata and Rhophodan bairnsdalensis (Figs 117c, 8%c) to strongly keeled in. Nguirea murphy! and Hedlevecancha delta. In Letomola coniortus whorl profile is altered by a supraperipheral sulcus (Fig. 67c). A weak supraperipheral sulcus is also present in Elsothera nautilodea (Pig. 106c), and Coenocharopa macromphala (Vig. 114¢). Sutures are generally weakly impressed al- though Cralopa straudensis has a very deep suture (Fig. 95a), and Gyrocochtlea vinitincta (Fig, 45a) and its congeners inihally have a weakly impressed suture which hecomes deeply furrowed as the last whorl descends, A more unusual development in Neairea cataliculata and Elsothera genihecata is a narrow channel al the sutural area (Fig. 12a). Apertural sinuses are developed in Cralopa and Elsethera. Sculpture - Spiral apical sculpture is prominent in Stmployea intensa, Omphaloropa varicasa, Seiomedea and Mussonula. S. intensa has low broad spiral cords. (Fig. 137d) while O. varicosd has the cords narrower and slightly higher (Fig. I3de), In Setomedea sencosiata and S, janae they ure reeularly spaced and similar to those of O. varicasa, whereas in S, monteitht (Fig. 41e). 5. nudicostata (Fig. 35e), and Setomedea sp. (Fig. 4c), the cords become squiggly, and vague curved radial ridges develop at the nuclear- postnuclear boundary. Radial clements on the latter part of the protoconch are more pronounced in Mussonula veray Iredale, 1937 (Fig. 18d) and MW. fallax (Fig. 20f). In Gyrocochlea paucilamellata apical sculpture is finely reticulate with spiral and radial elements equally prominent (Fig. Sle) whereas in 6. vrartracta (Fig. 430) and G. conveluta (Fig. 48e) the radials are low weak ridges which are raised at their intersection with the spiral cords to form tiny beads, In contrast, G. currisiana (Fig. 64a) has apical sculpture which initially is a Series of regularly arranged "pits" then becoming ir regularly, radially ribbed toward the end of the protoconch. Biomphalopa recava (Fig. 57e¢), B. concinna (Pig, 60e) and Nautiliropa omicron (Fig. 63d) have similar apieal pitting. A dif- ferent pattern of apical pits in Hedleveconcha delia has deeper and more crowded depressions (Pig, 23), Even in this case, the vague radial pattern of pits indicates probable derivation from a typical reticulate pattern. Leawebbia protoscrobiculate has a similar but finer pattern (Fig. 292). In Ngairea murphyt irregular radial wrinkles and shallow irregularly shaped pits combine (Fig, 9f); apparently a modification of radial ribs and spiral grooves seen in N. cor- licteala (Cox, 1866) and N, levicostata (Figs 6d, 15e). Leromoala contartus, has apical sculpture like that of N. merphyr but with an underlying radial pattern which is less conspicuous, produc- ing w matleate surlace (Pig. 67d), Spirul grooves on the protoconch are rare oc- curring in Neairea corticicola (Fig, 13e), N. canaliculata (Pig, 121), and to a lesser extent NV, levicostata (Fig. 6d). N. dorrigeensis has predominantly radial apical sculpture with very hg spiral lines visible in (he sutural area (Pig. Su). Apical sculpture most commonly consists af prominent radial ribs and less prominent spiral cords. Radials may be crowded (£/sothera, Cralopa, Diseacharopa), widely spaced (Egilomen) or widely spaced initially, becoming more crowded toward the nuclear-postnuclear junction (Ritophoden, Cocnecharopa), Spirals may be continuous, narrow, squiggly cords (Cralopa, Elsothera, Coenocharopa), short, wrinkle-like cordlets (Rhoplodon, Dis- cocharopy), or inconspicuous (Egtlomen). the cords are contifuaus they are usually visible on the apices of the radial ribs. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 7 Apical sculpture af Relacharepa appears reticulate under the light microscope, bur highly complex under the SEM. The radials are a series of curved ridges which become broader toward the nuclear-postnuclear sculpture (Fig. 146d). Spirals, while appeating continuous, are a series of short segments (Figs 140c, 146c). This apical sculpture was also reported in Microcharopa mimula Solem (1983, p. 70). Postnuclear sculpture involves variations on the pattern of major radial ribs and microsculp- tire of microradials and low spiral cords. Major departures are seen in species wilh an incomplete secondary urcter. Pcriostracal enhancements may he present and take the form of high blades on the major radials as seen in Nauziliropa micron (Fig, 63e) or the expanded peripheral extensions of Coenocharopa alata (Pig. 117e,h), A more significant change takes place in Setomedea which has periostracal setae along the length of the major ribs (Figs 32f, 38f, 41f). Major radial ribs may be broad and high as in Biomphalopa recava (Fig. 570, Gyrocochlea vinitincta (Fig. 450), Egilomen cachlidium (Fig. 1282) and Rhophodon peregrinus (Fig, 71f), have a high periostracal blade us in Nautiliropa emicron, Ompltalorepa variceasa, and Rotacharopa (Figs 63e, 1341. 140f), or be reduced with a low periostracal blade expanded aj the shell periphery as in Coenocharapa alata (Fig. 117g), and C. parvicostata (Fig. 120d). In Mussonula verax (Fig. 18¢), M. fallax (Fig. 20g), and Nyairea Jevicostata (Fig. be), the periostra- cal blade is high and continuous above a low radial tib, However, the blade is not vertical as in O. yaricosa, but instead is folded back toward the shell apex. In Setomedea the major ribs have two periostrical blades (Figs 32, 38f). Rib height reduction is scen in Lenwebbia protoscrabiculata which has very low radial un- dulations on adult whorls (Fig. 29f), Ngairea corticicola which has widely spaced low, radiul undulations with a deciduous periostracal blade (Fig. 15f), Letomela contortus which has an overlapping shingle-like arrangement of radial thickenings (Fig. 67¢), and Coenocharopa yes- subahensis which has very low, crowded radial tibs (Fig. 123g). Negairea murphyi (Fig. 9g), N. derrigoensis (Fig, 3¢), Hedleyoconcha delta (Fig. 23f) and H, ailaketoae (Hig. 26d) have crowded, reduced, and thread-like major ribs to ihe exclusion of microradials. Most species lave regular radial ribbing. An average pattern is shown by Siamplialopa recava (Fig. 57a) unl Setmedea xereastara (Fig. 32d). Reduction and ierease in rih counts vecur in hath small and large specyys of several genera, Widely spaced ribs of Rhophordan balrasdalensts (Fig. 89a), Egtlomen eochlidian (Figs 128a.c) and Gyracechlea paucilamellata (Fig, 51a), crowded ribs of Biomphalopa concin- na (Fig. 00a.d), Rhephodon minutissimus (Fig, 84a), und Rotucharopa kessneri (Fig. 143a), Gyrocochlea currisiana (Fig. 54a), Elsethera nautilodea (Fig, 1064) and Retacharopa den- silamellara (Fig. 146a), and the exceptional situation in Riophodon kempsevensis which his a rib count of 279.7 (Fig. 77a), are examples of the variation encountered. Species af Coenacharopa (with the exception of C. yes- s@bafrensis) haye reduced rib counts. However, the deciduous, periostracal nature of the tibs, combined with the worn stale of most of the available material, made rih counts in these species difficult. Reduction in primary radial sculpture may therefore involve reduction in rib counts throughs increased rib spacing, as well as a reduction in the height of the major ribs, Microsculpture typically consists of high crowded radial riblets and low. narrow crowded spiral cords buttressing the radials un either side e.g. Elsothera sericatula (Fig. 104f), Gyrecochlea vinilincta (Fig. 451), Biomphalepa concinna (Fig. 60f) and Cralopa stroudensis (Fig. 9Se). Rhaphodon and Rotacharopa have high microradials bul very low broad spiral cords with small clongate beads formed at their inter- section (Pigs 71f, 89e, 143f, 1461); Rhophodon kempsevensis (Fig. 77e), and Discocharapa aperta (Pig. 930) have the microradials divided into a series of shart segments; Sinployea intensa (Iredale, 1941) has the microradials more thread- like and spiral cords conspicuous with elongate beads at the mmtersections (Fig. 137e); and Egilamen cocliidium (Fig. 128¢) and E. globosa (Fig. 131f), have spirals more conspicuous thas microradials, This latter condition is taken a step further in Omphaloropa varicosa where microradials are very low and microspirals form large beads at the intersection such that intersti- tial areas. appear to be sculptured with spirally arranged beads (Miz 134f), Reduction in the height of mieroradials also oceurs in Coenocharope, Wowever, the microradial ele~ ment attains height by the addition of fine thread~ like periostracal blades e.g. C. alata (Fig. 117g). Microradials may be higher, equal in heighe or lower than (he microspiral cords, but (here are me beads formed and there is no buttressing. 8 MEMOIRS OF THE QUEENSLAND MUSEUM More complex departures from the above pat- terns of microsculpture are seen in species with an incomplete secondary ureter. Setomedea has conspicuous microradials which are higher than the thread-like microspirals, and although the microspirals continue up the sides of the microradials there is no buttressing (Figs 32f, 35f). S. janae has microradials with a scalloped profile (Fig. 38g) as a result of the microsculp- ture dipping into spiral furrows in the shell sur- face. These undulations are also developed in S. monteithi and Setomedea sp. Lenwebbia protoscrobiculata has only microspiral grooves on the early part of adult whorls, but a combina- tion of fine thread-like microradials and grooves on the latter part of the body whorl (Fig. 29e,f). Spiral grooves are present on shells of Ngairea corticicola (Fig. 15f) and N. canaliculata (Fig. 12g). On the other hand N. levicostata has microsculpture of fine spiral lines (Fig. 6e) sim- liar to Mussonula fallax (Fig. 20g). M. verax appears to lack the fine spiral lines of M. fallax, but this needs to be confirmed. Ngairea dor- rigoensis (Fig. 3e) and N. murphyi (Fig. 9a,e) lack microradials, but have many crowded, low, thread-like radials crossed by crowded thread- like microspirals. Low reticulate sculpture also occurs in Hedleyoconcha delta (Fig. 23f) and H. ailaketoae (Fig. 26d), where spiral cords and radials are fused. Hedleyoconcha exhibits an unusual growth-related change in sculpture; after about 4 1/2 whorls the reticulate pattern is replaced by a much less sculptured pattern of irregularly spaced radial growth ridges (Figs 23a, 26a). Apertural Barriers - Apertural barriers were recorded in Letomola contortus and Rhophodon. However, barrier formation is even more widespread among east-coast subtropical charopids. Several undescribed taxa with bar- riers live between Gympie and Rockhampton; until these are described, discussions on barrier formation will remain, skeletal. The form and surface characteristics of bar- tiers in Letomola contortus separate them from those in Rhophodon. Without exception Rhophodon spp. have numerous parietal and palatal barriers whose overall effect is to reduce the apertural area. In contrast L. contortus has only asingle, thin blade-like parietal barrier (Fig. 68a), no barriers on the outer lip margin, and a low ridge-like basal palatal (Fig. 68b). Further- more, the barrier surface in L. contortus has small rounded protrusions (Fig. 68c) whereas Rhophodon has narrow to broad transverse ridges angling up from the barrier surface (Figs 78c, 81d, 84d). R. kempseyensis (Fig. 78c), and R. minutissimus (Fig. 84d), display extremes of variation. These basic differences indicate separate derivation of the barriers in the two genera. In Rhophodon the number, size and shape of barriers as well as surface sculpture are variable. Generally the parietal barriers are more blade- like in form than the shorter crescent-shaped palatal barriers, The number of parietal barriers is least in R. bairnsdalensis which has a single elongate barrier (Fig. 90a), and ranges to the four barriers and two traces present in R, peregrinus (Fig. 71g). Typically there are three blade-like parictals - a horizontal lamella about halfway down the whorl margin, and two on the lower part of the parietal wall, deflected downwards. R. elizabethae has only two parietal lamellae but has a well-developed, low, blade-like superior parietal trace. In R. peregrinus the first parietal is deflected upwards while the second, third and fourth are directed downwards (Fig. 71g). An accessory parietal trace is also present in R. consobrinus (Fig. 74h). Usually the parietal bar- riers have a long anterior taper projecting beyond the edge of the aperture and an expanded posterior section with an abruptly tapered resorption edge. R. peregrinus has recessed parietals which have shortened or no anterior tapers (Fig. 71g). A similar shortening is seen in the deeply recessed second parietal of R. minutis- simus (Fig. 84g). The only other notable varia- tion is in R. colmani which has the parietals greatly swollen, particularly at the posterior edge, thus further reducing the apertural opening (Fig. 871). Columellar barriers range from well deve- loped, horizontal lamellae in R. elizabethae and R. colmani (Fig. 87h) to fine, almost thread-like in R. peregrinus and R, consobrinus. Palatal barriers vary from two in R. bairnsdalensis (Fig. 90b,c) to six in R. colmani (Fig. 87g). A superior palatal trace, situated very near to the palato-parietal margin, may also be present. The first palatal barrier is usually situated at the baso-columellar margin and is a short, high. crescent-shaped lamella. The shape in R. colmani is typical. Remaining palatal bar- tiers are similar in shape though somewhat reduced in height. In R. peregrinus the upper palatals have a slightly elongate anterior taper (Fig. 71h), while those in R. colmani have the anterior edge much more markedly tapered. The low thread-like upper palatal of R. bairnsdalen- sis (Fig. 90c) is a further variation. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 9 ANATOMY 44 of SI) species were dissected, Species nol dissected are Mussonela verax, Hedleyocoucha additu, Rhaphoden colmant, Discocharopa aperta, Elsothera nautilodea, and Coeno- charopa multiradiata. Available material of Mussonula fallax and Neairea levicostata had immature reproductive systems which provided iio details on genital structures. Poor preserva- tion led to obtaining only fragmentary detail of the penis interior of Sinployea intensa and pallial cavity of Coenocharopa parvicostata. Variations in the pallial and genital anatomy are moran reflect regional endemicity, With the exception of Rotacharopa and the predominantly southern Elsottera the patterns are conservative. Pallial Cavity - Typically the pallial cavity has a well-developed mantle collar with mantle gland extending onto the pallial roof, a weakly bilobed kidney in which the pericardial lobe is vaguely triangular and much longer than the rectal lobe, and a complete secondary ureter with ihe ureteric pore situated at the inner edge of the mantle collar adjacent to the anus.. The hindvein 18 conspicuous, bul lacks prominent branching. A major departure from this pattern occurs in Neairea, Hedleyoconcha, Lenwebbia, Mus- sonula, and Selomedéa which have an incom- plete secondary ureter thal is a very short reflexed tube with the ureteric pore situated at the rear of the pallial cavity in the angle between the kidney and the hindgut (Figs 7, (Oe, 21, 24g, 30e, 33g). Whether or not these taxa should be given separate subfamilial recognition ts uncer- tain and will remain so until more of the southern laxa are dissected, In most species the primary ureter isa simple ube running from the kidney apex to the rear of the pallial cavity. However, some taxa with an incomplete secondary ureter have undergone clongation of the primary ureter. This is most ubvious in Se¢amedea which has the initial part of the primary ureter coiled (Figs 33g. 360, 39f, 42f), and in Lenwebbia which has it con- spicuously reflexed (Fig. 30c), Less noticeable are the subUle variations seen in Ngairea and Mussonula (Figs 4e, 13e, 16f, 21) where some degree of reflexion also occurs. The kidney is generally weakly bilobed with a elongate pericardial lobe. In Neairea mur- phyi (Fig. We) and some species of Rhaphaden (Fiys 27f, 3fe} the rectal lobe is almost entirely absent and the pericardial lobe is markedly elon- gate, All these species have above average whorl counts, Bilobed kidneys with almost equal sized lobes are present in Egilomen (Figs 129d. 1321), Sinployea jinensa (Pig, 138f) and Lefomaly contortus (Fig. 69g). Species of Cralopa (Figs 960, 99¢, 102c) have the rectal lobe about 1/3 the length of (he pericardial Jobe. A mantle gland ts usually present although na gland was noticed in fedl/eyoconcha, Sinployvea and some species of Gyrocectilea, Nautilirapa, Elsotiera and Ratacharopa. GENITAL ANATOMY Ovetestis - Usually the ovotestis consists of two clumps of pulmutely clavate Lo finger-like jobes of alveoli embedded in the apical whurls of the digestive gland and oriented parallel to the plane of coiling, In species with an clevated spire such as those belonging to Ngatrea, Hed- leyoconcha and Lenwebbia the \obes are pusi- tioned at right angles.to the plane of coiling (Figs 5e. 10g, 24e, 27d, 30c). Gyrocochlea vinitincta which has a greatly enlarged shell with expanded whorls, also has the ovotestis more at right angles than parallel (Fig.46c). Although two clumps of alveoli are normally present, some proliferation of clumps takes place in the high spired and high whorled Nguiren suuirphst (Fig. Wd), and Hedleveconcha delta (Fig. 24¢). Single clumps. are present in the tiny Siaplovea intensa (Pig, 138e) and Letomola can- fortus (Fig. 69%c) while according to Solem (1983, p, 74) a single clump is present in the minute Diseocharopa aperta.. Within a clump af the ovotesuis the number of alveolar lobes varies from many in some of the larger species to the simple bilobed condition in the very small Ont phaloropa varicosa (Fig. 1350), Rliephacdar bairnsdalensis (Vig, 91d) and L cantarius (Fig. 69d). In Rotacharopa the ovotestis consists of two teardrop-shaped lobes (Figs l4tc, l44e. 147e) representing a significant difference fram the simple clavate alveoli in the Charopinae. Hermaphroditic Duct - Most species inves- tigated have a hermaphroditic duct which ts a simple tube leading apically from the carrefour region, along the parietal whorl margin to the ovotestis Jocated in the apical whorls of the digestive gland, In several species the tube is partially kinked, This is best illustrated in species ol Medleyoconcha (Figs 240, 27e) whieh have elevated, trochoidal shells, and also in Neairea dorrigoensis and Gyracochlea vinitincla (Figs 4d, 46d), Differences in the We gree of duct expansion, noted in some dissec- Hans. are most probably related to levels of tt MEMOIKS. OF THE QUEENSLAND MUSEUM reproductive activity and have lille phyletc relevance. The colour of the duct varies from Indescent pink to jridescent green, Talo and Carrefour - In a number of cases poor preservation made interpretation of this region difficult. Typically the talon is a circular to subcircular swelling situated atop a short stout stalk although in some species e.g. Hed- leyoeoneha delta, ie talon is finger-like (Fig. 27c), Flongation of the talon stalk occurs im Cyrocvelilea curtisiana (Fig. 55e), Rhophedan bairnsdalensis (Fig. 91e) and Rotacharopa den- sllamellata (Fig. 147d), while in Sinployea in- tensa and Egilomen cachlidium the stalk is extremely short (Figs 138c, 129e). The car- refour is typically an expansion in the talon stalk al its junction with the hermaphrodilic duct, Albumen Gland - The size of the albumen vland is variable and probably reflects the level of reproductive activity, Shape of the gland is uffected lurgely by the degree of indentation by the spermathecal head and intestinal loops. Prastate and Uterus - Nospecies showed any departure from the typical charopid patiern of (used prostate-ulerus, The prostatic alveoli are situated cither along the full length of the uterine chambers or compacted into a mass at the base of the albumen gland (Fig. [ 15a). The uterus consists of 4 lower chamber with thick walls thrown into folds and a thinner, glan- dijlar-walled upper chamber, In some cases this differentiation was extemally conspicuous while inathers, dissection was necessary to determine the areas of differentiation. Terminal Male Genitalia - Differences in this region include relative size and shape variation in the vas deferens, epiphallus and penis; shilts in the insertion of the penial retractor Muscle, modification of the vas deferens-epiphallis Junction; positional variation in the cpiphallus- penis junction; addition of structures; and major changes in the penis interior particularly under conditions of sympatry, Because of differences id preservation states between individual specimens, size and shape variations were not quantitied, The penial retractor muscle is generally a short tull originating on the diaphragm and inserting or the epiphallus near or at the epiphallus-penis junction, * The muscle is clongate in Biom- phalopa recava and Rhophndon cansabrinus bul differential contraction of individual specimens did not enuble full investigation of length uif- ferences, Major departures fromm this basic pat- tern include the shift of the insertion point onto the pems head or to a point well along the cpiphallus. The former condition is usually as- sociated with a shift of the epiphallic entrance to a sub-apical position on the penis e.g. Ngairea murphyi (Fig. |\0b), Rhophaden consohrinus (Fig. 75c). Gyrocochilea and Biomphalopa (Figs 46b, d9b, 52b, 55a, SKb, 61b), In Egilomen globosa (Fig, 132b), and Cralapa kaputarensis (Fig, 99b), the penial retractor muscle inserts c. 1/3 of the way along the epiphallus while in Seromedea nndicosiata (Fig, 36b), and Elsothera genniecata (Fig. 109a), the insertion point isc. 1/2 way along the epiphallus. In Roracharopa, the epiphallus is ditferentisted into ascending and descending arms and the retractor muscle is inserted cither at the point of reflexion (Figs l44a. 147a), or al the vas deferens-epiphallus junction (Fig. 1414). This latter shift in position may be related to the effects of increased whorl count. An unusual condition in Seromedea manterthr (Fig. 42a), and Gyracochlea curtisiane (Fig. 553i) has retractor muscle and terminal part of epiphallus intertwined. The vas deferens is typically a thit tube des- cending from the prostale-wlerus to the penieviducal angle before reflexing and giving rise to a more expanded epiphallus c. 1/2-1/3 of the way along the penis length, Exceptions are Gyrocochlea which has the vas deferens ex- pinded initially before becoming more siender prior To joining the epiphallus (Figs 46a, 49a, 55a}, and Cralepa which has the vas deferens expanded for its entire length (Figs 96a, 99a, LU2a). In most species the vas deferens enters the epiphallus apically through a simple pore, huwever in B/sothera sericatula (Fig. }05b) and F. gentthecala (Vig. 1094), the vas deferens enters the epiphallus laterally through a large cup-shaped pilaster. Externally this complex junction is a conspicuus swelling. The epiphallus is usually differentiated from the vas deferens as an abruptly expanded. mus- culur jube, generally shorter than the penis, in- ternally with longitudinal pilasters, and reflexed before entering the penis apically, Departures from the general patiern include Gyracocklea which has an epiphallus that is as Jong as the penis (Figs 46a, 49a, 52b, 55a), and Neairea murplivl (Fig Wa), Rhophodan kempseyensis (Fig 7¥a). Elsaihera genithecata (Fig. 109a), Nyatrea canalieulua (Vig, 13a), Seromedea nudicastata (Figs 36a.b). and Coenochurapa vwordidus (Fig, 112a), which have an cpiphaflus thar is much longer than the penis, In. SYSTEMATIOS OF AUSTRALIAN CHAROPIDAE il kempseyensis and N. murpliryi the elongate epiphallus may be a secandary modification re- lated to whorl count increment; in C. serdidus the change to Jonger epiphallus occurs under conditions of complex congencric sympatry and may be associated with species recognition; in E. genitfrecata the enlarged epiphallus correlates with overall size increase. Letomola cantortus, which has reduced whorl count, has elongate terminal genitalia with long slender cpiphallus and penis of equivalent length (Fig, 49a), However, the bulbous protoconch indicates descent from a larger ancestor and elongation in this case can be jnterpreted as secondary miniaturisation of a larger genital sys- lem. The epiphallus undergoes minor structural modification in Seromedea seticastata which has the terminal part of the epiphallus and apical section of the penis bound in a connective sheath (Fig. 33a.b), and §, janae which has the epiphal- Jus incorporated into the penial apex (Fig. 39a,c). Gyrocochlea, Coenocharopa macromphala (Fig. }15a), and C. alate (Fig. 118b), have the lerminal portion of the epiphullus bound to the penis sheath with thin strands of connective tis- sue, In Elsothera genithecata thick muscular sheath covers the terminal half of the epiphallus and the penial retractor muscle is inserted on the sheath (Fig. W¥a), In contrast to the minor adjustments seen in Seinmedea and Coenocharopa the epiphallic shealh of &. genithecaia represents a major additive strue- ture, A more significant deviation from the above patterns is seen in Retacharopa where the epiphallus hus ascending and descending arms of roughly equal length but with the latter greatly expanded, The penial retractor muscle is in- serted either at the point of reflexion or al the vas deferens-epiphallus junction (Figs 141a, 144a, \47a), This external change correlates with a significant change In Internal structure. Typical- ly the interior walls of the epiphallus are sculp- tured with simple longitudinal thickenings whereas in Rotacharopa there are large Irunsverse thickenings (Figs 141b, 144b), Entry of the epiphallus into the penis is usually apical, through a simple pore surrounded by a circular fleshy thickening or two fleshy lips. In (iyrocochlea, Biomphalupa and some Setomedea, the entry point is ina subapical posi- tion, with the penial retractor musele inserted on the penis head (Pigs 33e, 49b, 52b, 58b). These positional shifts are associated with elongation of the penis (Gyrocochlea, Biomphalapa) or change in epiphallus structure (Serontedea) and are tm contrast ta the positional shifts related to increased whorl count in Neu/rea murphyi (Fig 10b) and species recoynition in Rhophodon con- sobrinus (Fig. 75¢). Conical verges are developed in Gyrocechlea vinitineta (Fig. 46b) and Ngairea canaliculata (Fig. 13b). most prohably in response tn congencric sympaters. A tubular verge is seen in Omphaloropa varicosa (Fig. 1356). The penis is generally a short, muscular tube which may have an apical section expanded in response to complex internal pilasters or incor poration of the epiphallus, Exceptions include the elongate, sometimes reflexed, cylindrical penis of Greocochlea (Figs 460.6, 44a,b), ex- panded. barrel-shaped penis of Cralopa (Figs 96a, 99a, IN2ajy, and tubular penis of Coenocharapa (Figs 15a, | 18a, 24a). A more drastic change is seen in the penis of Roracharapa which has the typical apical bulb of other charopids and an unusual basal exten sion, ‘This basal extension, or preputial (ube ws iv is here named, is mast noticeable in R. den- stlamellaia (Fig, 1474). where it is almost twice the length of the penis proper. Function of this structure is unknown. A penis sheath is present in Gyrocochlea, Coenocharapa and some species of Rhophodon. The penis interior is variable between genera; and wilhin genera under conditions of cun- generic sympatry. However, there is generally an apical epiphallic pore and basal longitudinal pilasters, sometimes with a muscular apical cal- lar separating the penial bulb from the lower tubular section. In Ngajrea i muscular collar below the epiphalli¢ entrance effectively divides the penis into two chambers (Figs 4b, 10b, [3b, 166). while Hedlevoconcha has w grooved muscular thickening constricting the passage between the epiphallic pore and the basal portion of the penis (Figs 24c, 27b), The shift in epiphallic entrance (o a subapical position in Seramedea seticastata (Fig, 33c), S. mameithi (Fig. 42c), and Rhophedon consobrinus (Fig. 75e) has resulted in the developiment of an apical penial chamber which has developed ils own pilaster structure, most likely for species recognition. In contrast Neairea murphyi which also has a subapical epiphatlic entrance has an apical penial chamber without pronounced sculpture (Fig. LOb). Natutiliropa omicran which has a penis with an apical bulb separated from the main chamber by a thin muscular collar (Fig. 65c) also has the epiphallic entrance apical, though slightly lateral, with the penial retractor muscle inserted on the penial bulb. In addition to the epiphallic entrance the penial bulb contains a horseshoe- shaped pilaster. This pattern is broadly equivalent to that seen in S. seticostata and R. consobrinus. Elsothera sericatula (Fig. 105e) and E. genithecata (Fig. 109b), have penes which are also internally divided by a muscular collar. The epiphallic entrance is apical and the region be- tween the collar and the epiphallic pore is large and sculptured with short fleshy thickenings. This change to two quite differently sculptured penial chambers in Elsothera is a major struc- tural change and contrasts with some of the species recognition changes mentioned pre- viously. In Gyrocochlea, the long, cylindrical penis is internally differentiated into a lower portion with large, longitudinal pilasters and an area apicad of the simple epiphallic pore which has short, almost pustular, thickenings (Figs 49a, 52a). The exception is G. vinitincta with a conical verge (Fig. 46a). Biomphalopa recava has a single large longitudinal pilaster and short lon- gitudinal thickenings apicad of the epiphallic pore (Fig. 58b) but B. concinna lacks the apical thickenings and has a well-developed spatulate verge (Fig. 61b). Verges are also present in Setomedea nudicostata, Omphaloropa varicosa, and Rotacharopa kessneri (Figs 36b, 144b). The penis is typically sculptured with inter- generically-variable longitudinal pilasters which show marked intrageneric differences under conditions of sympatry. These differences may be subtle minor shifts in size and shape of pilasters as in Gyrocochlea, Ngairea and Setomedea, or may involve more drastic altera- tion of pilaster patterns as seen in Rhophodon, Coenocharopa, Egilomen and Rotacharopa. Gyrocochlea vinitincta and G. paucilamellata, are sympatric in areas of the Lamington National Park in the eastern Border Ranges, SEQ. G. paucilamellata retains the typical generic pat- tern of numerous anastomosing longitudinal pilasters (Fig. 52b) while G. vinitincta has an altered condition in which the pilasters are simple and unbranched (Fig. 46b). This change may seem minor but, together with the conical verge of G. vinitincta, would serve as an effec- tive species isolating mechanism. The microsympatric Ngairea corticicola and N. dor- rigoensis have numerous longitudinal pilasters, 2 MEMOIRS OF THE QUEENSLAND MUSEUM but these are strongly corrugated in N. cor- ticicola (Fig. 16b) and simple in N. dorrigoensis. In Setomedea monteithi, which is probably sym- patric with S. sp., short longitudinal pilasters are replaced by two short pad-like pilasters (Fig. 33c). More dramatic pilaster pattern alteration is seen in Coenocharopa sordidus and C. yes- sabahensis. Sympatry in Coenocharopa is high- ly complex with two to three species involved per locality. In C, yessabahensis an apical pad and large semi-circular basal pilaster (Fig. 124b) replace the more typical longitudinal pattern (Figs 115d, 118b), while the horseshoe-shaped pilasters of C. sordidus represent further varia- tion (Fig. 112a). In Rhophodon pilasters are typically fusiform longitudinal. R. peregrinus has a penis with an apical verge-like thickening and basal circular pilaster (Fig. 72b), while the sympatric R. con- sobrinus has longitudinal pilasters in the penis proper, a basal circular pilaster and shift in epiphallic entry (Fig. 75e). The two species of Egilomen are sympatric and have quite contrast- ing patterns of penial surface sculpture (Figs 129b, 132b). In the three species of Rotadiscinae the change from typical longitudinal pilasters to the pad-like pilasters of Rotacharopa annabelli (Fig. 141b), is an equivalent change to that seen among members of sympatric species pairs in the Charopinae. However a sympatric partner for this species has yet to be found. Of the monotypic genera, Lenwebbia and Om- phaloropa have simple longitudinal pilasters (Figs 30b, 135b); Letomola has an apical spongy thickening in addition to longitudinal pilasters (Fig. 69b); and the penis of Sinployea intensa was too poorly preserved to accurately detail internal features, though a basal semicircular pilaster (Fig. 138b) relates to the generic pilaster pattern (Solem, 1983). In Nautiliropa omicron the penis has several anastomosing longitudinal pilasters and a large apical pocket stimulator (Fig. 65c). This con- trasts with the conchologically similar Gyrocochlea, and effectively separates Nautiliropa and Gyrocochlea. The complex stimulator is different to the simple circular and pad-like pilasters developed in other genera dis- cussed above and indicates that it probably is not a simple species recognition feature. Terminal Female Genitalia - Observed varia- tion was confined to differences in the compara- tive lengths of the free oviduct and vagina. The SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 13 few species in which the internal wall sculpture was examined showed a repetitive pattern of longitudinal pilasters; no attempt was made at detailed analysis as presented for male genitalia. Length of structures may be affected by contrac- tion during preservation so comparisons are pre- sented qualitatively rather than quantitatively. In general, genera with elongate genitalia have long vaginae e.g. Hedleyoconcha, Gyrocochlea, Letomola and Rhophodon (Figs 24a, 46a, 69a, 72a). However, length changes in the vagina may also be related to species recognition. Rhophodon consobrinus with short vagina (Fig. 75a) in comparison with its congeners, is sym- patric with R. peregrinus and has altered ter- minal male genitalia. Coenocharopa, except C. sordidus (Fig. 112a), has average length vaginae; C. sordidus, which exists under condi- tions of complex congeneric sympatry, has a very short vagina. Short vaginae are also present in Ngairea and Lenwebbia (Fig. 30a). The free oviduct is typically a muscular tube equal in length to, or only slightly longer than, the vagina. Conspicuous exceptions are Ngairea (Figs 4a, 13a), Rhophodon consobrinus (Fig. 75a) and Coenocharopa sordidus (Fig. 112a), which have much longer free oviducts. Atrium - The atrium does not display any un- usual features apart from minor variations in length. The shortened condition in Rhophodon consobrinus (Fig. 75a) and the elongate structure in Coenocharopa sordidus (Fig. 112a) illustrate the extent of variation. Spermatophore - Sperm packets were found in five species. The most unusual spermatophore was that of Setomedea monteithi, which had a long chitinous tail (Fig. 42b). The others con- sisted of a clavate structure in Nautiliropa omicron (Fig. 65d), an arc-shaped, terminally hooked packet in Gyrocochlea paucilamellata (Fig. 52d), and a vaguely U-shaped packet in Rhophodon consobrinus and R. kempseyensis (Figs 75f, 79f). Although few were located, it is probably more than coincidental that the two Rhophodon species have similar looking sper- matophores. RADULA The usual pattern of radular structure is for a tricuspid central tooth, slightly smaller than the adjacent laterals, with long, lanceolate mesocone and short pointed ectocones; tricuspid lateral teeth which are similar in shape to the central; a transition zone of lateromarginal teeth in which the endocone becomes curved and clon- gate, the mesocone becomes shorter and the ec- tocone lengthens slightly; and marginals in which the endocone and mesocone are reduced, and the ectocone, and occasionally the en- docone, are split into a series of minor cusps. Setomedea nudicostata is typical (Fig. 35d,g,h). Variations fall into several categories. Minor differences in the size and shape of teeth occur in Biomphalopa and Gyrocochlea which have a slightly more elongate mesocone on the central and lateral teeth (Figs 45g, 48d, 57d). Elsothera has a large, broadly lanceolate mesocone on the central and lateral teeth (Figs 104g; 108g,h); Egilomen and Cralopa have the central tooth c. 1/2 the size of the laterals (Figs 98f, 101f, 128h, 131h); Omphaloropa has a small central tooth and reduced number of teeth per row (Fig. 134d,h): Rotacharopa and Rhophodon have slender, lanceolate mesocones on the lateral and central teeth (Figs 146g, 190d). In contrast to the average generic pattern Rhophodon kempseyen- sis has the mesocone of the central and laterals greatly expanded (Fig. 78e). This condition is also seen in Coenocharopa yessabahensis (Figs 123d.h). Significantly both R. kempseyensis and C. yessabahensis live on limestone rocks and the modification in cusp size probably relates to an associated shift in feeding habit. A more dramatic level of change in tooth shape is seen in Letomola contortus (Fig. 68d,e,f) and Nautiliropa amicron (Fig. 63c.f). L. contortus has a highly modified radula with a small central tooth which has the side cusps reduced to tiny knobs. In contrast, the first row of laterals has a greatly enlarged, broadly lanceolate mesocone and small side cusps. Furthermore there are very few lateral teeth, no transitional lateromarginal teeth, and serrate marginals (Fig. 68d). L. con- tortus lives on limestone and the enlarged mesocone on the laterals may be habitat related. However, the remaining modifications are major changes in radular structure which correlate with the many unusual conchological specialisations. In N. omicron the central and inner laterals have a long, slender, curved mesocone and almost obsolete side cusps (Fig. 63f), the outer laterals have the endocone lengthened and curved, and the marginals are unusual in having an elongate basal shaft with two cusps - the mesocone and endocone - in a crab-claw arrangement (Fig. 63c). The teeth are arranged in V-shaped rows as opposed to the straight line arrangement in other genera. These features are significantly different to the typical pattern of other log-dwell- ing species reviewed. 14 MEMOIRS OF THE QUEENSLAND MUSEUM Some unusual radular variations also occur among species with an incomplete secondary ureter. Lenwebbia protoscrobiculata has typical central, lateral and marginal teeth but lacks tran- sitional lateromarginal teeth (Fig. 29d). Hed- leyoconcha delta has a typically tricuspid central tooth, but with a marked flare of the basal plate (Fig. 23g); the flared extension is less obvious in the lateral and marginal teeth. Mussonula fallax (Fig. 20d,e), has a tricuspid central tooth with a broad anterior extension, laterals initially bicuspid, outer laterals and lateromarginals tricuspid with the basal extension further en- larged, and marginals typically multicuspid with an enlarged basal extension. A similar pattern is present in the conchologically convergent Ngairea levicostata, although, the basal exten- sion is more pronounced (Fig. 6f). In NV. murphyi the basal extension is absent in the central tooth but present as a conspicuous outwardly directed flare in the laterals (Fig. 9d,h). Negairea corticicola, N. dorrigoensis and N. canaliculata have radulae with a small, broadly triangular central tooth with a short broad basal extension, and laterals and marginals with a dis- tinctly flared extension (Figs 3f, 12d, 15d,h). These changes to tooth structure in Mussonula, Hedleyoconcha and Ngairea represent a change in interrow support which in Hedleyoconcha and Ngairea correlates with a shift to semiarboreal and arboreal lifestyles. The preferred microhabitat of Mussonula is unknown. SUMMARY It is possible to group the conchological and anatomical patterns outlined above, into levels of evolutionary significance (Solem, 1978). At the lower end of the scale there are changes which are related to species recognition. These include differences in the internal sculpture of the penis, minor positional shifts in terminal reproductive structures, and differences in shell morphology and radular structure associated with small habitat shifts. Examples are given in species accounts. Ata slightly higher level are those changes associated with colonising par- ticular adaptive zones. These include the periostracal setae of the damp habitat Setomedea; the reduced sculpture of the intersti- tial Coenocharopa; the strongly to moderately elevated shells of the semiarboreal and arboreal Hedleyoconcha, Lenwebbia and Ngairea; the strongly ribbed, flattened to biconcave shells of the under-log dwelling Gyrocochlea, Nautiliropa and Biomphalopa; the dentate, sculptured shells of the rock dwelling Rhophodon; and the dentate, secondarily modified shell of rock dwelling Letomola. Cor- responding anatomical changes include altera- tions in penis shape and internal penial structure, addition of major structures to the genitalia, changes in kidney shape, and radical modifica- tions in radular morphology. Above this level are basic changes in sculptural elements and large scale changes in anatomy, such as in Rotacharopa and possibly the group of genera with incomplete secondary ureter. SYSTEMATICS Family CHAROPIDAE Hutton, 1884 Solem (1983, p. 59) presented a detailed diag- nosis that is accepted herein. Subfamily CHAROPINAE Hutton, 1884 A broad definition of the subfamily is presented by Solem (1983, p. 70). The genitalia of the Charopinae reviewed here show a consis- tent pattern of muscular epiphallus with internal longitudinal thickenings, reflexed before enter- ing the penis through a simple pore, or oc- casionally a verge. The penis has an apical bulb containing the epiphallic entrance and a lower basal portion with longitudinal thickenings which may be modified under conditions of sym- patric species interaction. The pallial configura- tion includes a kidney in which the pericardial lobe is usually much longer than the rectal lobe and the secondary ureter is complete or rarely incomplete. Ngairea gen. nov. Oreokera Iredale, 1933 (part), p.54, nom. nud.; Iredale, 1937a (part), p. 318, nom. nud.; Iredale, 1941a, p. 261, nom. nud.; non Stanisic, 1987, p.2. ETYMOLOGY For my youngest daughter Ngaire. TYPE SPECIES Oreokera dorrigoensis Iredale, 1941. PREVIOUS STUDIES Iredale (1933) introduced Oreokera for Flam- mulina cumulus Odhner, 1917 and added Helix corticicola Cox, 1866 (Iredale 1937a) and Ore - SYSTEMATICS OF: AUSTRALIAN CHAROPIDAE 15 okera dorrigoensis Iredale, 1941. Stanisic (1987) validated Oreokera, including only F. cumulus, and one other north Queensland species. Ngairea is introduced to include H. cor- ticicola, O. dorrigoensis, two new species from southern Queensland, and ’Helix’ murphyi Cox, 1864 from southern NSW. The latter was pre- viously included in the punctid form-group Paralaoma Iredale, 1913 by Iredale (1937a, 1941a). DIAGNOSIS Large to very large, diameter 5.46—-7.40 mm, with 4 1/8 to 5 7/8 normally to tightly (murphyi) coiled whorls. Height 2.86—5.35 mm. Apex and spire moderately to strongly (murphyi, levicos- tata) elevated, body whorl descending slightly. Protoconch almost flat, 1 5/8 to 1 3/4 whorls. Apical sculpture of curved radial ribs, reduced (canaliculata) or modified (murphyi), with in- cised spiral grooves (canaliculata, corticicola), weak raised spiral cords (/evicostata) or without spiral elements (dorrigoensis, murphyi). Post nuclear sculpture of crowded, protractively sinuated radial ribs which may be reduced in some species (corticicola), Microsculpture of crowded spiral cords (murphyi, dorrigoensis), incised spiral grooves (corticicola, canaliculata), or fine spiral lines (levicostata). No microriblets present. Sculpture continuous on the base but reduced. Umbilicus narrow to widely U-shaped (murphyi), rarely closed (dor- rigoensis), Sutures impressed, maybe chan- nelled (canaliculata). Whorls weakly angulate to strongly carinate (murphyi), sometimes rounded (canaliculata). Aperture roundly lunate to subquadrate. Lip simple, sometimes slightly thickened; columella vertical, reflected toward umbilicus. Colour yellow-horn, with red flam- mulations or without (murphyi and canaliculata.) Foot and tail broad, bluntly rounded posterior- ly; a caudal horn sometimes present (levicos- tata). Animal creamy-white with orange-grey speckling on the sides of the foot and on the mantle collar. Pallial roof and visceral hump with variable black speckling. Ommatophores black. Kidney broadly to narrowly (smurphyi) triangular; rectal lobe vestigial or absent (mur- phyi). Ureter sigmurethrous with incomplete secondary branch. Ureteric opening at the rear of the pallial cavity in the angle between kidney and hindgut. Genitalia with ovotestis of two or more (murphyi) clumps of palmately clavate lobes of alveoli, separated by digestive gland and oriented at right angles to the plane of coiling. Terminal male genitalia with muscular epiphal- lus, sometimes much longer (canaliculata) than the penis, entering the penis apically, or subapi- cally (murphyi). Epiphallic pore central in mus- cularised pilaster or opening through a conical verge (canaliculata). Penial retractor muscle in- serted on the epiphallus (dorrigoensis), penis- epiphallus junction (corticicola, canaliculata), or on the penis (murphyi). Penis with an apical bulb internally separated from a lower tapered, tubular section by a muscular collar. Internal penial sculpture of broad, corrugated (cor- ticicola, canaliculata) or slender (murphyi, dor- rigoensis) fleshy, longitudinal pilasters. Female reproductive tract without unusual features. Central tooth of radula broad, triangular with tiny ectocones, or slender lanceolate (/evicos- tata, murphyi) with prominent ectocones. DISTRIBUTION AND ECOLOGY Ngairea extends from warm temperate closed forests in the Illawarra region of southern NSW to the subtropical and temperate rainforests of northern NSW, and drier subtropical forests of southern Queensland. With the exception of N. murphyi and N. levicostata species distributions are well known. Most species live under the bark of fallen trees. N. corticicola is sympatric with N. dorrigoen- sis from Dorrigo to the eastern Border Ranges; N, levicostata is sympatric with N. dorrigoensis in the Koreelah State Forest region of the western Border Ranges. N. dorrigoensis and N. corticicola were collected from under the same piece of bark at Upper Pine Creek, Canungra, SEQ. In N. dorrigoensis and N. levicostata dif- ferences in shell shape and radula indicate diver- gent microhabitat preferences. PATTERNS OF SHELL VARIATION Size and Shape - Adult Ngairea are some of the largest Australian subtropical charopids with mean shell diameter 5.80-6.90 mm. Average whorl counts range from 4 1/2- to 5 1/2-. N. murphyi has the highest mean whorl count (5 1/2-), smallest mean diameter, but has a smaller whorl cross section. Reduced height in N. murphyi is the result of smaller shell size but N. /evicostata has a taller shell through spire elevation. In N, murphyi and N. dorrigoensis decreased shell height correlates with keel development on the body whorl. The umbilical opening is smallest in species with a distinctive keel (NV. dorrigoensis and N. 16 MEMOIRS OF THE QUEENSLAND MUSEUM levicostata) although N. murphyi has a secon- darily enlarged umbilicus resulting from reduc- tion in size of whorl cross section. Sculpture - Apical sculpture consists of curved, crowded radial ribs which may be pronounced, reduced or modified into vague radial ridges and wrinkles. Less prominent spiral elements may be present as incised spiral lines or raised cords. In WN. corticicola (Fig. 15e) intersections of radials and spirals form a series of pits. Postapical sculpture is predominantly radial, with ribs varying from bold, broad, regularly spaced (Fig. 12a), and high, crowded (Fig. 6e) to very crowded (Figs 3e, 9g). Various types of spiral sculpture are also present and in N. cor- ticicola (Fig. 15f), incised spiral lines become notched, approximating some helicarionid taxa, e.g. Westracystis Iredale, 1939 and Delinitesta Iredale, 1933 (Solem, 1982; Stanisic, 1988). One of the more interesting developments in the shell of Ngairea is the canalised sutures seen in N. canaliculata, Solem (1983, pp. 9, 108) recorded this feature in the Pacific Island charopid Sinployea proxima (Garrett, 1872). Canalisation is due to detachment of part of the body whorl and in N. canaliculata results in a relatively wide channel. Variation in shell morphology is greater than in other genera included in this study indicating a long, complex period of evolution. N. murphyi, which has developed in isolation from the main group of species exhibits some major departures from average patterns. PATTERNS OF ANATOMICAL VARIATION Variations in anatomy involve shape changes in the pallial structures, size differences in Treproductive organs, species interaction modifications in the terminal male genitalia, and diet related shifts in radular morphology. COMPARISONS Ngairea and Mussonula may be confused and are sympatric in and around the eastern part of the Border Ranges. Mussonula has a strongly exsert protoconch with prominent spiral cords and vague radial ridges prominent toward the nuclear-post nuclear shell boundary (Figs 18d, 20f). Lenwebbia is sympatric with Ngairea be- tween Colosseum Creek and the Bobby Range, SEQ, and bears some similarity. However, it has a strongly punctate protoconch, adult sculpture of spiral grooves and fine thread-like radials (Fig. 29e—g), lacks the internal division of the penis, and has the kidney apex divided by a portion of reflexed ureter (Fig. 30b,e). KEY TO SPECIES OF NGAIREA 1.Shell with prominent radial ribs v.00... 2 Shell without prominent radial ribs; base al- mostsmooth (Fig. 15b) wc. corticicola PZ SUMUPSS SINGS 5 sdepacsicedseconiogysieriseonqvowbeanntacace 3 Sutures channelled (Fig. 12e) .... canaliculata 3.Umbilicus nearly closed to slightly open; peripheral keel obtuse occ eeeeeeseeeeees 4 Umbilicus widely open (Fig. 9b); peripheral keel acute (Fig. 9C) wesc murphyi 4.Microsculpture of prominent spiral cords (Fig. 3e); spire lower, mean H/D ratio 0.56.0... lesplosegsizendceigg Maskvseyavrolgsngtrsevaaves dorrigoensis Microsculpture of weak spiral lines (Fig. 6e); spire higher, mean H/D ratio 0.73.....0..0..00 Ngairea dorrigoensis (Iredale, 1941) comb. nov.(Figs 3-5; Tables 2, 3) Flammulina corticicola (Cox); Hedley, 1912 (part), p. 264, pl. 7, figs 46-48. Oreokera dorrigoensis Iredale, 1941a,p. 265, fig. 5 COMPARISONS N. dorrigoensis differs from N. corticicola in its more prominent radial sculpture, more angu- late periphery, narrower umbilicus and micros- culpture of crowded microspiral thickenings (Fig. 3e).N. levicostata has a more elevated shell and microsculpture of fine spiral lines (Fig. 6e). The apical spiral cords, smaller size and lack of prominent spiral cords on the adult whorls of Mussonula Iredale, 1937 readily separate mem- bers of that genus from N. dorrigoensis. TYPE MATERIAL HOLOTYPE: AMC63517, Dorrigo Scrub, NSW., Height of shell 3.45 mm, diameter 6.23 mm, H/D ratio 0.55, D/U ratio 14.49, whorls 4 1/2-. PARATYPE:AMC635 13, same locality as holotype. VAULE DRANGE DF VARIATION IN NGAIREAMUSSONUTA, LENWEDDTA AND TEDEBYOCONCHA (MEAN AND RANGE) DU RATIO UMBILICAL wibnt HLS, 1UL5/MM SY/WwW RATIO WIDTH nobyY WHORL (mm) SPIRE HEIGHT DIAMETER HD RATIO PROTRUSION (nvm) WHORLS NUMBER OF SPECIMENS NAME Gm) (rom) (mm) SYSTEMATICS OF AUSTRALIAN CHAROPIDAE (126-2199) 1291 1459 (124 P3905) = iz = a a) a se (25-204) 227 (030-066) 1973 SFScsrsrs a 5 ae ae ae (424 aay Pinte (430-5 1/2) AMS protevceotieutare Hedlepornnchal canalicular darriyaensiy Newrra Neairen carfivicela anlaketoae “Tevidwstdia Mucrsurrictar Np rrurphye Neairva verax Mics délia Nedleyncunchy aire fatlas Lenwebhun Negarea OTHER MATERIAL. NEW SOUTH WALES: Platypus Creek, New England N.P., (1 AMC142440, 27 Feb 1961, D-F. McMichael); Acacia Plateau Rd, c. 12.2 km from Acacia Ck ~ Killarney Road, SEVT/MVF (28°21'S, 152°24°E) (1, AMC128560, 15 Mar 1981, J. Stanisic, D. Potter); Beaury §.F., c. 15 km W of Urbenville, 810 m (28°27°S, 152°24°E) (3, AMC142475, 18 May 1976, P.H. Colman, I. Lach): Marengo S.F.,.c. 40 km NW of Dorrigo, 1020 m (30°06'S, 152°25°E) (5, AMC 142476, 10 Feb 1976, J.B. Burch, P-H. Calman); SE of Tabulam, tributary of Busby’s Creek (29°02'S, 152°43°E) (2, AMC137757, 29 Aug 1982, ABRS - AM/QM); The Glade, Dorrigo N.P., 600 m, temperate rainforest (30°22'20"S, 152°43'°40"E) (i, QMMO16440, 12 Nov 1983, ABRS - AM/QM):; Tooloom Scrub, Beaury S.F., CNVF (28°29'S, 152°24’E) (2, OMMO10939. 15 Mar 1981, ABRS - AM/QM);: Dorrigo (6, AMC142443); Dorrigo, under bark and logs (1, AMC63788, Dee 1910, S.W. Jack- son); Darrigo (1, AMC 142444, ex Helms); Moonpar S.F., Dorriga (1, AMC142441, 4 Apr 1960, D.K, McAlpine); Booyang (= Booyong?) Richmond River (1, AMCS786, ex Hedley); Washpool S.F,, SW of Casino. dry eucalypt forest (29°13730"S, 152°29°E) (1, AMC136879, 29 Aug 1982, ABRS - AM/QM); Wonga Walk, The Glade, Dorngo N.P., under bark of fallen tree, rainforest (30°22'S, 152°44’E) (1, QMMO17137, 6 Mar 1987, J. Stanisic, D. Potter); Mills ‘Rd, Moonpar S.F., under bark of fallen tree, rainforest (30°13°S, 152°39"E) (1, QMMO17141, 7 Mar 1987, J. Stanisic, D. Potter). SOUTH EAST QUEENSLAND «¢c. 1 km Eof Carr's Lookout, Koreelah $.F., under bark of fallen tree, NVF (3, OMMO16470, 3 May 1986, J. Stanisic); Mt Glorious, litler, NVF (1, QMMO16130, 26 Jan 1986, J. Stanisic); Mt Glorious, ¢. 3 km SE of summit, NVF (27°20'S, 152°,46'E) (1, QMMO11992, 2 Dec 1982, ABRS - AM/QM): c. 28.5 km from Goomburta, Goomburra S.F., 600 m, CNVF/Palms (27°59'S, 152°21'E) (3. AMC136812, QMMO12690, 7 Dec 1981, ABRS - AM/QM); Fred’s Road, MI Mee, rain- forest (27°05'S, 152°43'E) (2, QMMO16460, 14 Apr 1980. J. Stanisic, N. Hall, A. Green); Maitala N.P., ML Glorious, liller (1, QMMO11852, 20 Jun 1982, MSA Party); Lower Ballanjui Falls circuit, Lamington N.P., litter, NVF (1, OMMO16448, 3 Oct 1986, J. Stanisic, D. Potter); Canungerah (= Canungra), pine scrub (5, AMC32997, Jul 1908, S.W. Jackson); Mt Tamborine (2, AMC 142442); Maiala N.P., Mt Glorious, CNVF (1, QMM06304, 1 Oct 1976, M.J. Bishop); Mt Mis- take, Goomburra S.F., under bark of fallen trees, CNVF (2, QMMO16571, 5 May 1986, J. Stanisic); Running Ck, Lion's Road, via Rathdowney, under bark of fallen (ree, rainforest neat picnic area (2, 18 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Ngairea dorrigoensis (Iredale, 1941). a—c, Dorrigo Scrub, NSW. AMC63517, holotype; d-e, c. 1 km E of Carr’s Lookout, Koreelah S.F., NSW. QMMO16470; £, Upper Pine Ck, Canungra, SEQ. QMMO16544. a—c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, central (top left) and lateral teeth. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 19 lmm FIG. 4. Ngairea dorrigoensis (Iredale, 1941). a, c—e, Maiala N.P., Mt Glorious, SEQ. QMMO6304. b, Upper Pine Creek, Canungra, SEQ. QMMO16583. a, genitalia; b, details of penis interior, c, ovotestis; d, hermaphroditic duct; e, pallial cavity. Scale lines as marked, ltr lr Orel EIS TFORRL 971 L7L MEMOIRS OF THE QUEENSLAND MUSEUM bya ort Pr OLLVa Wa 20 (eR 12-6741) ram 871 Ie sro Le (re-0e'0) ClO OF LED (wu) HLGIM ‘vorlawna 4 - 90 = ~ 6c0 ae a 670 (ZE0-L1'0) = - SPO OF09TO (0-970) a ZENUFIED ~- “ 60 res “? 870 > 7 670 = ~ 60 “=< + 90 om os cal) ud = ceo “ ae 0 WW/SAIY See OLLVY MAMI/dS (p98 "XOD) JAY VIMIVON ONV ‘AON “dS VIVISOMANT VAM) SB we woz (pr't-z0'z) ATT OFNOLT (tvz-€6' 1) TOLUFELT sft (ww) HLCIAN TYOHA AGOF oso aso “oo 104 650 650 (L9°0-zp'0) FL0'07095°0 (ps'0-05°0) ROO'OFLIO ran) 650 650 ps0 oso oso £90 oso (wus) NOISNYLOYd AadS 6rd £50 8s0 £90 oL0 oL0 ico 6s'0 (60-250) £0007 0850 (z9'0-25'0) 1e0'0F 950 V0 6s'0 6S0 sso OLLVY d/H (ZONVY GNV WS ‘NV3IN) oF's ses eos oro (ss°9-gk'5) £27'0F01'9 (OP'L-ze'9) 961 UF WNL PIL 6e9 lek 469 Los oe'9 ws £79 (ww) waLLAWVIG 987 6lre 61 OLE SOE 6L coe one (L8°¢-9¢'¢) OLVOFESE (z9°e-19'¢) £TCOFRG'E Ley RLE £0'P Sty I9'E Ole 9E'E SPE (wu) LHOIH “p/P R/S & “BILE R/LE (R/L 601 +Z/1 b) “PEP +B/IL B/LS a/Sh STYOHM VON (P61 ‘TIVGAUI) SISNTODIAYOM VAMIVON NI NOILVINVA TV9O01- € F18V.L SNAWIOddS dO da egWON £691 ONWO suey Aowny R967FLOWY BUEMIQUIEL 1A 1600ST OWY (adhioan) Z9H7FI OWV BD IW idydanu vasoSy £SIL1 OWWO dep s,weqauiuuns (adAyojon) 65601 ONWO “A'S YE}PBIOy DIDISOI AA) Dasa 8rr9y| ONWO “dN YOu] ZL+91 OWWO 4D Suruuny L667£ DWV eadunues PSII ONWO 4D aug saddy, O9S8z1 DNV neared BIDRIy SLe7hl OWV 6£601 ONWO ‘a’§ Aneag LSLLEL OW wejngey, Ovbzbl OWY “AN Puridug man 9LP7Pl OWV £1S¢9 DNV (adAjojoH) LISE9OWY otiu0g Sisuao3iuop pansy ANYN SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 21 25°s 27°S 30°S 31°5 154°E FIG, 5. Distribution of Ngairea dorrigoensts (Iredale, 1941). QMMO16472, 1 Jan }980, V. Kessner); Upper Pine Creek, Canungra, NVF/Araucaria, under bark (15, OMMO16544, 1 Oct 1986. J, Stanisic, D, Potter; 3, QMMQ16583, 29 Sept 1986, J. Stamisic, J, Chasel- ing). DIAGNOSIS Shell depressed, large, diameter 5.71-7.40 mm (mean 6.50 mm), with 4 1/8 to 5 1/8 (mean 4 1/4-) normally coiled whorls. Apex and spire moderately elevated, height of shell 3.36—-4.62 mm (mean 3.78 mm). H/D ratio 0.52-0.64 (mean 0.58), Body whorl descending slightly in front. SP/BWW 0.17-0.39 (mean 0.31). Protoconch with | 5/8- to | 3/4+ whorls, mean diameter 1065.0qm at | 1/2 whorls. Apical sculpture of low curved radial ribs, without spiral clements (Fig. 3d). Post nuclear sculpture of numerous protractively sinuated radial ribs whose inter- stices are 5—7 times their width. Microsculpture of crowded raised spiral cords (Fig. 3e). Sculp- ture continuous on the base (Fig. 3b). Umbilicus small. diameter 0.30-0.68 mm, (mean 0.46 mm), D/U ratio 9.26-21.83 (mean 14.59), to closed, Sutures slightly impressed. Whorls. flattened above and below an angulate periphery (Fig. 3c), Body whorl keeled, more angulate in juvenile specimens. Aperture ovately lunate. Lip simple, becoming thickened at the basal margin, Columella thickened and reflected over the um- bilicus. Parietal callus white with brown flam- mulations, pustulose. Colour yellow-horn with reddish-brown flammulations above, continued below the periphery and onto the base injuvenile specimens, but fading toward the umbilicus in adult specimens. Based on 16 measured adults, Epiphallus internally with longitudinal pilasters, entering penis through « large mus- cularised pilaster (Fig. 4b). Penis (Fig. 4b) a short cylindrical muscularised tube, more swol- len apically; internally with apical muscular col- lar consisting of 5-6 pad-like thickenings, and slender longitudinal fleshy pilasters in the lower regions. Penial retractor muscle a shortened tuft inserting on the epiphallus near its junction with the penis. Free oviduct muscular reflexed, much longer than vagina (Fig. 4a). Radula (Fig. 3f) with central tooth having a large, broadly triangular mesocone and reduced ectocones. Based on six dissected specimens (QM- MO06304, OMMO16471, OMMO16470, AMC137757, AMC142476). RANGE AND HABITAT Under the bark of rotting logs in moist sub- tropical notophyll vine forests of cool wet high- land areas of the Great Dividing Range (from New England, NSW to the Mistake Mountains, SEQ); the Border Ranges, and the D’ Aguilar Range. SEQ: penetrating adjacent drier subtropi- cal forest (low microphyll vine forest, semi- evergreen vine thicket) in the Koreelah Creek and Richmond Range areas, northern NSW. REMARKS Northern representatives of N, dorrigoensis are larger than their southern counterparts and have the umbilicus more closed (Table 3). Pos- sibly the warmer climate provides a longer growth period in northern populations, 22 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. Ngairea levicostata sp. nov. a-c, Along Acacia Plateau Rd, Koreelah S.F., NSW. QMMO10959, holotype; d-f, Cunningham’s Gap, SEQ. QMMO17153. QMMO12684, paratypes, a-c, entire shell; d, apical sculpture; €, post nuclear sculpture; f, lateral teeth. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 23 Ngairea levicostata sp. nov. (Figs 6-8; Tables 2, 3) EryMoLocy Latin levis, smoath; costa, tib; referring to reduced microsculpture on major ribs of adult whorls. COMPARISONS N. levicostata is most similar to NV. dorrigoen- sis but differs in having a higher spire, fewer whorls with different sculptural features, a protoconch with curved radial ridges and fine, weak, crowded spiral cords and in lacking the microsculpture of crowded wavy spiral cords present in.V. dorrigoensis. Species of Mussonula spp. are separable from N. levicostata by their exsert protoconchs and apical sculpture of bold spiral cords (Figs 15d, 20f). TYPE MATERIAL Hoiotyre: OMMO10959, Koreelah S.F., NENSW, along Acacia Plateau Rd, 850 m (28°21'S, 152°24°E), 15 Mar 1981, AM/QM - ABRS 1981. Height of shell 4.79 mm, diameter 6.03 mm, H/D ratio 0.76, D/U ratio 13.35, whorls 4 5/8. PARATYPES: Cunninghams Gap, SEQ, near top, 755 m, NVF/Arawearia (28°04°S, 152°24'E) (3, QMMO12684, QMMOQ 10994, AMC128610. 16 Mar 1981, AM/QM - ABRS 1981); Cunningham's Gap, SEQ. leaf litter (1, QMMO17153, 18 Jun 1979, G. Annabell). DIAGNOSIS Shell trochoid, large, diameter 5.63-6.30 mm (mean 5.97 mm) with 4 1/4 to 4 5/8 (mean 4 1/2-) loosely coiled whorls, Apex and spire strongly elevated, height of shell 3.95-4.79 mm (mean 4.37 mm). H/D ratio 0.70-0.76 (mean 0.73). Body whorl descending more rapidly, SP/BWW ratio 0.36—-0.44 (mean 0.40). Protoconch with 1 5/8 whorls, mean diameter 1119.0u.m at 1} 1/2 whorls. Apical sculpture of low curved radial ribs crossed by raised spiral cords (Fig. 6d). Post nuclear sculpture of numerous crowded, high, protractively sinuated radial ribs (Fig. 6e), Fine raised microspiral lines present but no microriblets. Sculpture continued onto the base (Fig. 6b). Umbilicus narrow U-shaped, diameter 0.45-0.47 mm (mean 0.46 mm), D/U ratio 12.46-13.35, (mean 12.91). Sutures impressed, Whorls flattened above and rounded below an angulate periphery (Fig. 6c). Aperture sub- quadrate. Lip simple with a slight thickening at the baso-columellar margin. Columella almost Imm FIG. 7. Neatirea levicostata sp. nov. Cunningham's Gap, SEQ. QMMO12684, paratype. Pallial cavity. Scale line as marked, vertical, reflected toward the umbilicus, Parietal callus well developed. Colour yellow-horn with adhering dirt particles. Based on 2 measured adults. Foot and tail moderately broad, tapering posteriorly with a weak caudal horn. Manile collar thickened with a4 strongly developed glan- dular zone (Fig. 7). Kidney vaguely triangular c. 1/3 length of pallial cavity, rectal lobe reduced. Pulmonary vein inconspicuous. Reproductive system immature. Radula (Fig. 6f) with central tooth having a slender lanceolate mesocone and small but prominent side cusps: inner laterals with slender mesocone, endocone absent and weakly developed ectocone, Based on one dissected specimen (QM- MO12684). 153°5 154°E FIG_8. Distribution of Ngairea levicostata sp. nov. 24 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 25 FIG. 10. Ngairea murphyi (Cox, 1864). c.4km E of Fitzroy Falls on Kangaroo Valley - Fitzroy Falls Rd, NSW. QMMO16932. a, genitalia; b, details of penis interior; c, talon and hermaphroditic duct; d, ovotestis; e, pallial cavity. Scale lines as marked. FIG. 9. Ngairea murphyi (Cox, 1864). a-c, Mt Keira, Wollongong, NSW. AMC142962, neotype; e-g, Woodhill NSW, AMC142968; d,h, c. 4km E of Fitzroy Falls on Kangaroo Valley-Fitzroy Falls Rd, NSW. QMMO16932. a-—c, entire shell; d, central and lateral teeth; e, spire; f, apical sculpture; g, post nuclear sculpture; h, lateromarginal teeth. Scale lines as marked. ZA MEMOIRS OF THE QUEENSLAND MUSEUM RANGE AND HABITAT N. levicostata is known from two localities on the cool wet highlands of the Great Dividing Range in northern NSW and SEQ where it is sympatric with N. dorrigoensts. The micro- habitat is unknown, REMARKS N. levicastata is erected to assist comparative and biogeographical discussions: its conchologi- cal features.are distinctive. Although the general shell facies relates to Mussonula, N. levicostata is included in Ngairea primarily on form of the protoconch, Ngairea mucphyi (Cox, 1864) comb. nov. (Figs 9-11; Tables 2, 3) Helix murphyi Cox, 1864, p. 37; Cox, 1868, p.23. Paralaoma mirphyr (Cox); Iredale, 1937a, p. 314; Iredale, 1941a. p. 264, COMPARISONS N. murphycis a southern outlier distinguished within the genus by its higher whorl count and wider umbilicus (Table 2). N. dorrigoensis which it most resembles, has a tiny to closed umbilicus, weaker keel, fewer whorls and pratoconch with curved radial ridges (Fig. 3d). Apical sculpture involves irregular radial rugosities and spiral wrinkles producing a pitted effect (Pig. 9f). Anatomically distinctive are the penis with epiphallic pore located subapically and the penial retractor muscle inserted apically (Fig. 1b). PREVIOUS STUDIES Cox (1864) introduced the name without an accompanying figure and referred to it again without figures in 1868. A further listing (Cox, 1909. p. 54) was accompanied by an explanation for the lack of figures: “I fear the types of this species have disappeared from our (Australian) Museum Collection. The specimens of il which I possessed were unfortunately smashed while in the artists hands”. The large whorl count, small size, widely open umbilicus, sharply keeled body whorl and reticulate sculpture, referred to in Cox’s original description, together with the type locality of Wollongong, NSW, effectively fix Cox's name to the species discussed below, The neotype was collected at Mt Keira, Wollon- gong, which I consider to be near the original type locality. Iredale (1937a, p. 314) placed A. murphyi in Paralaoma Iredale, 1913, withoul explanation, later (Iredale, 1941a, p. 264) admitting that no known specimens of H. murphyi existed and that this decision was based on broad details of Cox’s description. [In Paralaoma the shell is much smaller, more fragile, with fewer whorls and bold spiral cords on the protoconch, Although morphology of NM. muwrphyi is unusual, penial morphology, shell sculpture and peculiar bady colour are considered sufficient to endorse the new generic placement. TYPE MATERIAL Nearyre: AMC142962, Mt Keira, Wollongong, NSW, Collected by C.F. McLauchlan, 6 Noy 1948, Height of shell 3.70 mm, diameter 5.88.mm, H/D ratio 0.63, D/U ratio 4.19. whorls 5 7/8. OTHER MATERIAL c. 4 km E of Fitzroy Falls on Kangaroo Valley - Fitzroy Falls road, NSW, temperate rainforest, under bark of fallen tree (5, OMMO16932, J. Stanisic, 4 Jan 1987): Woodhill, c. 4 mls NW of Berry. NSW, mixed forest, E side of range (1, AMC142968, W. Ponder, 14 Now 1970): Mt Cambewarra, nr Nowra. NSW (1 AMC142964_L. Price..30 Oct, 1963): Mt Keira, Wol- longong, NSW, from rainforest at scout camp (1, AMC1429b62, C.F. McLauchlan. 6 Nov 1948), DIAGNOSIS Shell depressed to elevated trochoid, moder- ately large, diameter 5.46-6.05 mm (mean 5.80 34's 34°30'5 KS Sf) alt. oe oe > 500m ahec 1sp°F r50°30'E TEs Fic 11, Distribution of Ngairea murphyt (Cox, 1864) in (he IHawarra region, New South Wales, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE mat) with 5 to 5 7/8 (mean 5 |/2-) tightly coiled whorls, Apex and spire weakly to strongly elevated, height of shell 2.86-3.70 mm (mean 3.24 mm), H/D ratio 0.49-0.63, (mean 0.56). Body whorl not descending in front, SP/BWW (),32-0.65 (mean 0.44). Protoconch with ir- regular curved radial ridges and pits, spiral ele- ments absent (Fig. 9f). Post nuclear sculpture of numerous, crowded, protractively sinuated radial ribs (Fig. 9¢, g), whose interstices are ahoul 5—7 times their width. No microriblets. Microsculpture of numerous, crowded, narrow spiral cords. Sculpture continuous on the base but reduced (Fig. 9b), Umbilicus relatively wide, V-shaped, diameter 1,.28-1.58 mm (mean 1,39 mm), D/U ratio 3.84—4.39 (mean 4.17), Sutures impressed, Whorls flattened above and below a cdrinate periphery (Fig. 9c), Aperture ovately Junate to subquadrate, Lip simple with a slight thickening at the baso-columellar margin. Calumella almost vertical, slightly reflected toward the umbilicus. Parietal callus well developed, white, pustulose. Colourcreamy-yel- low horn without flammulations. Based on four measured adults. Kidney (Fig. 10e) with clongate pericardial jabe and no rectal lobe, 1/2 length of the pallial region. Ovotestis (Fig. 10d) three or more clumps of creamy-white palmately clavate lobes of alveoli. Epiphallus (Fig. 10a) twice length of the penis with internal longitudinal pilasters, entering penis subapically through a muscular pilaster (Fig, 10b). Penis (Fig. 10b) short, cylindrical, muscularised, internally with an api- cal muscular collar and fleshy, longitudinal pilasters below the collar. Penial retractor muscle (Fig. 10a, b) inserted apically on the penis. Vagina relatively long (Fig. 10a), Radula (Fig, 9d, h) with mesocone of central tooth slender lanceolate and ectocones small but prominent. Based on 2 dissected specimens (QM- MO LY632). RANGE AND HaAdliiTat N. murpliyi occurs in an area which has been greatly allered and disturbed by white-settlement (Baur, 1957, pp. 191, 195). Its distribution en- compasses the remnant patches of moist warm temperate and cool subtropical vine forests of the Illawarra region, NSW, from Wollongong south to Mt Cambewarra, near Nowra, The original specimens Were taken from under stones; live material has been collected from under bark of a fallen 1ree (QMMO16932). ta —4 REMARKS The high whorl count has not been observed in any other east coast charopid with a trochoid shell, The North Queensland helicarionid Thes- kelomensor (Iredale, 1933 has a trochoid shell, wide umbilicus and high whorl count but the similarity is convergent. The few available specimens of N, murphyi show a greal deal of variation in shell height brought about by chan- ges in spire protrusion (Table 3), Ngairea canaliculata sp, nov, (Figs 12-14; Tables 2, 4) Flammulina corticicola (Cox); Hedley. 1912 (part). p. 204, but not figures ETYMOLOGY Latin canaliculus, small channel; for the chan- nelled sulures. COMPARISONS Readily identified by its channelled sutures, low spire, rounded periphery and sculpture of low, broad radial ridges crossed by fine, crowded microspiral grooves (Fig, 12g), Lenwebbia pro- toscrobiculata which is sympatric with N. canaliculata has similar shell shape but differs markedly by having a punctate protoconch, fine thread-like radial riblets on the budy whorl, and simple sutures (Fig. 29c—g). TYP MATERIAL HoLatyre: OMMOL6454, Beauty Spot 98, Kroom: bil Tops, SEQ. Collected by G.B. Monteith, 29 Sep 1983, Height 3.87 mm, diameter 6.55 mm, H/D ratio 0.59. whorls 4 5/8. PARATYPES! SOUTH EAST QUEENSLAND - QM- MO17154, 1 specimen, same data as holotype; Kroombil Tops, TA 47 creek (1, OMMQO 16467, 9-19 Dec, 1983, G.B. Monteith, G. Thompson); Kroombil Tops. under palm fronds, NVF/Palms (1. OMMO14890, 8 May 1984, J. Stanisic, D. Potter); ML Fort William, Kalpowar $,F,, NWF (24°39°S, 151° 2 E) (2, OMMO12598, AMC136757, 4 Sept 1982, ABRS-AM/OM; Camp Creek, S of Miriam Vale, SW side ol Bobby Range, MVF (24°36'20"S, 151°33'E) (3, OMMO13290, AMC 137913, 6 Sept 1982, ABRS - AM/OM); Rainforest walk, Kalpowar S.F.. MVF/Araucaria, (24°41°8, 151°21°E) (4, AMC- 136742, QMMO12574, 4 Sept 1982, ABRS - AM/QM): Bobby Range S.F., NVF (24°37'S, 151°32°E) (2, OMMO17151, AMC136509, 6 Sept 1982. ABRS-AM/QM): Bulburin (Austral) S.F., S of Gladstone, 380 m (24°34'S, 151°29'E) (1, MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 29 FIG, 13. Ngairea canaliculata sp. nov. a, ce, c. 12.1km from Granite Creek crossing on Bobby Range-M1 Perry Rd, SEQ. QMMO 16468. paratype. b, Mt Fort William, Kalpowar S.F..SEQ. QMMO16469, paratype. a, genitalia: b, details of penis interior; c, ovotestis; d, hermaphroditic duct; e, pallial cavity. Scale lines as marked. FIG. 12. Ngairea canaliculata sp. nov. a—c, Beauty Spot 98, Kroombit Tops, SEQ. QMMO16454, holotype; d-h, summit, Mt Fort William, Kalpowar S.F.. SEQ. QMMO16469, paratype. a—c, entire shell; d, central and lateral teeth; e, details of spire showing channelled sutures; f, apical sculpture; g. post nuclear sculpture; h, details of base and umbilicus. Scale lines as marked. i MEMOIRS OF THE QUEENSLAND MUSEUM 23°S \ Rockhampton 24°s 25°S 26°S 27°S x! 150°B 151°E 152°E 153°E FIG. 14. Distribution of Neairea canaliculaia sp. nov. AMC142465_8 May 1975, 1.B. Burch, W.F. Ponder. PH. Colman); Bulburin (Austral) S.F., S of Gladstone, 540 m, CNVF-A1 forest type (24°31'S, 151°29°R) (1, AMC 142466, J.B. Burch, W.F. Ponder, P.H, Colman): Bortlearee senib, west of Gladstone, on the ground under leaves (1, AMC32998, Aug 1908, 8.W. Jackson); Colosseum Creek. 16 km-S of Mitiam Vale (2, AMC142445, 25 May 1958, L. Price); Kal- powar S.F., litter. MVF/Araucaria (9, OMMO16459, 4 Jul 1984, D, Potter, J. Stanisic, K. Emberton); Mt Larcom, on rocky slope, litter, SEVT (2, QMMO16443, 9 May 1984, J. Stanisic, D. Potter); Summit, Mt Fort William, Kalpowar S.F., under bark, CNYF (4, OMMO 16469, 3 Jul 1984, J. Stanisic, D, Potler, K. Emberton); c. 12.1 km from Granite Creek crossing on Bobby Runge - MI Perry Road, under bark of fallen trees, NVF/Palms (24, OMMO16465, 16 Sept 1985, J. Stanisic, D. Potter). OTHER MATERIAL SOUTH EAST QUEENSLAND: Colosseum Creek, SW of Miriam Vale, rainforest along creek (24°23°30"S, 1S1°27°E) (2, AMC137890, QMMO 13223, 6 Sept 1982, ABRS - AM/OM); Sum- mit af Mt Booroon Booroon, on rocky slope, litter, MVFi/Araucarta (2, QMMO16444, 17 Sept 1985, J. Stanisic. D. Poller); Kroambit Tops. 13 km W of ‘Chapman’s’. NVF (24°25°S., 1351°02°E) (1, Curtis (3, AMC63752. ex Cox): near Colosseum Ck, 16 km S of Miriam Vale (4. AMC142446, 24 May 1958. 1. Price); Miriam Vale (f, AMC152447, ex Musson), DIAGNOSIS Shell depressed, moderately large, 5.97-7,39 mm (mean 6.53 mm) in diameter, with 4 1/2 to 4 3/4 (mean 4 5/8-) normally coiled whorls. Apex and spire moderately elevated, height of shell 3.28-5,35 mm (mean 3.83 mm), H/D ratio ().53-0.83 (mean 0.59). Body whorl descending only slightly in front, SP/BWW ratio 0,180.32 (mean 0,24). Protoconch flat, shiny, with | 5/8 to | 5/8+ whorls, mean diameter 1169.0j.m al 1 1/2 whorls. Apical sculpture of very weak, low, curved radial ridges crossed by irregularly spaced incised spiral lines (Fig. 12f). Post nuclear sculpture of numerous, regularly spaced, low, broad. protractively Sinuated radial ribs (Fig. 12e), whose interstices are about 7-9 times their width. No intervening microriblets. Microsculpture of incised spiral lines (Fig. 12g). Sculpture continuing onto base (Fig. 12h) with spirals fading toward the umbilicus. Umbilicus small, diameter 0.43-0.94 mm (mean 0.71), D/U ratio 7.21-15.63 (mean 9.40). Sutures distinctly channelled. Whorls rounded above and below the periphery (Fig, 12c). Body whorl without any noliceable angulation, even in juvenile specimens. Aperture ovately lunate. Lip simple, becoming thickened at the baso-columellar mar- gin. Columella thickened, twisted and slightly reflected over the umbilicus. Parietal callus pus- tulose with some flammulations. Colour deep golden-horn with regularly arranged reddish- brown flammulations above, continuing below the periphery but fading near the umbilicus. Based on 21 micasured specimens. Vas deferens (Fig. 13a) a very thin tube deli- cately convoluted along its length. Epiphallus (Fig. 13a) very long. muscularised. reflexed half way alone its length, internally with longitudinal pilasters, entering penis apically (Fig. 13b) through a conical verge (Fig. 13b). Penial retrac- lor muscle Jong, inserted at the epiphallus-penis junction. Penis (Fig. 13b) a cylindrical, mus- cularised tube, swollen apically, internally with anapical fleshy collar and low, corrugated, spon- gy longitudinal pilasters. Free oviduct (Fig. 13a) strongly musecularised, about 3-4 times the length of the vagina. Vagina very short, internal- ly with fleshy longitudinal lamellae. Atrium short, without unusual features. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE Central tooth of radula (Fig. 12d) with broad triangular mesocone and small ectocones. Based on 4 dissected specimens (QM- MO14890, OMMO16468, OMMO 16469). RANGE AND HABITAT Neairea canaliculata inhabits warm moist Subtropical notophyll vine forests of the ranges and mountain caps between Colosseum Creek and Mt Latcom, SEQ. This region also has large areas. of drier sub-tropical forest (vine thickets and microphyll vine forests) but NV, canalicnlata shows little tendency to occur outside of the moister areas, The species has been collected live under bark and in the rolled up stems of discarded palm fronds. REMARKS This species was first collected by Sidney W. Jackson in 1908, but included under Flarnulina vorticicola (Cox, 1866) by Hedley (1912). It is distinguishable from that species by its chan- nelled sutures and radially sculptured base. The channelled suture is an unusual development seen in a limited number of Australian, New Zealand and Pacific Island species. Ngaivea corticicola (Cox, 1866) comb. nov. (Figs 15-17; Tables 2, 4) felix corticicala Cox, 1866a, p. 374 .Cox, RGR. p. 149, pl.7. figs 77a. Oreakera corticicola (Cox); Iredale, 19374, pais: fredale, 1941a, p. 264. COMPARISONS N. corticicala can be distinguished from N. dorrigoensis by its less prominent radial sculp- ture, almost smooth base. more rounded periphery, microsculpture of incised spiral grooves (Fig. 15f) and penis with corrugated, longitudinal pilasters and a continuous. weakly muscular, apical collar.N, canaliculata (Fig. 12) has similar microsculpture to N. corsicicola but differs in having low broad radial ridges, fewer whorls, less elevated spire, channelled sutures, more rounded shell periphery and an epiphallus almost twice as long as the penis (Fig. 16a). PREVIOUS STUDIES Hedley (1912) attempted to clarify Helix cor- ucicola Cox, 1866 but'he considered specimens from Canungra, SEQ, to be conspecific and figured a representative, This was the species later described by tredale (19414) as Oreokera dorrigoensis. N, corticicola occurs microsym- patrically with V, dorrigoensis at Pine Creek, Canungra, but the two species can be distin- guished by sculpture on the base of shell, which is reduced in the former and strongly radial in the latter. TPE MATERIAI LECTOTYPE: AMC6352U. Lismore, NSW. Cox cole lection, Height of shell 3,87 mm, diameter 6.47 mm, H/D ratio 0.60. D/U ratio 10.78, whorls 5. PakaLecroryres; AMC6351 1, 3 specimens, same ditla as lecloly pe, OTHER MATERIAL NEW SOUTH WALES; Wollongbar (1, AMC142431, Aug 1905): Richmond River (4, AMC142432. ex Cox, Petterd); Rous Mill, Richmond Rv., (6. AMC142433, 1900, S.W. Jackson); Lismore, Richmond Rv., (9, AMC142434, ex Brazier); Rich- mond Rv., (1, AMC142435_ ex Helms); upper Tweed River (7. AMC14243h. ex Cox, Petterd): Lowanna, near Dorigo (1, AMC 63816, M. Ward): Borngall CX. Bellinger River, under bark (1, AMC142437, ex Cox); Opposite Kelly selection, 16 km fram Murwillumban, in gully on main toad (3, AMC63835); Red Scrub Flora Reserve, Whian Whian S.F,, 210 m (28°38°S, 153° 19 E) (13, AMC 142473, 15 Mav 1976, P-H. Cul+ man, |, Loch); Terania Creek, Whian Whian S.F., 340 m. (28°34°S, 153°19°E) (1. AMC)42474, 16 May 1976, P.H. Colman. 1. Lach): Dorrige Range, (4- AMC 142438, Apr 1957, L. Price); Ballina, in rotten wood (10, AMC142439, ex Cox): Big Scrub, Whian Whian State Forest. litter, SNVF (1, OMMO16441,, 15 Noy. 1983, ABRS - AM/QM): Upper Wilson's Creek, litter, NVF (28°32°S, 153°24°E) (1, OMMO 16443, 16 Nov 1983, ABRS - AM/QM), Nightcup Range, under bark of fallen tree (28°34'S, 153°20'E) (2, OMMO16466, 15 Nov 1983, ABRS - AM/OM); The Glade, Dorrigo N.P., under log, cool temperate rainforest (30°22°20"5, 152°43°40"E) (1, OMMO14766, 12 Nov 1983. ABRS - AM/QM); Swan's Road, Bruxner Park Flora Reserve, under bark of fallen logs and in palm fronds on ground, subtrupi- cal rainforest (22. OMMO14767, QMMO16757, 14 Nov. 1983, ABRS - AM/QM): Wonlgoolga Ck Flora Reserve, Wedding Bells S.F_, under leaf in subtropical raintorest (1. AMC152252. Mar 1983, M. Sheu); Cur- Tumbin Valley Nature Reserve, under log in rainforest (3. AMC 152254, Jul 1977. M. Shea): Lismore Serub. Lismore. under birk of fallen trees, remnant rainforest (4, OQMMO17135.9 Mar 1987, J, Stanisic, D, Potter); Wonga Walk, The Glade. Dorrigo N.P., under bark of fallen trees. rainfores! (35, QMMOQI7146, 6 Mar 1987. J. Stanisic, D. Potter); Never Never Prenic Area, a i) MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE FIG. 16. Negairea corticicola (Cox, 1866). Slopes of Mt Hobwee, Lamington N.P., SEQ. QMMO16462. a, genitalia; b, details of penis interior; c, talon; d, hermaphroditic duct; e, ovotestis: f, pallial cavity. Scale lines as marked. FIG. 15. Ngairea corticicola (Cox, 1866). a-b, Lismore, NSW. AMC63520, holotype: c, Tullawallal Circuit, Lamington N.P., SEQ. QMMO16465; e-g, Red Scrub F.R., Whian Whian S.F., NSW. AMC142473; d, h, Slopes of Mt Hobwee, Lamington N,P., SEQ. a-c, entire shell; d, central (bottom left) and lateral teeth; e, apical sculpture; f, post nuclear sculpture; g, details of umbilicus; h, lateromarginal teeth, Scale lines as marked. MEMOIRS OF THE QUEENSLAND MUSEUM 34 08 ee vee ve'# (.V6-LVe) SUS'UFELR SUL Ce lt-£6'2) LEN VHT {ez-e-L0'8) HOOF 6FL (09'8-66°9) MEE'OFRR'L (sp's-1P'L) O70 LER RLOL (ro'R-1IC'K) OL OFLF'R fre (s0'6-12°L) Plz 0FSP'R ech Un (pR'6~tt'h) SS7'0705'6 ROOT tt rcailt an fost as frail OLLva vd seu sR'0 iO oR) (zot-2e) SsU'1F28' ro" (ax'0-S5°0) LOVOFIL'Y wo roo (6R°0- S80) O10'079R'0. 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AA Jaddr) r £SL91 ONWO qed uxTg z £LeZhl ONY “A'S URI BETH AN (ad Mojo) t O¢SE9 DWV SUOUIST] 0919040 Dasnin z Spezrl OWY 42213 tunassojo 1 P91 ONWO WOE] IW 9 6S¢91 ONWO 1 TPLYEL OWY 1 86871 OWWO z #871 ONWO “y'Saemodyey 99pzhlL OWY S9bZtl OWV IsIZ1 ONWO O6zEl ONWO atuey Aqqog 1 L9¢91 ONWO i FSILLOWAOD {ad Kojo) 1 rSp9l ONWO sdoy uqwoory oInpnnpouns vas0dN SNIWIOddS 40 WaHIWAN JNYN SYSTEMATICS OF AUSTRALIAN CHAROPIDAE \ 26°S 27°S 28°S 29°S 30°S 31°S 151°E 152°E 153°E 154°E FIG. 17. Distribution of Ngairea corticicola (Cox, 1866) Dorrigo N.P., under bark of fallen tree, rainforest (1, QMMO17142, 6 Mar 1987, J. Stanisic, D. Potter); Booyong Flora Reserve, via Lismore, under bark of fallen trees. rainforest (21, QMMO17005, 9 Mar 1987, J. Stanisic, D. Potter). SOUTH EAST QUEENSLAND - Lower Ballanjui Falls Circuit, Lamington N.P., litter, NVF (3, QMMO16447, 3 Oct 1985, J. Stanisic, D. Potter): Tullawallal Circuit, Lamington N.P., CNVF (3, QMMO16465, 2 Oct 1985, J. Stanisic, D. Potter); Warrie Circuit, Springbrook N.P., under bark of fallen tree, CNVF (4, QMMO16464, 6 Apr 1986, J. Stanisic); Orchid Bower, Lamington N.P., 900 m, CNVF (8, QMMO6264, Apr 1976, M.J. Bishop); Nagarigoon Falls Circuit, Lamington N.P., CNVF (2, QMMO16463, 20 Apr 1986, J. Stanisic); Warrie N.P., Springbrook, CNVF (28°00°S, 153°00°E) 15, QMM06300, 4 Oct 1976, M.J. Bishop); Nagarigoon, Lamington N.P., 800 m, CNVF (28°14°S, 153°13°E) (13. QMMO6085, Apr 1976, M.J. Bishop): Mt. Hob- wee Circuit, Lamington N-P., litter (3, QMMO16445, to wv 1 Oct 1985, J. Stanisic, D. Potter); Natural Bridge N.P., NVF (28°13°S, 153°14°E) (1, QMMO10456, 18 Mar 1981, ABRS - AM/QM): Tullawallal Circuit, Lamington N,P., litter, NVF (1, QMMO16446, 2 Oct 1986, J. Stanisic, D. Potter); Coomera Falls, Lamington N.P., under bark of fallen tree, CNVF (2, QMMO16461, 19 Apr 1986, J. Stanisic); Slopes of Mt Hobwee, Lamington N.P., under bark of fallen trees, CNVF (17. QMMO16462, 19 Apr 1986, J. Stanisic); Upper Pine Creek, Canungra, under bark, NVF/Araucaria (1, QMMO16545, 1 Oct 1986, J. Stanisic, D. Potter: 7, QMMO16903, 21 Jan 1987, J. Stanisic, D. Potter: 3, QMMO16584, 29 Sept 1986, J. Stanisic, J. Chaseling); Araucaria Track, Binna Burra, under bark, NVF (5, QMMO16577, 27 Sept 1986, J. Stanisic): Lower Ballanjui Falls Track, Binna Burra, Lamington N.P., under bark (5, QMMO16550, 25 Sept 1986, J. Stanisic); Coomera Falls Circuit, Binna Burra, Lamington N.P., under bark, NVF (11, QM- MO16574. 26 Sept 1986, J. Stanisic). DIAGNOSIS Shell depressed, large, diameter 5.88-7.82 mm (mean 6.90 mm), with 4 3/8 to 5 1/2 (mean 4 3/4+) normally coiled whorls. Apex and spire moderately elevated, height of shell 3.36-4.62 mm (mean 3.90 mm), H/D ratio 0.50-0.63 (mean 0.56). Body whorl descending slightly in front, SP/BWW ratio 0.22-0.39 (mean 0.31). Protoconch with 1 5/8 to 2 whorls, mean diameter 1056.8um at 1 1/2 whorls. Apical sculpture of curved radial ribs crossed by incised spiral lines, most noticeable near the sutures (Fig. 15e). Post nuclear sculpture of low, weak protractively sinuated radial ribs and ridges, oc- casionally with a vertical periostracal blade (Fig. 15a), interstices 3-5 times their width and without radial riblets. Microsculpture of incised spiral grooves (Fig. 15f). Sculpture reduced on base to almost smooth (Fig. 15b). Base shiny to dull. Umbilicus small, 0.55-1.02 mm (mean 0.81 mm) in diameter, D/U ratio 6.99-11.31 (mean 8.61). Sutures impressed. Whorls flattened above and rounded below the periphery. Body whorl with a weak keel (Fig. 15c), becoming less pronounced in larger specimens. Aperture ovate- ly lunate. Lip simple, slightly thickened at the basal margin, becoming more so at the columel- la. Columella thickened, twisted and slightly reflected over the umbilicus. Parietal callus pus- tulose. Colour creamy-white to yellow-horn, upper surface with regular brown-red flammula- tions fading at the periphery, base creamy-white. Based on 34 measured specimens. Vas deferens giving rise to a long, muscular J0 MEMOIRS OF THE QUEENSLAND MUSEUM epiphullus (Fig. 16a). Epiphallus reflexing api- cally to enter the penis through a simple pore (Fig. 16b) surrounded by a spongy pilaster, in- ternally with longitudinal pilasters, Penis (Fig. l6b) slender, cylindrical, muscular, slightly swollen apically, internally with an apical fleshy collar and strongly corrugated, longitudinal pilasters. Penial retractor muscle long, inserting on the epiphallus prior to its entry into the penis. Radula (Fig. 15d,h) with broad triangular mesocone and reduced ectocones on central tooth. Based on four dissected specimens (QM- MQ16462, OMMOA330). RANGE AND HABITAT N. corticicola inhabits warm subtropical notophyll vine forests of the coastal lowlands and foothills between Dorrigo and Lismore, the Dorrigo Plateau, and the eastern McPherson Ranges. Its northern limit appears to be Mt Tam- borine, SEQ. On the basis of material collected late last century and early (his. century, it would appear that the Big Scrub formed an important part of the range of N. carticicola. In spite of the fact that much of the Big Scrub was cleared by early cedar-getters and more recently for agricul- ture, N. certicicola is still common in some of the remnants e.g. Booyong Flora Reserve, near Lismore, NSW. The species lives under the bark of rotting logs. REMARKS Southern representatives of N, certicicola (QMMOI7I46, AMC142438) have compara- tively lower whorl counts than northern specimens (Table 4). Mussonula Iredale, 1937 Musvonule Iredale, \937b,p, 13 TYPE SPECIES Mussonula verax Iredale 1937; by original designation. PREVIOUS STLDIBS Iredale (1937b) related Mussouula to Paralaoma Vredale, 1913 which is a punetid eroup with a spirally lirate protoconch, Although Iredale (1937b) noted spirals in M. verax he lujled to notice the less conspicuous apical radial clements referred to in this study (Figs 18d, 20f). Combined with general shell sculpture and details of the radula (Fig. 20d, e), the apical sculpture places Mussonula in the Charopidae, DIAGNOSIS Shell small ta moderately large, trochoid, adult shell diameter range 4+.81-5.63 mm, with 3 7/8 to 4 1/8 normally coiled whorls, last whorl des- cending. Height of shell 3.32-4.45 mm. Apex and spire strongly elevated. Protoconch strongly exsert. Apical sculpture of numerous bald (verax) to fine (fallax) spiral cords, and very weak radial ridges which become more prominent near the nuclear-post nuclear junc- tion, Post apical sculpture of numerous. high, protractively sinuated radial ribs with periostra- cal blades inclined toward the shell apex. Some trace of weak incised spiral lines (fallax) but no microriblets present. Sculpture continuous on the base. Umbilicus narrow U-shaped. Sutures impressed. Whorls rounded above and below a strongly angulate periphery. Aperture sub- quadrate. Lip simple, columella reflected slight- ly toward umbilicus. Colour yellow-horn to brown with adhering dirt particles, Pallial cavity with incomplete secondary ureter and kidney with vestigial rectal lobe. Genitalia unknown, Radula as for Mussonula fallax. DISTRIBUTION AND ECOLOGY Mussonula extends from Byangum on the Richmond River. northern NSW to the Gympie forestry area, SEQ. It inhabits moist lowland notophyll vine forest in the south, and drier microphyll vine forest (witha raucaria) and vine thickel in the north. The two species are al- lopatric, Little is known of the microhabit for although C.T. Musson collected the type series of M, veray arboreally, and a single live specimen of M, fallax was found under timber among rocks, all other specimens have come from litter samples. The irochoid shell und dark animal colour indicate an arboreal or semi-arboreal ex- istence. However the adhering dirt particles on the shell Suggest that it is a ground dweller, Possibly it lives among triable carth, between and under rocks. and forages for food on vertical surfaces above the ground. COMPARISONS The extremely prominent radial ribs (Figs 18e, 20g). strongly angulate periphery, elevated spire, exsert protaconch with strong spiral cords (Figs 18d. 20f), and almost total lack of mictos- SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 37 c FIG. 18. Mussonula verax Iredale, 1937. a—c, Scrub at North Pine, SEQ. AMC63770, holotype; d-f, c. 2km E of summit, Mt Nebo, SEQ. QMMO16518. a-c, entire shell; d, apical sculpture; e, post nuclear sculpture: f, terminal region of protoconch. Scale lines as marked. AM MEMOIRS OF THE QUEENSLAND MUSEUM culpture in Mussonula are a combination of fea- tures not found in any other east coast charopid. Oreokera Stanisic, 1987, from north Queensland has the same shape and postnuclear ribbing as Mussonula but is larger and has quite prominent, reticulate microsculpture (Stanisic, 1987. pls Ie.d;2e,d.e). Hedlevoconcha Pilsbry. 1893 has a similar shaped shell but has strong reticulate sculpture and a punctate protoconch (Figs 23e, f: 26d). Mussonula verax Iredale, 1937 (Figs 18-19: Tables 2, 5) Thalassia delta (Pleitter); Hedley and Musson, 1892, p. 553 - misidentfication. Mussonula verax Iredale, 1937b, p. 13, pl. 1. fig. 11. COMPARISONS M. verax can be distinguished fram M. fallax by its larger size and bolder, more regular, apical spiral cords (Figs 18d, 20f), Ngairea levicostata has a similar shell to M. verax but is slightly larger with apical sculpture of curved radial ribs and weak spiral cords (Fig. 6d); N. darrigoensis which is sympatric with M. verax in the southern part of the D’Aguilar Range, SEQ. is larger, has a less elevated spire, and microsculpture of prominent spiral cords (Fig. 3e). TyPF MATERIAL HOLOTYPE: AMC635U1, in scrubs, urboreal. North Pine River, South Queensland, Coll. C.T. Musson. Heivht of Shell 4.54 mm, diameter 5.46 mm, H/D ratio 0,82, D/U ratio 13.29, whorls 3 7/S+, PaRATYPES: AMCA3770, 5 spectmens. Scrub at North Pine, SEQ. Coll. by CT, Musson. OTHER MATERIAL Fred’s Rd. Mt Mee, SEQ, rainforest, leaf hitter, (27°05°S.. 152°43°E) (1. OMMO7340, 14 Apr P980, J, Stanisie. N. Hall, A. Green); MI Nebo, SEQ, c. 2 km E of summit. NVF/Araucaria, leat liter (27°24°S, 15S2°97°E) (1. QMMO16S18. 13 Auge 1980, J. Stanisic¢); Wratten ’s Camp, Wrailens $.F., SEO. NVF, fitter (26°17'°S, 152°20'E) (1. QMMONVISOO. 17 Jul |980. J. Stanisic. A. Green); c. 0.5 km E of Mt Mia turnoff on Mt Mia - Kilkivan Rd, SEQ (26°15°S, S227 Er(1. OMMO12320, | Sept 1982, AM/QM - ABRS 1982): Maiala N-P.. Mt Glorious, SEQ, NVF (1, OMMQ11602, 20 Jun 1982, MSA Party). DIAGNOSIS Shell trochoid, small, diameter 5,045,603 mm 153°5 154° FIG. 19, 1937. Distribution of Mussonule verax Iredale, (mean 5.36 mm) with 3 7/8 to4 (mean 4-) loosely coiled whorls. Apex and spire strongly elevated, height of shell 4.03-4.45 mm (mean 4.31 mm), H/D ratio 0.79-0,.82 (mean 0,81). Body whorl descending more rapidly. SP/BWW ratio 0.52- 0.57 (mean 0.54). Protoconch strongly exsert with | 1/2 to | 1/2+ whorls, mean diameter 1370.5 pom at 1 1/2 whorls. Apical! sculpture of about 20 continuous spiral cords and curved radial ridges appearing in the last 1/8 whorl (Pig. 15d). Post nuclear sculpture of high, crowded protractively Sinuated radial ribs. 90-120 (mean 104.8) on the body whorl (Fig. [Se). Ribs/mm 5.68-6.75 (mean 6.20), Major ribs with slender backward curved periostracal blades, interstices 7-2) times their width. no radial riblets. Micras- culpture absent. Sculpture continuous on the base (Fig. L&b). Umbilicus very narrow, U- shaped, slightly obscured. diameter ().31-0.4) mm (mean (0.35 mm), D/U ratio 13,.29-17.19 (meun 15.27). Sutures impressed. Whorls rounded above and below an angulate periphery. Aperture subquadrate, Lip simple. columella iwisted, vertical. slightly covering the umbilicus. Parivtal callus pustulose, well developed. Colour yellow-horn with adhering dirt particles. Based on 4+ measured udulls. Anatomy unknown. TAULE5- LOCAL VARIATION IN MUSSONULA VERAN IREDALE, 1937 AND MUSSONULA FALLAY SILNOV (MEAN, SEM AND RANGE) UMINEICAL wip (rn) SP/LWW BODY WHORL. StIRE, NUMUER OF NAME DAI RATIO RULS/MM 1uns RATIO wibtHt (min) PROTRUSION HEIGHT DIAMETER HID RATIO (remy (mm) WHORLS SULCIMENS (mm) SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 39 (15.33-17,19) V2hath Yh at M92 (ALA! nS2 AAs. (5.68-6.97) ca ms ci i a 2 (00-97) (052-0055) Lal ethid as (1771.85) hor (OTIS kY2-1M) TRULIA) a2 (O.KD-DKT) S.172t. 7007 (5,04-5.29) 5.46 W1b20,0M) (4.034 24) 445 (44) 37+ Nurth Pine Seruhs AMC 6.451 Miuxsoniele verax (Heokptype) AMC 63770 Mt Mia vlysed 7K 445 Sas nay hoe 177 4b OMMO 12320 Massenola fallax Byangum a4 OKA 162 45 72 O64 jal 4h AMC 63471) (Holotype) Houmbiask S.Kb OM 342 an ony thot ANB. AMC 142449. RANGE AND HABITAT M. verax inhabits notophyll vine forests of the D°Aguilar Range and drier microphyll vine forests of the Gympie region, SEQ. REMARKS C.T, Musson collected this species during an expedition to SEQ in 1887 which was funded by noted malacologist Dr J.C. Cox. The species was misidentified as Thalassia delta (Pfeitfer, 1857) by Hedley and Musson (1892), Iredale (1937b, p. 13) noted the error and erected the genus. Although the type locality of the species is given broadly as "scrubs at North Pine River", the itinerary of Musson’s trip suggests that a more specific locality may be Mt Mee, SEQ. Mussonula fallax sp. nov. (Figs 20-22; Tables 2, 5) ETYMOLOGY Latin fallax, fallacious; referring to its having been previously identified as M. verax. COMPARISONS M. fallax differs from M. veraxby having finer, more irregular apical spiral cords (Fig. 20f) and a smaller shell. Hedleyoconcha delta (Pfeiffer, 1857) has a trochoid shell butis much larger with greatly elevated spire and has postnuclear sculp- ture of low radial ribs and spiral cords in a reticulate pattern (Fig. 23e, f). TYPE MATERIAL HoLotyre: AMC63870, No. | Scrub at Byangum, NSW, Cox coll. Height of shell 3.41 mm, diameter 4.32. H/D ratio 0,64. D/U ratio 6,18, whorls 4 1/8-. PARATYPES: Mt Warning N.P.. NSW (28°24°S, 153°16°E) (1, AMC129289, 18 Mar 1981, ABRS - AM/QM): Upper Coopers Ck, Huonbrook, NSW (1, AMC142449, 26 Aug 1936, A. Musgrave, E, Troughton); Burleigh Heads N.P., SEQ, vine thicket (4, QMMO6157, QMMO6160, Oct 1976, M.J. Bishop); Terania Ck, NENSW, NVF, under piece of wood among rocks (1, QMMO17323, Aug 1987, J. Stanisic); Richmond Rv., NSW (1, AMC142448), DIAGNOSIS Shell small, trochoid, diameter 4.81—5.32 mm (mean 5.07 mm) with about 4 1/8- normally coiled whorls, last whorl only slightly descend- ing. Apex and spire strongly elevated, SP/BWW ratio 0.39-0.45 (mean 0.42). height 3.32-3.41 mm (mean 3.37). H/D ratio 0.64-0.69 (mean 0.67). Protoconch exsert, slightly shiny with | 40 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 4] lmm FIG. 21. Mussonula fallax sp. noy, Teramia Creek. NSW. QMMO17323, paratype. Palbal cavity. Scale line as marked. ]/2-1 5/8- whorls, mean diameter 1 149.5j.m at 1 1/2 whorls. Apical sculpture of 20-25 inter- rupted spiral cords with numerous, low, curved radial ridges forming toward the end of the protoconch (Fig. 20f), Post nuclear sculpture of numerous, high, protractively sinuated radial ribs, with periostracal blades which are folded toward the apex (Pig. 20g). No microriblets. Microsculptute consisting of vague incised spiral lines (Fig. 20g), Sculpture continued on the base (Fig. 20b). Rib interstices variable, 5-15 times the width of the ribs. Umbilicus narrow, V-shaped, diameter 4.51-5.32 mm (mean 5.07 mm), D/U ratio 5.86-6.18 (mean 6.02). Sutures impressed. Whorls rounded above and below a strongly angulate periphery. Lip simple. columella dilated and slightly twisted toward the umbilicus. Aperture subquadrate. Parietal callus strongly developed. Colour brown, often covered with dirt particles, Based on 2 measured adults. Foot and tail short and broad, bluntly tapering posteriorly. Body colour dark with sides of foot and outer margins of the sole purple-grey, Upper body and neck black. Ommatophores purple- FIG. 20. Mussonula fallax sp. nov, a, No. | scrub at Byangum, NSW, AM(C63870, holotype; d-e, Terania Ck, NSW. QMMO17323, paratype; fh, Burleigh Heads N.P., SEQ. QMMO 6157, paratype. a—c, entire shell; d, central and lateral 1eeth: e, mar- ginal teeth: [, apical sculpture; g. post nuclear sculp- ture; h, details of post nuclear microsculpture, Scale lines as marked. grey. Pallial roof black. Pallial cavity occupying c. 1/2 whorl apically. Mantle collar well developed with large glandular zone extending onto the Jung roof. Kidney with triangular pericardial lobe and rectal lobe present as a tiny vestige abutting hindgut. Ureter sigmurethrous without complete secondary branch. Ureterie pore situuted adjacent to the hindgut. anterior to the small rectal lobe of the kidney (Fig. 21). Reproductive organs immature. Central tooth of radula (Fig. 20d,e) with lan- cealate mesocone and small. pointed ectocones; inner Jaterals with endocone absent and more prominem ectocone. Outer laterals and mar- ginals typical, Based on one dissected subadult (QM- MO17323). RANGE AND HABITAT M. fallax inhabits the notophyll vine forests of the moist foothills af coastal northern NSW, and the unusual coastal vine thicket at Burleigh Heads, SEQ, Its microhabitat is not fully known. REMARKS In the absence of dala on the terminal genitalia the two species are separated on the difference in protoconch sculpture. Habitat destruction, particularly in the Big Serub area of north-east- ern NSW, has had significant effects on the distribution of M. fallax. Iredale (1941b, p. 2) cited specimens of M. verax from the Richmond River, narthern NSW. These are M. fallax. 154°= FIG,22. Distribution of Mussonula fallax sp. nov. 42 MEMOIRS OF THE QUEENSLAND MUSEUM Hedleyoconcha Pilsbry, 1893 Hedleyoconcha Pilsbry, 1893, p.18; Iredale, 1937a, p. 321; Iredale, 1937b, p. 13; Iredale, 1941a, p. 264; Iredale, 1942, p. 35; Iredale, 1944, p. 317; Burch 1976, p.133. TYPE SPECIES Helix delta Pfeiffer, 1857; by original desig- nation. PREVIOUS STUDIES Pilsbry (1893) placed Hedleyoconcha in the Flammulinidae where it remained until Iredale (1942) erected the Hedleyoconchidae for it. Solem (1983) included Hedleyoconchidae with his expanded Charopidae. I follow Solem and consider that Hedleyoconcha can be derived from more generalised charopids. DIAGNOSIS Shell relatively large, diameter 5.38- 10.08 mm with 5 1/8 to 6 5/8 whorls. Spire and apex strong- ly elevated (Figs 23c, 26c), height of shell 4.62- 6.47 mm, last whorl descending. Protoconch of 1 1/2 to 1 3/4 whorls, pitted (Fig. 23e). Post nuclear sculpture of low, protractively sinuated, radial ribs and low rounded spiral cords in a reticulate pattern (Figs 23f, 26d). Sculpture al- tered after 4 whorls, continuous on base but much less pronounced (Figs 23b, 26b). Um- bilicus narrow, partially obscured by the reflected columella. Sutures lightly impressed. Whorls slightly flattened above and rounded below a strongly angulate to thread-carinate periphery. Aperture ovately lunate. Lip simple, columella dilated and slightly reflected over the umbilicus. Colour straw to cream-yellow with a darker spire. Periostracum deciduous. Foot and tail broad, rounded posteriorly. Caudal horn present. Colour creamy-white with some pigmentation on the pallial roof. Om- matophores black. Kidney elongate, vaguely tri- angular with a reduced rectal lobe. Ureter sigmurethrous with an incomplete secondary branch. Ureteric opening in the angle between the kidney lobes. Ovotestis several clumps of palmately clavate lobes of alveoli, oriented at tight angles to the plane of coiling. Terminal male genitalia with thin vas deferens and large, reflexed, muscular epiphallus, entering penis through a large fleshy pilaster opposite a grooved muscular thickening which acts as a collar. Penis proper with long slender corrugated pilasters. Penial retractor muscle inserted at the penis-epiphallus junction. Radula with tricuspid central and lateral teeth which have long mesocones, short endo- and ectocones, and well- developed anterior flares; marginals typical. DISTRIBUTION AND ECOLOGY Hedleyoconcha has a classic relict distribution with a widespread species in northern NSW and southern Queensland, an altitudinally restricted species on the summit of Mt Bellenden-Ker in north Queensland and a further species on Lord Howe Island, off the coast of New South Wales. Although similar distribution patterns are seen among some flightless groups of insects such as the Peloridiidae (Monteith, 1980) this is the first time a land snail group has been identified as having such an unusual and biogeographically complex distribution. COMPARISONS The high spire, punctate apical sculpture (Fig. 23e) and distinctive low reticulate post nuclear sculpture (Figs 23f, 26d) distinguish Hed- leyoconcha from other subtropical and tropical charopids. Oreokera Stanisic, 1987 has an elevated spire but apical sculpture of spiral cords, prominent post nuclear radial ribs and fine, reticulate microsculpture (Stanisic, 1987, pls 1,2). Mussonula is much smaller and has prominent apical spiral cords (Figs 18d, 20f) and crowded, high, curved radial ribs on the post nuclear whorls (Figs 18e, 20g). Ngairea lacks the punctate apical sculpture seen in Hed- leyoconcha and has much finer post apical sculp- ture (Figs 3e, 6e, 9g, 12e, 15f). The species show minor differences in shell features but are readily segregated on the basis of geography. KEY TO SPECIES OF HEDLEYOCONCHA 1.Shell with umbilicus wider (Fig. 26g), mean D/U ratio 4.60; Lord Howe Id ......... addita Shell with umbilicus narrower, mean D/U ralio greater than 6.50 v.cccecseeeseseeeees 2 2.Shell with strong reticulate sculpture on early whorls (Fig. 26d): protoconch weakly punctate (Fig. 26d); Mt Bellenden Ker, NEQ rigthivrcaneislopeardinetebige sevh teasannenegusgneey sedi ailaketoae SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 43 Shell with weaker reticulate sculpture on early whorls (Fig. 231); protoconch strongly punctate (Fig. 23e): southern Queensland and northern New South Wales y.c.u esses della Hedleyoconcha delta (Pfeiffer, 1857) (Figs 23-25; Tables 2, 6) Helix delta Pfeiffer, 1857, p. 386 Helix conoidea Cox, 1864, p. 21; non Drapurnaud, 1801. Helix fenestrata Cox, 1866, p.a74: non Sowerby. 1841. Charopa (Thalassia) delta (Pieifter): Tryon, 1886, p. 215. pl. 63, fig, 76, Flammulina (Hedleyaconcha) delta (Pfeiffer): Pilsbry, 1893, pp. 18-19. Hedleyacancha delta (Pfeiffer), twedale, 1937a, p, 321; Iredale, 1941a, p. 35. Hedlevoconcha duona Iredale, 1937b, p, 13, pl, fig. 13: Iredale, 19414. p. 35, fig. ¥. COMPARISONS H. delta can be distinguished from H. ailaketoae by its larger size, less prominentadult sculpture and more conspicuous apical sculp- ture. Anatomically the differences are minor with slightly more corrugated penial pilasters and a more prominent apical penial bulb present in H. ailaketoae (Fig. 27a-b). H. della is not casily confused with any sympatric charopids. Mussonula verax (Fig. 18). M. fallax and Ngairea levicostata all have elevated shells and occur in the NSW-QLD border areas. However they have prominent, fine, crowded radial ribs on the adult shell whorls (Figs 6e, 18e, 20g). spiral cords (with radials in N. levicosiata).on the pratoconch (Figs 6d, 18d, 20f), are smaller, and fee a less prominent peripheral keel (Figs 6c, 19c. 20¢). Ngairea murphyi resembles A, delta in gross conchological detail but is smaller, with more tightly coiled whorls, less elevated spire,, more open umbilicus, finer adult sculpture and apical sculpture of fine pits and wrinkles (Fig. Gab, e-g). H. delia is microsympatric with several helicinids - Pleuropoma jana (Iredale, 1937) from the Port Macquarie area, NSW and P. draytonensis (Pfeiffer, 1857) from far northern NSW and southern Queensland. Both species have elevated shells similar in colour to #7. delta and live on the leaves of low shrubs. However they may be distinguished in having strongly incised spiral grooves on the protoconch and adult whorls, and no umbilicus, The introduced and widespread Pacific helicarionid Coneuplec- ta calculosa (Gould, 1852) is also microsym- Patric with H, delta in parts of northem NSW and southern Queensland but it is much smaller, transparent, yellow-hor in colour, and has. a ecceperstip ie of numerous fine incised spiral lines, PREVIOUS STUDIES H. delia was originally described from the Drayton Runge (= Toowoomba), SEQ. Cox (1864, 1866) introduced names for two NSW populations (conaidea and fenestrata) which he later (Cox. 1868) synonymised with H. delta. Iredale (19374) placed them in synonymy with Helix scandens Cox, 1872, a helicarionid, then (1937b) realised the error but considered the NSW specimens specifically separate as H. duona on the basis of their broader base, nar- rower umbilicus and weaker sculpture. In this study specimens of Hedleyoconcha were dissected from numerous localities (including Wentworth Falls, west of Sydney. NSW) and no perceptible differences in genital anatomy were noted. Differences in shell size and sculpture are mosaic and fall within the range of a single widely distributed species. Hence Iredale’s separation is considered unsustainable and H. duona is placed in synonymy with J. delta. TYPE MATERIAL LecToTyre, BMNH1982246, Drayton Range, North Austrilia. Collected by Mr Stutchbury. ex H. Cuming collection, Height of shell 6.64 mm. diameter 7.56 mm, H/D rativ 0.88. D/U raliv 7.67. whorls 6 1/4, PARALECTOTYPFS: BMNH1982246, 2 specimens, same collection data as lectotype. HOLOTYPE of ditona: AMC63526. Terrigal, near Gosford. NSW. No other dala, Height of shell 6.30) mm, diameter §40 mm. H/D ratio 0.75, D/U ratio 10.24, whorts 6. OTHER MATFRIAl NEW SOUTH WALES: Frank’s land, Terrigal, Gos- ford. in serub, on trees (2. AMC142450. S. W, Jack- son, 23 Mar 19/3); Dungog $.F,. among orchid roats on fullen tree (lL, AMC142455, W. Dowling, 19 Jun 1962): Lismore (1, AMC142458, Lower): In valley down Wentworth Falls, rainforest by creek (1, AM- C142453, D.K, McAlpine): Sassafras Gully, Springwoud (2, AMC 142452, C.F, McLauchlun, 8 May 1948): Dorrigo Scrubs, on underside of leaves on trees (2, AMC142454. Dec 1910, S'W. Jackson); Byron Bay serubs (7, AMC 142457, 1904, S.W, Jack- son): 1 1.4km § of Nymboida, on Grafton-Armidale MEMOIRS OF THE QUEENSLAND MUSEUM 44 2mm SYSTEMATICS OF AUSTRALIAN CHAROPIDAE FIG. 24. Hedleyoconcha delta (Pfeiffer, 1857). Dandabah, Bunya Mts, SEQ, OMMO6068. a, genitalia; b, details of penis/epiphallus region; c, details of penis interior; d, hermaphroditic duct; e, ovotestis; f, talon; g, pallial cavity. FIG 23. Hedleyoconcha delta (Pfeiffer, 1857). a-c, Drayton Range, North Australia.) BMNH1982246, lectotype: d, Terrigal, near Gosford, NSW. AMC63526, holotype of duona; e—h, Dandabah, Bunya Mts, SEQ. QMM0O6063. a-d, entire shells; e, apical sculpture; f, post nuclear sculpture; g, central and lateral teeth; h, marginal teeth. Scale line as marked. MEMOIRS OF THE QUEENSLAND MUSEUM 46 cst O0'b OR Ly6 (es'x-zO's) SSTUFRT'R (86°6-€L'R) STP OFITG creat) o's 0L'6 ROU (onnl-19) HLPOHPER (60'6-$$'9) ERLUFOO'R Lvl OlLvY n/a 70 ce) (eo 26°0) GEO OFA (z6'0-2L'0) OOVOFTR'D zRO RRO (RL'0-240) OEOUFSLO wo 890 (01-260) 870'0700'1 (pS 1-76'0) LOTUFELL 660 (wu) HLLGIA “Worn WHW/StIRI Sard ra) on 50 £90 (z9°0-09'0) O07 190 (¥2°0-98'0) OOL0F PIO 670 #90 (E9°0-05°0) SworLs0 (89°0-65°0) 920'0F P90 LL0 (820-890) 1en0FzL'0 (88'0-49°0) #LO0FZLO 90 OLLVY AAME/dS Ltt rare Liz LUZ (p6'7-98'7) OPO0F06'7 (19'7-Se-2) O€ POFRET 697 197% (697-287) SRUOFI9Z (g'7-Se'O) OLOOFRET raya (697-197) £20097 (69°7-SEO) ENN 0FSSZ £Lt (wu) HLGIM “TNOHM ACOA Ist trl sel er Lut (g6'1-97'1) SEE OFOTT set Lut (o9'1-se1) STLOFRF'I (09 -€e'D GPOOFIS'T £61 (ovz-Le) ILOOF16) (sez-18 1) 9STOFSR'I 61Z (uu) NOISNYLOUd auldS 420 6L0 LL0 sz0 (620-820) SOO 0F6LO (9L'0-€2'0) sioorszo seo 6L0 (EL°0-ZL'0) SUMOFEL'O {es'0-18'0) 90007780 8L0 (ex'0-€L'0) IZO'OFRL'O (62'0-1L'0) LEOOF PLO Bg0 OLLV¥ G/H (40NVa GNV Was ‘NvaW) phot TIVGaM VIC VHONODOATIGSH ONY ‘AON ‘AS TVOLINVIUVY VHINODOAT 1TH (L881 ‘WASAAdd) VLTIG WHINOIOATIGIH NI NOLLVRAWA TWOOT- 9 TTAVL #69 68'9 LoS 189 $2'8 (L0°8-68"9) 06S 08F'L OF an (95'L-26°9) O6ZOFLTL (L¥9-27'9) PL0'0F9E'9 SOL (ip6-ze'L) ELE'OF0E'R (80°01-#6'L) Zp9'0378'8 ook (wa) YALAWVIG RED us wor ers (Le'9-6£°9) OPpOOF EPS (eV9-v0's) SPSOF 65'S 0c9 9's (sos-r0's) SSTOFOE'S (6z's-£1's) FOF 17'S Les (689-779) 6Pl FLED (g6"L-£9"S) £ELOFES'9 wy +H1S (z/1 9-4/1 9) we9 (p/1 901-R/L S) +9 9 (+8/1 9 9) +9 +B/LS veo (+R/€ 9- 9) 19 (8/5 9 OI-B/L S) +B/19 He STYOHM favds! ONY (adAjoya7q) RRREY OWY “SP IMO} PA] puppy pyuor0ka)pe]] zosst ONWO (ad Kioyo}4) $SZ11 ONWO JI wapuryjog IW apojaynpIn DYIWOIOAA|PA}] nq = SNAWIDadS: AO WaAIN 0se9z OWV SUPA UHOMIUDAA, OStZh1 OWV 9zS£9 DWV qetinay, £6491 ONWO auenboeyw uo £8L91 ONWO “Wa Rd JUxXTUR 9979 OWWO aduey puowysy 1601 ONWO “a's (aneoq s6zZ1 ONWO #909 OAWO STW BAUng, (ad4yopaq) OFT7ZR6I HNL adury uojAtig Dyap DY MOIOAa|paH aIWYN SYSTEMATICS OF AUSTRALIAN CHARGPIDAE 47 rh?s is s9°S 315s er a 32°S xc SSeS 33°8 a4*a \\\ > 800m 149° 0 SO*E = o1S2"E | 1SO"H | ASSFE | ASASE FIG, 25. Distribution ol Medlevecuncha delia (Pfeil fer, ISS7), Ru, in rainforest beside road (1, AMC 142456, 20 May 1976. PH. Colman & 1, Loch); Wilson River Reserve. via Bellangry (1, AMC142451, 25 Sept 1981, D.K. McAlpive): Richmond Rv (1, AMCI42461, Cox coll); Port Stephens, on island in port (7..AMC63730, Oct 1892, ex Bland, ex Cox); Port Macquarie (4, AMC36565, ex Hedley), Col’s Cave, Carrai’S.T., in liter near entrance (1, AMC121693, 20 Feb 1980, PH, Colman); Carrai Cave, Carrai S.F., in litter vine forest (8. AMC121701, ‘foonumbar Forest Road c. 34 km from Kyogle, ‘Toonumbuar S.F.. CNVF (28°33°8, 152°45°E) (1, AMCI28501. 15 Mar 1981, AM/QM - ABRS). Newell Falls, Dorrigo NVP. (3(°24'S, 152°45°E) (1, AMC128337, 12 Mar 1981, ABRS ~ AM/QM), Acacia Plateau Road, ¢, 12.2 km from Acacia Ck-Kil- larney Road. Koreelah $.F., SEVT (28°217S, 152°24°E) (3, AMC128562, OMMO10960, 15 Mar 21 Feb 1/980, PH, Colman): 1981, AM/OM - ABRS); Mt Warning N.P. (28°24°S, 153°16°E) (1, AMC129290, 18 Mar 1981, AM/OM - eres Tooloom Scrub, Beaury $.F., NVF_(28°29"S, $2°24 E) (6, AMC128538. QMMO10941, 15 Mar oy AM/QM - ABRS); Wiangarie 8.F.,¢. 25 km NE Kyogle. 1000 m (28°23°S, 153°06'E) (2, AMC142480, 17 May 1976, P.H. Colman, |. Loch); Koreelah Ck, Beaury S.F..c. 15 km W of Urbenville, 530 m (28°21°S. 152°20°E) (3, AMC 142477, 18 May 1976, PH. Colman, 1. Loch); Cherry Tree North §.F., c. 40 km W of Casino, 400 mo (28°54°S, 152°45°E) (24, AMC142478. 19 May 1976, PH. Colman, I. Loch); Beaury S.F.. ec. 1S km W of Urbenville, 810 m (28°27 8, 152"24°E) (17, AMC 142479, 18 May 1976, PH, Colman, 1, Loch); O°Donnell Ck, nr Eden Creek. 15 km N of Kyogle (1, AMC131453, 13 May 1981. N. Moulds, C. Smithers); Ounmbali Serubs. nr Gas- ford (1. AMC 152163, Oet 1904, S.W, fucksun); New- castle (1, AMC 152162, 1404, S.-W. Jackson); Moonie Moonie Creek, ne Gustord (3. AMC142933, on tree trunk in cainforest, G6 Dee 1979. B, Day; |, AMC142931. 26Jan 1984. D. McAlpine): Wentworth Falls, Blue Mountains (2, AMC142938. 12 Oct 1965, DK. McAlpine): Gibraltar Range (1. QMMO17256, 10 Nov 1980, G.B, Monteith; 2, QMMO17257, If Nov 1980, R. Raven); Booyong Scrub. via Lismore (28°49°S, 153°30°E) (1, OMMOQ6337, 14 May 1977, M.J, Bishop); Richmond Range S.F. (28°41 8, 152°44°E) NVE (17, OMMO6266, 19 Apr 1976, MJ. Bishop); Upper Wilson's Creek (28°32'S, 153°24"E), in litter (1, OMMO16791, 16 Nov 19%3, AM/QM — ABRS); Bellingen River (30°27°S, 152°32°E), rain- forest in gully (1, QMMOQ10835, Mar 1981, AM/QM - ABRS); Wentworth Falls (2, AMC26350, ex C. Laseron) Sherrard Falls, Dorngo, 600 m, rainforest, in litter (1, OMMO16777,, 13 Nov 1983, AM/QM - ABRS); Swans Rd, Bruxner Park Flora Reserve, litter, NVF (3 QMMO16783, 14 Nov 1983. AM/OM - ABRS):; Whian Whiun $.F., ¢. 7 km Sof Dorroughby, NVF (1, OMMOIG788, 15 Nov 1983, AM/OM - ABRS): Sea Acres Nature Reserve, Port Macquaric, littoral rainforest with palms (1, OMMQ16793, 9 Dec ORS. PHL Colman): Bytrill Creek Tributary, on Meb- bin forest roud (28°27'S, 153°12°B) (1, OMMO1(407, Mar 1981, AM/QM - ABRS),. QUEENSLAND: Near monument, Cunningham Gap NPL, NVF (28°04°S, 152°24°E) (1. AMC128611. 16 Mur 1981, AM/QM-ABRS); Bunya Mountains N.P.. (1, AMC142464, 8 Feb 1961, D.P. McMichael): Cooluhunia pine scrubs, nr Ringaroy,.on pine trunks (2, OMC 142460, Jun L908, SW. Jackson; 5, AM- C142402, Jun 1908,ex Helms, S.W. Jackson); Laitle Nerang Creek (3, AMCS35505, ex Brazier); Dandabah, Bunya Mis (26, AMC 136617, OMMO12295, 31 Sept 1982, AM/QOM-ABRS); Halls Plain, E of Emu Vale, 48 MEMOIRS OF THE QUEENSLAND MUSEUM CNVF, crawling on log (1, QMMO16898, 4 May 1986. J. Stanisic, J. Chaseling); Binna Burra, Lamington N.P., crawling on ‘cordylines’ (4, QM- MO14976. 10 Mar 1986, D.G. & N.G. Potter): Mt Hobwee. Lamington N.P. (28°13°S, 153°13°E), 1150 m. MFF. (1, QMMO6077, Apr 1976, M.J. Bishop): Toowoomba (27°34'S, 151°57°E) rainforest, (1, QM- M06276, 25 Sept 1976, R. Raven); Dandabah, Bunya Mts (26°53°S, 151°35°E) Araucaria/NVF (90, QM- MO6068. FMNH 206312, 5 Mar 1976, M.J. Bishop): Upper Pine Creek, via Canungra, NVF/Araucaria, litter (2, QMMO16541, 1 Oct 1986, J. Stanisic, D. Potter): Tullawallal Circuit, Binna Burra, Lamington N.P., NVF, under logs (1, QMMO16736, 2 Oct 1985, J. Stanisic, D. Potter, J. Chaseling); Dandabah, Bunya Mts, behind picnic area (26°53’S, 151°36°E) (11, AM- C136617, 31 Sept 1982, AM/QM-ABRS). DIAGNOSIS Shell trochoid (Fig. 23c), diameter 6.22—10.08 mm (mean 7.77 mm) with 4 5/8—6 5/8 (mean 6 1/8+) normally coiled whorls, last whorl des- cending. Apex and spire strongly elevated. SP/BWW ratio 0.36-0.88 (mean 0.66), height 5.04-7.98 mm (mean 6.01 mm), H/D ratio 0.71- 0.88 (mean 0.78). Protoconch of 1 1/2-1 5/8 whorls, mean diameter 875.0u.m at | 1/2 whorls. Apical sculpture of irregular pits and vague radial rugosities (Fig. 23e). Postnuclear sculp- ture of low spiral thickenings crossed by numerous low protractively sinuated radial ribs in a reticulate pattern (Fig. 23f), becoming less pronounced after the first four whorls (Fig. 23a). Sculpture continued on the base but reduced (Fig. 23b). No additional microsculpture. Um- bilicus small, diameter 0.62—1.54 mm (mean 0.89) D/U ratio 6.55—11.28 (mean 9.01), partial- ly obscured by the reflected columella. Sutures lightly impressed. Whorls rounded above and below a strongly angulate to thread-carinate periphery. Aperture ovately lunate. Lip simple, columella dilated and reflected over the um- bilicus. Parietal callus well developed. Periostracum strongly deciduous. Colour (with periostracum) creamy-yellow: (without) whitish-pink, often with a darker brown to pink apex and spire. Based on 20 measured adults. Genitalia with epiphallus (Fig. 24b) reflexing before entering the penis apically through a fleshy pilaster (Fig. 24c). Penial retractor muscle insert- ing adjacent to the penis/epiphallus junction. Penis (Fig. 24c) long and slender, muscular, with an apical bulb containing a grooved, muscular thickening. Penis proper with spongy lon- gitudinal pilasters. Free oviduct thick, mus- cularised, longer than vagina (Fig. 24a). Vagina short. internally with longitudinal pilasters. Radula (Fig. 23g, h) with tricuspid central and lateral teeth which have long lanceolate mesocones and well developed anterior flares. Based on 6 dissected specimens (QM- M06068, FMNH206312). RANGE AND HABITAT H. delta inhabits cool subtropical notophyll vine forests of the Great Dividing Range from Dorrigo, NSW to the Bunya Mountains, SEQ; the humid warm notophyll vine forests of the coastal region of northern NSW; warm temperate rainforests in the Gosford and Blue Mountains area of central NSW; and some mar- ginally drier subtropical forest (microphyll vine forest, semi-evergreen vine thicket) in southern Queensland. The distribution of H. delta at the southern end of its range is not well known. REMARKS H. delta has a caudal horn. This structure is typical of some New Zealand arboreal charopids and is common in Australian helicarionids. The shell of H. delta has an apical portion of about 4 whorls in which the sculptural features are quite prominent, but on the remaining whorls sculpture is almost absent. A similar pattern is seen in H. ailaketoae. This is not gerontic growth and its significance, if any, is unknown. Hedleyoconcha ailaketoae sp. nov. (Fig. 26a—e; Tables 2, 6) ETYMOLOGY For Dr Aila Keto in recognition of her efforts for rainforest conservation. COMPARISONS H. ailaketoae is distinguished from H. delta by its smaller size, fewer whorls, less elevated shell, weaker apical sculpture, and more prominent reticulate sculpture on postnuclear whorls (Fig. 26a-d), The sympatric Oreokera cumulus (Odhner, 1917) has an elevated. keeled shell but is larger, has fewer whorls, apical spiral cords, FIG. 26. a-e, Hedleyoconcha ailaketoae sp.nov. a-c, Summit. Mt. Bellenden Ker, NEQ. QMMO11254. holotype: d-e, Mt. Bellenden Ker, NEQ, QMMO1S802. QMMO11273, paratypes. a-c, whole shell: d. apical-postapical sculpture: e, central and lateral teeth. f-h, Hedlevoconcha addita Iredale, 1944, AMC63488, lectotype. 49 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 50 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 27. Hedleyoconcha ailaketoae sp.nov. Mt. Bellenden Ker, NEQ. QMMO15802, paratype. a, genitalia; b, details of penis interior; c, talon; d, ovotestis; e, hermaphroditic duct; f, pallial cavity. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 31 Bell Pesk North r] Ss Edmonson Sy NIN a ee. Pel lenden Nt Bab ince RRA Wa EN . fercle Frere Smuth Peck \ Malt. kd S 145" Aste ho? ADE FIG. 28. Distribution of Hedleyaconcha ailaketoae sp, nov. and postnuclear sculpture of prominent radial ribs with reticulate micrasculpture (Slanisic, 1987, pls 1.2). TYPE MATERIAL HoLotyPr: OMMQO11254, Mi Bellenden Ker. NEO. Summit. 1560) m. SMVFT. Collected Earthwatch/- OM, 17-24 Oct 198). Height of shell 5.13 mm. diameter 6.8] mm, H/D ratio U.75. D/U ratio 9,47, whorls 5 /s+. PARATYPES: Mt Bellenden Ker, NEQ. Summit TV stn. 1560 m (17° 16'S, 145°51°E) rainforest (2, OM- MO1S802, 29 Apr 1983, G.B. Monteith, D. Yeates): Mi Bellenden Ker, NEQ. summit, 1560 m. SMVFT (1. OMMO1 1273, 1-7 Nov 198), Earthwarch/QM). DIAGNOSIS Shell trochoid (Fig. 26c), diameter 5.97-6.81 mm (mean 6.39 mm) with 5 1/8 normally coiled whorls. Apex and spire strongly elevated, SP/BWW ratio 0.54-0.63 (mean (0.61), height 4.62-6.15 mm (mean 4.88 mm). H/D ratio 0.75— 0.77 (mean 0.76). Protoconch 1 1/2 normally coiled whorls, mean diameter 965p.m at 1 1/2 whorls, Apical sculpture of weak irregular pits (Fig, 26d). Postnuclear sculpture of prominent, spiral cords crossed by crowded, protractively simuated radial ribs in a strongly reticulate pat- tern Which becomes less pronounced after c. 4 whorls (Fig, 26b). Sculpture continuous on the hase but somewhat reduced. No additional microsculpture. Umbilicus very small. diameter (72mm, D/U ratio 8.30-9.47 (mean 8.89), par- tially covered by columella. Sutures lightly im- pressed. Whorls rounded above and below a thread-carinate periphery. Aperture ovately lunate. Lip simple, columella dilated and reflected over the umbilicus. Parietal callus well- developed. Periostracum deciduous, Colour (with periostracum) straw yellow, spire darker. Based on 2 measured adults. Epiphallus (Fig. 27a) Jong. entering penis api- cally through a fleshy circular pad. Penis (Fig. 27b) shorter than epiphallus, slender with cor- rugaled, longitudinal pilasters (Fig. 27b), apical- ly with a penial bulb containing entrance of epiphallus (Fig. 27b) and a muscular grooved collar. Penial retractor muscle inserted at the penial bulb/epiphallus junction. Free oviduct muscularised, much longer than vagina (Fig. 27a). Vagina short with longitudinal pilasters internally. Radula (Fig. 23¢) with tricuspid central and lateral teeth which have long mesocones and strongly developed anterior flares, No data avail- able on marginal teeth. Based on 1 dissected adult (QMMO15802). RANGE WWD HABITAT H. ailaketode has only been found to date in simple microphyll vine fern thickets and forests on the summit of Mt. Bellenden Ker, NEQ. Its microhabitat is unknown. REMARKS Discovery of H. ailaketoae on Mt. Bellenden Ker was unusual considering that its neurest relative is almost 1400 km to the south. Pre- viously Stanisic (1987) reported on the al- titudinally restricted charopids from this region. Hedlevoconcha addita Iredale, 1944 (Fig. 26f-h; Tables 2. 4) Hedlevoconcha addita Iredale. 1944, p. 317, pl. 19. fig, 19 COMPARISONS In general appearance, H. addita bears some resemblance to H. delta. The two type specimens are Worn and only a few traces of vaguely reticu- late adult sculpture are visible, However, reticu- late postnuclear sculpture is not sufficient to conclusively link this species with the two main- land congeners, The keel of . addita is much more pronounced than in either H. delta or H, ailaketoae and may indicate that the overall similarity of 47. addira lo these species is conver- a MEMOIRS OF THE QUEENSLAND MUSEUM gent. but in the apsence of other evidence, the original allocation of fredale (1944) is retained. TYPE MATERIAL Lectotyre: AMC63488. Mt Gower, SSE, Lord Howe Island, PARALECTOTYPE: AMC 150093, same collecuon dats as lectolype. DIAGNOSIS Shell moderately large, trochoid (Fig. 26g), diameter 6.89—6,98 mm (mean 6.94 mm) with 6 to 6 1/8 (mean 6+) normally coiled whorls, the last whorl descending slightly. Apex and spire strongly elevated, SP/BWW ratio 0.63—0.67 (mean 0.65), height 5.38-5.46 mm (mean 5.42 mm). H/D ratio 0.77-0.79 (mean 0.78). Protoconch of | 3/4 whorls, mean diameter §82.52m1 at 1 1/2 whorls, Apical sculpture worn on avatlable specimens. Post nuclear sculpture also mostly worn. Body whorl with remnant patches of periostracum and sculpture consisting of low crowded spiral cords and weak protrac- tively sinuated radial growth ridges. Sculpture continuous on the base as crowded spiral cords and weak protractively sinuated radial growth ridges (Fig. 26g). No additional microsculpture visible. Umbilicus narrow. V-shaped, only slightly covered by the columella (Fig. 26h), diumeter |.48—1.54 mm (mean 1.51 mm). D/U ratio 4.53466 (mean 4,60). Sutures only slight ly impressed. Whorls rounded above and below a strongly carinale periphery. Aperture quadrate to roundly lunate. Lip simple, columella dilated and partially reflected over the umbilicus. Paric- tal callus developed. Periostracum deciduous. Colour (with periostracum) straw yellow, (without periostracum) chalky whitish-pink, With 4 darker spire. Based on 2 measured adults. Anatomy unknown, Lenwebbia gen, nov, ERY MOLOGY Vor Dr Len Webb in recogmuon of his pioneer- ig studies On rainforest evolution. TYPE SERIES Fommebhin protuservbienlata sp, ny, DI\GNONS Shell senatl, moderately elevated (hig. 29c) average divmeter 435 mm ind relatively few (avergye + L'S+) rounded whorls. Protoconeh sculpture of numerous dimples and vague radial ridges (Fig. 29g). Adult sculpture of spiral grooves, broad radial undulations and very fine thread-like radial ribs (Fig. 29e.f), Base with incised spiral lines and weak, raised radial growth lines. Sutures impressed. Umbilicus small (Fig. 29b). Lip simple, columella dilated and deflected toward the umbilicus. Poot and tail broad. bluntly tapering posterior- Iv. No caudal horn or foss. Body colour (in preservative) grey. Ommutophores black, Pallial root with some black speckling. Kidney orange, triangular, c. 1/2 length of the pallial cavity. Rectal kidney lobe reduced. Ureter sig- murethrous with am incomplete secondary arm and reflexed primary arm (Fig. 30e). Ureteric opening located al the rear of the pallial cavity in the angle between the kidney and the hindgut. Ovotestis (Fig. 30c) several clumps of palmately clavate lobes of alveoli, oriented at nght angles to the plane of coiling, Terminal male genitalia with short muscular epiphallus (Fig. 30b) enter- ing penis apically through a simple pore (Fig. 30b). Penis elongate, internally with few, large longitudinal pilasters (Fig. 30b). Free oviduct (Fig. 30a) longer than vagina, otherwise typical. Radula (Fig. 29d) with mesocone of central and lateral teeth long, lanecolate. Marginal teeth typical, COMPARISONS Lenwebbia differs from Neairea in its smaller size, more weakly sculptured shell and more strongly pitted upex (Fig, 29g), Anatomically, the retlexed primary ureter (Fig. 30e) and bright- lv coloured kidney of Lenwebbjq are distinctive. In the Mudlo Gap area, SEQ, Lenmwebhia is sym- patric with Caenocharopa whose worn shells are grossly similar, However Coenocharopa has an apes with curved radials and taised spiral cords (Figs }11t, 114. 120f-n), adult sculpture of fine radials and spiral cords (Figs | 11g. 114g, 120d), and more shouldered whorls. The sympatric helicarionid Tareevstiy tredale. 1937, has a protoconch with incised spiral lines and adult sculpture of incised spiral grooves which have egular notches along their length. Lenwebbia protoscrobiculata sp, nov, (Figs 29-31: Tables 2, 7) ETYMOLOGY Latin serobiculus. a litthe diteh; for the punctate protoconch. PAULL 7 LOCAL VATUIATION IN LEN WEHHID PROTOSCHUMICULA TA SPONOV, (MULAN, SUM AND KANGH) SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 53 COMPARISONS L, protoscrabiculata most closely resembles Ngatrea canaliculata which is sympatric with the former in the northern parts of its range. N. canaliculata lacks the punctate protoconch of L. pratescrobiculata, has prominent post nuclear radial ribs (Fig. 12a), channelled sutures (Fig. 12e) and flammulated shell. In the southern part of its range L. protoscrobiculata is sympatric with Coenocharopa macromphala and C. par- vicestata and may be confused with worn dead shells of these latter two species. However Coenocharopa has apical sculpture of curved radial ribs and low spiral cords and is smaller (Figs 114e-g; 120d-h). Anatomically a complete secondary ureter (Figs 115a, 121a) is sufficient to distinguish these species from L, protoscrobiculata, DIU RATIO $a SYR20, 126 (545-652) SR SAT 54 SAL (5,386.43) UMHILICAL (imum WIDTH OF OS O.7S,006 (0.02-0.70) (0.720.102) Wad nro (74 42 RADMM RIDS APA RAN 35 AN 2010 (Moa) (hits (a2 zt) 0.25 On nla TYPE MATERIAL HOLOTYPE: QMMO17048, c. 8 km from Kalpowar on Fireclay Rd, Kalpowar S.F.. SEQ (24°43'S, 151°21°E) leaf litter, MVF/Araucaria. Collected by J, Stanisic, A, Green, 15 July 1980, Height of shell 2.94 mm, diameter 4.49mm, H/D ratio 0.63, D/U ratio 5,71, whorls 4 3/8. PARATYPES; SOUTH EAST QUEENSLAND - QMMO17047. same data as holotype. 36 specimens; Mt Mudlo, Kilkivan S.F. (26°01°S, 152°13°E) leaf litter. NVF/Araucaria (34. QMMO8359, 17 Jul 1980, J. Stanisic, A. Green); Mudlo Gap, Kilkivan S.F., litter MVF/Araucaria (3, OMMO16843, 5 Jul 1984, J. Stanisic, D, Potter, K.Emberton); Mudlo Gap S.F. (26°01°S, 152°14°E) MVF/Araucaria (7, QM- MO12365. AMC1]3665, 2 Sept 1982, AM/QM- ABRS): Clifton Range S.F.. SW of Fairlies Knob, Araucaria/MVF (26°34’°S, 152°16'E) (2, OQMMO- 12386, AMC136681, 2 Sept 1982, AM/QM-ABRS),; Dawes Range. MVF/Araucaria (24°28°S, 151°07'E) (2, QMMQ12652, 4 Sept 1982, AM/QM-ABRS); c. 4 km below summit of Mt Fort William on Mt Fort William Rd, Kalpowar S.F., litter, under rocks, logs, MVEF/Araticaria (7, QMMO17010, OMMO16839, 3 Jul 1984, J. Stanisic, D, Potter, K. Emberton); rain- forest walk, Kalpowar S.F, (24°41°S, 151°21°E) MVF/Araucaria (1, QMMO12578, 4 Sept 1982, AM/QM-ABRS); base of Mt Woowoonga, (25°26°S, 152°06°E) MVF/Araucaria (5. QMMO12447, AMC136720, 3 Sept 1982, QM/QM-ABRS); Mt Goonaneman, via Childers, rainforest, 670 m, litter (2, QMMQ17258. 3-7 Nov 1980. R. Raven, V. Davies); Mt Goonaneman, rainforest (2, QMMO17259, 6 Feb 1981, G,B. Monteith); Mt Colosseum, 6 mls S of Miriam Vale (12. AMC 142928, 6 Jun 1964, L. Price); Kalpowar S.F., NE of Monto. SEQ (24°42’S, Dai (2.262. 2c (Lli-tay z a = ca i= o 3 wnt! (mm) 4 aay 3 4 (ape (0.30.55) bat oax ow Haren (UMA) Thoth PROTRUSION (im) (64 0007 Whnl-Oa7h ted ennor (.aL-u.07) HD RATIO thy? HAS on DIAMETER (omen oe) AAD EOD (4174.09) aay as ay AN ISK (Mit.74) 2M ITD (2hA-IS) 29H ate i (250-21) Bey (4 WKeld Whe) + Es 4 414 4dr alihe a Whe NUMINKK Ob SUL CIME NS Mi Winona MMO 12447 FE = = z > (MMO) | 2052 NAMI: = = Dawes Runge Jenwodehin 54 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 53 cos SO Er FIG. 30. Lenwebbia protoscrobiculata sp. nov..c. 4km below summit of Mt. Fort William on Mt. Fort William Rd, Kalpowar S.F., SEQ. QMMO17010, paratype. a, genitalia: b, details of penis interior; c, structure of ovotestis; d, details of hermaphroditic duct and talon; e, pallial cavity. Scale lines as marked. FIG. 29. Lenwebbia protoscrobiculata sp.nov. a—c, c. 8km from Kalpowar on Fireclay Rd, Kalpowar S.F., SEQ. QMMO17048, holotype; d, Summit of Mt Fort William, Kalpowar S.F., SEQ. QMMO17010, paratype; e-g, Mt Mudlo, Kilkivan S.F., SEQ. QMMO8359, paratype. 4c, entire shell; d, lateral and marginal teeth; e, early post nuclear sculpture; f, late postnuclear sculpture: g, apical sculpture. Scale lines as marked. aH MEMOIRS OF THE QUEENSLAND MUSEUM 24°S 25°S Kingaro 1 XSAN fSjait. > 500m 150°E 151°E 152°E 153°= FIG. 31. Distribution of Lenwebbia pratoscrebiculata Sp. Nov. 151°21°E) vine thickel/scrub(8, OMMO 13146, 4 Sepi 1982. AM/QM-ABRS); Camp Creek, SW side of Bobby Range, S of Miriam Vale, SEQ (24°36'20"S, 151°33E) (1, QMMO13287, 6 Sept 1982, AM/OM- ABRS}; Limestone Ck. W. of Childers, SEQ (25°15730"S, 151°S§'E) (2, QMMO13009. 3 Sept 1982, AM/QOM-ABRS). OTHER MATERIAL SOUTH-EAST QUEENSLAND: Balburin S.F. (24°32°S, 151°29'E) rainforest (1, QMMO17260, & Mar 1977, QM party); summit Mt Booroon Booroon, SW of Miriam Vale. MVF/Araucaria, litter (1, OMMO168al, 17 Sept 1985, J. Stunisic, D. Potter); Dawes Range, MVF/Araucaria (24°28°S, 151°07'E) (1, AMC136781, 4 Sept 1982, AM/QM-ABRS). DIAGNOSIS Shell small, depressed, diameter 3,96-4.69 mm (mean 4.35 mm) with 3 7/8 to 4 3/8 (mean 4 1/8+) normally coiled whorls. Last whorl des- cending only very slightly, Apex and spire elevated (Fig. 29c), SP/BWW ratio 0.12-0,25 (mean 0.19) height 2.56-3.15 mm (mean 2.79 mm), H/D ratio 0.61-0.71 (mean 0.64). Apical sculpture highly modified, consisting of numerous, crowded dimples with a few, vague. radial ridges mainly atthe terminal portion of the protoconch (Fig. 29g), Protoconch of 1 1/2 ta 1 1/2+ whorls, mean diameter Y06.9.m at 1 1/2 whorls, Post nuclear sculpture of spiral grooves, closely spaced. occasionally interrupted and crossed by weakly protractively sinuated radial ridges (Fig. 29e). Secondary sculpture of fine, crowded, radial growth ridges particularly on the body whorl (Fig. 29f). Umbilicus small, barely covered by the reflection of the columella, diameter 0.62-).82 mm (mean 0.75 mm), D/U tatio 5.34—6.52 (mean 5.85 mm). Sutures im- pressed, Whorls rounded. Aperture roundly lunate. Lip simple, columella dilated, slightly thickened and deflected toward the umbilicus. Parietal callus not strongly developed. Base glossy, rounded. with a sculpture of incised spiral striae and weak radial growth lines. Colour yellow-brown with a darker, grey spire in some specimens. Based on 24 measured adults. Genitalia with epiphallus entering the penis through a simple pore (Fig. 30b). internally with longitudinal pilasters. Penis a muscular tube with spongy longitudinal pilasters (Fig. 30b), Penial retractor muscle inserting at the penis/epiphallus junction. Free oviduct (Fig. 30a) a long, thick- ened muscular tube, internally with longitudinal thickenings. Radula (Fig. 29d) with tricuspid central and lateral teeth in which the mesocone is long and lanceolate. Based on 3 dissected adults (QMMO17010, AMC142928), RANGE AND HABITAT L. protoscrobiculata inhabits the drier sub- tropical microphyll vine forests (with Araucaria emergents) between Mudla Gap and Bobby Range, SEQ, and rarely, moister subtropical notophyll yine forests where these are in jux- taposition with drier rainforest. Little is known about the microhabitat of £. protescrobiculata because most available material consists of dead shells collected from litter, However, some live specimens have been taken from the upper sur- face of rotting logs. The comparatively reduced sculpture of L. protoscrobiculata is not indica- tive of life under logs but rather suggests that the Species hves in curled leaves. or under bark of irees. REMARKS At first glance L. protoscrobiculata appears more like a helicarionid than a charopid. Reduced radial sculpture (Fig, 29a), shell colour. and coiling pattern are not typically charopid. However the peculiar apical sculpture (Fig, 29g), SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 47 reticulate adult microseulpture (Fig. 29f) and anatomy (Fig. 30), leave no doubts about its true alfinities. The incomplete secondary ureter (Fig, 30e) is inconsistent with living in drier habitats but in the case of L. protoscrobiculata the primary ureter is lengthened by reflexion at the kidney apex. L. pratescrobiculala appears-to be a derivative of the Ngairea group. The distinctive punctate ir incised spiral grooves on the past nuclear shell and coiling pattern of reduced number of rounded whorls can all be readily derived from Neairea. Anatomically, the differences from Negairea are not great. Setomedea Iredale, 1941 Sefomedew lredale. 1933, p, 53 nom, nud.; lredale, 1937a, p. 329, nom, nud.; Iredale 19414, p. 267; Iredale. 19416, p. 1: Burch, 1976, p. 133, Seronredia (error) Kershaw, 1950b, p. 7, TyPrE SPECIES Sureria seticostata Hedley, 1924; by original designation. PREVIOUS STUDIES Iredale (1933) introduced Sefomedea for Sulerta seticostata Hedley, 1924 without any uccompanying description. Subsequently Iredale (1941a, p. 267) provided a barely mini- mal description and [ consider that the name should date from that publication. Iredale (1937a, p. 329) included 'Endodanta’ aculeata Hedley, 1899 from Wollongong, NSW without giving reasons. Sefortedea, herein, includes S, sericostata, three new species from Queensland, and a possible new species from the Atherton Tableland. ‘Endodonta’ aculeata Hedley, 1899 is excluded to be dealt with later. DIAGNOSIS Small to medium sized Charopidae. adult shell diameter 2.56-5.71 mm, with 3 5/8 to 5 5/8+ normally to loosely (¢tudicostata) coiled whorls. Height of shell 1.36-3.37 mm. Apex and spire slightly (seticostata) to strongly (monteith!) elevated. Body whorl descending in front, Fratocanch shiny. rounded, with | 3/8 to | S/8+ whorls. Apical sculpture of numerous slightly (seticostata, janae) to strongly (nudicostata, monieithi) squiggly spiral cords with a few vague radial undulations mainly near the end of the pratoconch. Postnuclear sculpture complex, of broad spiral furrows (except in nudicosrate) and numerous, moderately spaced, complex radial ribs formed by the enlargement of 2-3 radiul riblets. The major ribs have periostracal blades. scalloped (caused by the spiral undula- tions in the shell surface) or normal (nudicos- tata), with or without (mudicastata) regularly spaced slender setae along their length. Micrus- culpture of fine, crowded radial riblets, 7-15 between each pair of major ribs, and numerous, crowded spiral cords on the radial ribs. Sculpture continuous on {he base. Umbilicus wide U- shaped ta narrow V-shaped (nudicostata). Su- lures impressed, Whorls shouldered above and rounded below a slightly compressed periphery. or rounded above and flattened at the basal mar- gins (nudicostata), Aperture roundly lunate, Lip simple, columella slightly thickened. Parietal callus developed, Colour yellow-horn lo browny-vellow with reddish-brown flammula- tions and suffusions. Foot short and hroad, bluntly tapered posteriorly. No caudal horn or foss developed. Colour of animal (in preservative) creamy; dark in audicosiata, Ommatophores (in preservative] black (sericostata, nudicostata) or white (janee, monteri), Kidney broadly triangular to elon- gate. Ureter sigmurcthrous, Secondary ureter in- complete with ureteric opening at rear of the pallial cavity adjacent to the hindgut. Initial part of the primary ureter reflexed and coiled about the kidney apex. Ovatestis two clumps of pal- mately clavate lobes of alveoli located in the apical Whorls of the digestive gland or above the stomach (nudicestaia), oriented parallel to the plane of coiling. Terminal male genitalia with a muscularised epiphallus which may be short (monteithi), moderately long, very long (/udicostata), or modified and partially incorporated into the penial apex (janve). Epiphallic pore simple, entering bulb of penis apically or subapically (seticasram, monteithi), or entering directly into lower penial chamber (jartze). Penial retractor muscle inserting on the penial apex (rtonteithr), on the epiphallus near the penial bulb, or on the epiphallus well removed trom the penis (nudicostata). Penis with an apical bulb separated from the main penial chamber by a muscular collar, or without a muscular collar (nudicostuata). Apical bulb with pilasters (wr teltht, Seticostata), vetge (madicastata) or with incorporated epiphallus (/anae). Penis proper with fleshy. longitudinal pilasters or low lon- gituclinal pustular thickenings (udicostata). Radula with tricuspid central tooth that has a 58 TABLE 4 - RANGE OF VARIATION IN SETOMEDEA (MEAN AND RANGE) UMBILICAL SP/BWW RATIO BODY WHORL NUMBER OF NAME D/U RATIO WIDTH RIBS/MM WIDTH RIBS (mm) H/D RATIO [-4 = aw = < a WHORLS SPECIMENS (mm) E £ (mm) (mm) hy (2.81-3.13) (15-154) 6.15 (5.42-7.16) 5 (61-94) 0.20 (0.14-0.24) (1.15-1.32) 1.23 0.25 (0.17-0.30) (0.47-0.59) 0.52 (3.41-4.47) 2.05 391 (1.75-2.34) 5- (458-5 1/4) 13 stata Setomedea MEMOIRS OF THE QUEENSLAND MUSEUM Setomedea 3 (2.68-3.68) O1-1.31) 59.67 (04-0.17) ( (4.57-6.87) (49-80) 1.36 491 (3.22-3.55) (4.12-6.71) #92 (8.30-9.33) (70-89) .21-10.32) (t 15-1,24) O15 (0.04-0.21) ( 490,58) e = du rae 1.87 (1,66-2.08) 45/4- janae Gal ( 3.71) (4 1/2-10 4.9/4) Setumedea 5 /4- 19 montettht (3.62-5.71) ( (2.56-3.11) 155 (1.36-1,70) (2.04-3.37) 3 T/R+ (35/8-t0 41/4) (4.1/2-5 5/K+) nudicostata Setomedea slender, lanceolate mesocone and smaller pointed ectocones. Otherwise typical. DISTRIBUTION AND ECOLOGY Setomedea has a disjunct distribution from Dorrigo, NSW, to Shiptons Flat, near Cooktown, north Queensland. It inhabits moist temperate and subtropical rainforests of northern NSW and southern Queensland, subtropical forests of the Mackay/Prosperine area, and humid tropical forests of north Queensland between Tully and Cooktown. Setomedea lives under and in logs and oc- casionally may be found under bark with Ngairea. Species show a preference for very moist habitat where decomposition is well ad- vanced. Large numbers of individuals can be found under the same log or piece of bark. PATTERNS OF SHELL VARIATION The umbilicus is typically wide U-shaped. Departures from this pattern are seen in S. nudicostata (Fig. 35b) from SEQ and Setomedea sp. (Fig. 44b) from the Atherton Tableland, north Queensland. Variation in postnuclear sculptureis small. The most dramatic change is the loss of perio- stracal setae and spiral furrows in Setomedea nudicostata (Fig. 35f). Periostracal setae are rare among the Charopidae and only a small number of Australian and New Zealand taxa posses them. On their own they do not indicate phylogenetic relationship. However in the case of Setomedea, anatomical patterns unite the group, and indicate that this character was derived from a common ancestor. PATTERNS OF ANATOMICAL VARIATION The incomplete secondary ureter of Setomedea is similar to that found in Oreokera, Hedleyoconcha, Mussonula, Lenwebbia and Ngairea. However, the apical region of the kid- ney has the pericardial lobe well developed (Fig. 36e), The initial section of the ureter is reflexed and coiled around the tip of the kidney giving the kidney apex an S-shaped appearance. While some genera (e.g. Lenwebbia), may have the initial part of the ureter reflexed, only Setomedea has this unusual coiling. The terminal male genitalia of Setomedea show variations in the internal structures of the penis, shape and size of the epiphallus, and the insertion of the penial retractor muscle. Typical- ly the epiphallus is a short, reflexed, muscular SYSTEMATICS OF AUSTRALIAN CHAROPIDAE SY tube entering the penis either apically or subapi- cally (Figs 42a, 33b). In S. janae part of the epiphallus ts incorporated into the pental com- plex (Fig. 39c) and may represent spatial adjust- ment in response to reduced whorl count. The apical and femule genitalia of Seromreden are without unusual features except in S. nudicostata which has the free oviduct very long and the ovotestis situated in the digestive gland just above the level of the stomach. The shift in position of the ovotestis from apicad in the diges- tive gland is probably related to reduced whorl count. COMPARISONS Sctomedea can be distinguished from other subtropical and tropical charopids with spirally lirate protoconchs by size, shell colour. and adult sculpture - Oreokera has a much larger trochoid shell (Stanisic, 1987, pls 1,2): Sinployea has a smaller, monochrome brown shell with more numerous, simple, radial ribs (Fig. 137a, e); and Omphalorapa is smaller in size, has broader apical cords, wide cup-shaped umbilicus and few. widely separate major ribs on the body whorl (Fig. 134a.b,e). KEY TO SPECIES OF SETOMEDEA 1 Shell with periostracal setae 0. cece 2 Shell without periostracal setae (Fig. 35a. f) ddoachoh baicoh doatestigen yolct oa cbartonnesethekew jtudicostata 2.Shell with looser coiling (Fig, 44a); fewer whorls (Fig. 44a); mean whorl count 3 7/8+ (Table 8)... rae Selomedea Sp. Shell with tighter coiling; many whorls; mean whorl count greater than 4 1/2 2.4.3 3.Apical sculpture af regular Spiral cords .......... 4 Apical sculpture of squiggly spiral cords (Fig. 41). cccseeseesset sees ree teesteessiess. Montemthi 4, Major ribs with simple periostracal blades (Fig. BOF), assesses cea csatevessessaseaecavereenseness SCH COSIAIA Major ribs with scalloped periostracal blades (Figs ABE) vsrvscsnecenesevessessgenerserseneseenseennes janae Setomedea seticostata (Hedley, 1924) (Figs 32-34; Tables 8. 9) Suteria sericostara Hedley, 1924, p. 221. pl. 32. figs. 41-44. Setomedea seticostata (Hedley): Iredale, 1937a, p, 329: [redale, L941a. p, 268, fig, 4; Iredale, 1941b, p. 1. Setomedia (stc) seticostata (Hedley); Kershaw, 19566, p. 7. COMPARISONS S, seticostata is characterised by a flammu- lated shell with spirally corded protoconch and adult sculpture of complex radial ribs, microspiral cords, microradials, spiral grooves and prominent periostracal setae. 5. janae has fewer whorls with strongly sealloped periostra- cal blades and more prominent radial grooves (Fig. 38a, e-g); S. monteithi is larger with squiggly apical cords and a more protruded spire (Fig. 41c, e). Anatomically the incomplete secondary ureter and coiled, reflexed primary ureter immediately distinguish S. seticostata from other sympatric charopids reviewed in this study. PREVIOUS STUDIES Hedley (1924) tentatively assigned this species to the New Zealand Suteria Pilsbry, 1892 ot! the basis of overall similarity to Suzeria ide (Gray, 1850) which also has periostracal setae. However, different apical sculpture of S. ide indicates that setal processes in the two species are convergent. Type MATERIAL. HOLoTyPE; AMC64511, Dorrigo, NSW, under logs. Collected by §.W, Jackson, Nov. 1910. Height ofshell 1.87 mm, diameter 3.58 mm, H/D ratio 0.52. D/U ratio 2.90, whorls 4 5/&+. PARATYPES: AMC63685, 3 specimens, same locality data as holotype. OTHER MATERIAL NEW SOUTH WALES: Clarence River (5, AMC 142467, Cox coll.}: Tooloom Scrub, Beaury §.F, (28°29°S. 152°24°E). NVF (7, QMMO10937, AMC128533, 15 Mar 1981, AM/QM - ABRS), Woodenbong Ck, § slope of Mt Clunie (28°20'S, 152°30'°E), rainforest (15, QMMO10979, AMC1I28585. 15 Mar 1981, AM/QM - ABRS); W slope Wilsons Peak, top of Condamine River valley (28°16°S, 152°28'E) (2, AMC128635, 16 Mar 1981, AM/QM - ABRS): N of Dorrigo, c. 1.5 km along 60 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 61 Es WS =O \ \\ r \Gz A D FIG. 33. Setomedea seticostata (Hedley, 1924). a,b. e.g. c. 17km from Tannymorel. on Tannymorel-Cambubal S.F, Rd. SEQ. QMMO12705; c, Cunninghams Gap, SEQ. QMMO12678: d, f, c. km E of Carrs Lookout. Koreelah S.F.. SEQ. QMMO16477, a, genitalia; b, details of penis/epiphallus junction: c, details of penis interior; d, hermaphroditic duct: e, ovotestis; f, talon: g, pallial cavity. FIG, 32. Setomedea seticostata (Hedley, 1924). a-c, Dorrigo, NSW, AMC63511, holotype; d, Acacia Plateau, Koreelah S.F_, NSW. QMMO10952: e-f, Mt Glorious, SEQ. QMMOQ14164: c. 1km E of Carr’s Lookout, Koreelah S.F., SEQ, QMMO16477, a-c, entire shell; d, whole shell showing setae; e, apical sculpture: f, post nuclear sculpture; g, central and lateral teeth; h, marginal teeth. Scale lines as marked. i MEMOIRS OF THE QUEENSLAND MUSEUM T j 26°S 28°S 29°S 30°S 151°E 152°E FIG. 34. Distribution of Setowmdea seticosrara (Hed- ley, 1924) Kelly's Road (30°U7S, 152"41'E), raintorest (6, AMC1 28320, 11 Mar 1981, AM/OM - ABRS); SE of Tabulam, tributary of Busby's Ck (29°02'S, 152°43°E) (12, QMMO12944, AMC137759, 29 Aug 1982, AM/QM - ABRS): MI Warning (28°24°S, 153°17'E) rainforest (22, OMMO10499, AM- C129302, 19 Mur 1981. AM/QM - ABRS); Toonum- bar S,F., ¢. 34 km trom Kyogle on Toonumbur forest toad (28°33°S, 152°45°E) (3, QMMO)0908, AMC 128498, 14 Mar 1981, AM/OM - ABRS); Wian- garie S.FL, c. 25 km NE of Kyogle (28°23'S, 153°06'E), 1000 m (7, AMC142481, 17 May 1976, PH, Colman, 1 Loch); Red Serub F.R.. Whian Whian §.F, (26°38'°S, 153°19°E), 210m (1, AMC 142484, 15 May 1976, PH. Calman, I. Loch); Dorrigo (23, AMC63760, S.W. Jackson); Koreelah S,F,, Acacia Plateau, (28°21°S, 152°24'E), SEVT/MVF (47, AMC128554, OMMO10952, 15 Mar 1981, AM/QM - ABRS), Beaury S.F., c. 15 ken W of Urbenville (28°21°S, 152°24'E), 810m (2, AMC 142483, 18 May 1976, PH, Colman, |. Loch): Rocky Ck, Red Scrub PLR. (28°38'S, 153°20'E), rainforest (9, OMMO10404, AMC128865, 20 Mar 198), AM/QM - ABRS); Terania Ck (28°34'S, 153°19 E), 340m (1, AMC142482, 16 May 1975, P.H. Colman, 1. Loch), Ballina (2, AMCLIIOS8, ex C, Hedley); Tweed River (4. AMC142485, ex Coa, ex Petterd); Murwillumbah, under logs (1, AMC28488, 3 Dec 1907, C.F. Laseron); Lismore (2, AMCS784, ex C. Hedley); Dorrigo (5, AMC63772. S.W,. Jackson); Bryan Bay (3, AM- €4312, ex Cox); Big Scrub, Murwillumbah (3, AM- C103795, Lower, ex Cox); Richmond River, Lismore, in scrub under decayed timber (1, AMC63872, ex Brazier), Sherrard Falls, Dorrigo, warm temperate raintorest, litter, 600 m(1,QMMO 16776, 3 Noy 1983, AM/OM - ABRS), c. 1.5 km E ulong Kelly's Rd in dense rainforest, Cloud's Creek §.F.. (30°07'S, IS2*41°E) (3, OMMO10773, AMC128320, Mai 1981, AM/QM - ABRS); along Croftby Rd, top of Condamine River Valley, (28°15°S, 152°29'R) (1, QMMO10540. Mar 1981, AM/QM - ABRS); c. 5.5 km E of Dorrigo, (30°23°S, 152°44'E) (4, OMMO10796, AMC128351, Mar 1981, AM/QM - ABRS 1981); Tooloom Scrub, Beaury S.F., under logs, NVF (3, OMMO 16900, 4 May 1986. J. Stanisic, J, Chaseling); Mt Glennie 16 km E of Woodenbong, 900 m (3, OMMO11933, 24 Oct 1982. G.B. Monteith, D. Yeates); c. 1.5 km from summit of Mt. Warning, (28°24'S, 153°16"E)(3, QMMO10515, AMC129319, Mar 1981, AM/QM - ABRS): Mt Warning, CNVF (28"24'S. 153°16"E) (2. QMMO12737, AMC 1368231, 15 Dec 1981, AM/QM - ABRS); Swan's Rd, Bruxner Park Flora Reserve, subtropical rainforest, under logs (6, OMMO14768, 14 Nov 1983, AM/OM - ABRS), SOUTH-EAST QUEENSLAND: Lower Ballanjui Falls track. Binna Burra, Lamington N.P., under logs (3, O€MMO1655 1,25 Sept 1986, J. Stanisic); Burleigh Heads N.P., Under Joys, vine thickel (2, QM- MO16922. 21 Jan 1987, J. Stanisic, D. Potter); ¢, 3 km SE of summil Mt Glorious. NVF (27°20°S, 152°46°E) (3, OMMO11990. OMMO1 L994. AMC136419, 2 Dec 1982, AM/OM - ABRS):¢, 25,2 km from Gaom- burra, Goumburra S.F., NVF/Palms/Araucaria (27°59°S, 152°21°E) (5, OMMO1N121. AMC 124618, 16 Mar 1981; 7, OMMO12691, 7 Dee 1981, AM/QM - ABRS); Curtis Falls Cireoii, Mt Tamborine, liter, CNVF (1, QMMO16436, 23 Feh 1986, J, Stanisic, J, Chascling), Mt Hobwee Cirenit. Binna Burra, Lamington N.P., litter NVF (10, OMMO18868, | Oct 1985, J. Stanisic, J. Chaseling, D. Poller); Tullawallal Circuit, Bina Burra, Lamingion N.P.. under logs. NVF (9. QMMQ16737, 2 Oct J98S, J. Stunisic, B- Potter, J, Chaseling): Border Track, Biona Burra, Lamington N.P., under logs. NVF (5, OMMOLA&S7I. 10 Mar 1984, J, Stanisic); Maiala N.P,, Mt Glorious, NVF (6, QMMO11603, OMMO1 1848, 20 Jun 1982, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 63 MSA party); Koreelah S.F., Acacia Plateau, under E 23 == logs, NVF/MVF (1, QMMO16796, 4 May 1986, J. 2 a S25 5 5 £2 & = & Stanisic. J. Chaseling): Lower Ballanjui Falls Circuit, 5 iid veins i ea “ ™ Binna Burra, Lamington N.P., NVF, litter (7, QM- MO16892, 30 Sept 1985, J. Stanisic, D, Potter. J. 3_ ge $= =e Chaseling); O°Reillys, Lamington N.P.. (28°14°S, 25>. + Fin 2 = 32 = 32 ~ 153°15'E) MVF (13, QMMO10427, 17 Mar 1981, ZF 35 255 25 75 = =2 = AM/OM - ABRS): Mt Glorious, NVF, litter (3, QM- MO14164, 28 Nov 1983, J, Stanisic); Araucaria 25 Track, Binna Burra, Lamington N.P., under logs, NVF =z ae ere F 23 _, (7, QMMO16578, 27 Sept 1986, J. Stanisic); c. 1 km 2 “ ‘ aa #2 © Eof Carrs Lookout, Koreeluh S.F., under logs, NVF (10,QMMO16477,3 May 1986, J, Stanisic, J, Chasel- = ing): Border Track, Binna Burra, Lamington N.P.., P| sa under logs, NVF. (9, QMMO16877, J. Stanisic, 31 z af Si) #2 , #4 = Mar 1984); Hall's Plain, E of Emu Vale, under logs, CNVF, (4, O€MMO 16895, 4 May 1986, J. Stanisic, J. coe ans es Chaseling) Cunninghams Gap, (1, QMM0Q6070, 21 zc Ff 38 22 Mar 1977. M.J. Bishop); Binna Burra, Lamington s 28 88 228 28 82 8 22 = NP., CNVF, 850 m (28713'S, 153°12°E) (1, OM- 2 M0O6073, Apr 1976, M.J. Bishop); The Knoll N.P., Mt z 3 7 . Tamborine. palm gully (27°55'S, 153°10°E) (1, QM- =_ 2. ha 38 M06305, 3 Aug 1976, M.J. Bishop); initial part of EE 422 YE RE ES 44 8 S32 = Kweebanj Caves Walk, Binna Burra, Lamington N.P. =e 5 oI = under logs. NVF (3, QMMQ16884, 9 Mar 1984, J. $2 Stanisic); Cunninghams Gap, near monument, 755 m, s2 ¢ 2 8 28 NVF (28°04°S, 152°24°E) (8, QMMO12678, AM- 22 SEF <— Ele == 22 = FE = C136807,7Dec 1981,J, Stanisic, 2, AMC128405, 16 == “2 Se Fee 5 5 8S 5 FS = Mar 1981, J, Stanisic, D, Potler; 3, QMMO16901, 5 = May 1986. J. Stanisic, J. Chaseling); Upper Pine : s "a es 2a Creek, via Canungra, under bark of fallen tree, rain- zg g —_ 29. Le an 53 ., 33 _ _ forest with Araucaria (6, OMMO16904, J. Stanisic, 3 2 S§ BEB £2 2 8 88 8 D. Potter, 21 Jan 1987): Coomera Falls Track, Binna g Burra Lamington N.P.. NVE, under log (1, QM- a a 4 ibs MOQ16575, 26 Sept 1986, J. Stanisic); Nagarigoon 3 z a ee Pp] | Falls Track, Binna Burra, Lamington N.P., under logs, = 2£ ss de ct 22 2 22 = NVE (1, OMMO16883, 20 Apr 1986, J. Stanisic, J. ‘ é ns . Chaseling): ¢. 17 km from Tannymorel, on Tan- nymorel-Cambubal S.F. Rd, 1000 m, MVF/Araucaria But ae £3 3& (3. QMMO12705, AMC136819. 7 Dec 1981, Se ops dp es 22 2 FS 3 AM/OM-~ ABRS 1981); Mt Glorious, in litter and ~ ses oS oes = "under logs, NVF (3, QMMOQ16128, 26 Jan 1986, J. 4 Stanisic, J. Chaseling); Natural Bridge N.P.,(28°13°S, 4 2 = = 153"14'E). NVF (7, QMMO10452, OMMO10461, g é< fe 22 i Fs 7.00 TIT) esse cates ses eeseeduetec costes ter eesenee PUAN Post nuclear sculpture of more widely spaced ribs: shell smaller (mean diameter < 7,00 3.Ribs on body whorl very widely spaced (Fig. 49y); mean ribs/mm 1.96.,.... paucilamellara Ribs on body whorl moderately spaced (Fig. 48a): mean ribs/MiM 3.92 yur convelura Gyrocochlea yinitincta (Cox, 1868) (Figs 45-47; Tables 11,12) felix vinitineta Cox, 1868, 9.18, pl 1. figs 6, 6a, Diplomphalus Wuitinctus (Cox), Tryon, 1885, p. 115. pl. 24, figs 88,89, Monomphalus vinitinctus (Cox); 359, Endodonta vinitincta (Cox); Shirley, 1921, p. 34. Gyrovociilea vinitineta (Cox); Hedley, 1924, p. 218, pl. 31. figs 28-31, Iredale, 1937a, p. 322; Iredale, 194 1a, p.268, fig. 6; Kershaw, 1956a, p. 138. Ancey, 1888, p. COMPARISONS The large size, simple sutures, beaded apical sculpture, and strongly depressed spire differen- tiate this species from the sympatric Elsorhera venithecata, which has a flat to slightly elevated Spire With strong curved radial ribs on the apex, channelled sutures and very inflated body whorl, G. vinitineta can be distinguished from G. pauctlamellata, and G. convolura by its larger size and more numerous radial ribs. Nautiliropa amicron has the general shell form of G, vinitineta but is smaller, with greater number of tibs/mm, body whorl that does not descend, and flammulated shell. TYPE MATERIAL HOLOTYPE: AMC63490), Upper Richmond River, NSW, Height of shell 4.87 mm, diameter 7.98 mm, /D ratio 0.61, D/U ratio 3.99, whorls 4 1/2+, PARATYPE: AMC103624. same data as holotype. OTHER MATERIAL Richmond River, NSW (5, AMC139754; 3, AM- C139769, Petterd, ex Cox); Byron Bay, NSW (5, AMC139753, S,.W. Jackson); Big Scrub, Murwillum- bah, NSW (1, AMC63837, Lower, ex Cox); Lismore, NSW (2, AMC139752. ex C. McLauchlan, Bent Coll); Red Scrub Flora Reserve, Whian Whian S_F.. NSW (28°38'S, 153°19°E), 210 m (3, AMC139751, 15 May 1976, P,H. Colman, I. Loch): Terania Creek, NSW (28°34°S,,153°19°B), (3. AMC153725, 16 May 1976, P.H. Colman, I. Loch): Mt. Warning N.P., NSW (28°24°S, 153°16°E) NVEF/Palms (2, QMMO10487, Mar 1981, AM/QM-ABRS; 2. QMMO12735, AMC136830, 15 Dec 1981, AM/QM-ABRS). Upper Tallebudgera Ck. SEQ, 600 m (2, QMMO15990, 9 Dec 1984, G.B, Monteith); Warrie Circuit, Springbrook N,P.. SEQ, CNVF, under log in litter (1, QMMO16902,.6 Apr 1986, J, Stanisic); c. 1.6 km N of Numinbah Valley Turnoflt, on Burleigh- Springbrook Rd, SEQ, NVF/Palms, litter (2, QM- MO17319, 17 Dee 1980, J, Stanisic); Natural Bridge N.P., SEO (28°13°S, 153°14°E), CNVF (1, QMMO- 6281, Oct 1976, M.J. Bishop). LAMINGTON N_P., SEQ; Border Track, near O'- Reillys NVF, under logs (3, QMMO16875, 31 Mar 1984, J. Stanisic); Orchid Bower, Binna Burra, NVF, under stone (1, OMMO16864, 19 Apr 1984, J. Stamasic. J. Chaseling); Tullawallal Circuit, Binna Burra, NVF, under logs (7, O€MMO 16739, 2 Oct 1985, J. Stanisic, D. Potter, J. Chuseling); Border Track, Binna Burra, NVF, under log (3, OMMO16869, 10 Mar 1984, J, Stanisie): Araucaria Track, Binna Burra, NVF, under logs (1, QMMOT6864, 19 Apr 1986, J, Stanisic, J. Chaseling); Tullawallal Circuit, Binna Burra, NVF, under logs (7, QMMOQO16739,2 Oct 1985, J. Stanisic. D. Potter, J. Chaseling): Border Track, Binna Burra, NVF, under Jog (3, OMMO16869, 10 Mar 1984, J. Stanisic); Arauearia Track, NVF, under logs (3, QMMOT6579, 27 Sept 1986, J_ Stanisic): O’Reillys (28°14°S, 153°1S°E) MFF (3, QOM- MO10431, 17 Mar 1981, AM/OM-ABRS): Binna Burra (28°13°S, 153°12°E) B50 m. CNVF (1, QMMO6078, Apr 1976, M,J. Bishop), 80 MEMOIRS OF THE QUEENSLAND MUSEUM —— = ee cs a are a go, ge) 7 “ee teehee a] ret ve : arity alll qerrevterte® ae ren SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 81 FIG. 46. Gyrocochlea vinitincta (Cox, 1868). Mt Warning N.P., NSW. QMMO12735. a, genitalia; b, penis interior; c, ovotestis; d, hermaphroditic duct; e, pallial cavity. Scale lines as marked. FIG. 45. Gyrocochlea vinitincta (Cox, 1868). a—c, Upper Richmond R, NSW. AMC63490, holotype; d, g—h, c. l.6km N of Numinbah Valley turnoff, on Burleigh-Springbrook Rd, SEQ. QMMOQ17319; e-f, O’Reillys, Lamington N.P., SEQ. QMMO10431, a—c, entire shell; d, jaw: e, apical sculpture; f, post nuclear sculpture; g, central and lateral teeth; h, marginal teeth. Scale lines as marked. PAWL LU) ORANG ILOL VARLATION IS ZEROCOCHIEA, NAUTTIRQTA, HIOMIHALOPA AND LETOMOLA 82 MEMOIRS OF THE QUEENSLAND MUSEUM = el sd od ad hd gt ¥e En 8 #5 Sa Se88 23 88°58 38 ~£f 8 8 S28 § E22 23 of 24 sd se 22 53 So otS Hos Ae we So eS see eee = = TS az a ef oF z &4 22 3f 22 G2 24 23 2 =z = = = 2 = = = i = = = 7 i = => Ss z car nz ni Sf st SF <7 = 22 =2 HR 8 28 Fe 22 = Ey o cae $2 i 452° 153°R 154°E =< zz nx == FiG 47. Distribution of Gyrocochlea vinitincta (Cox, - | a: a oe eee 1868). e297 € 28 4 2 = a | e 4 os - = pei po ay gq a2 o4 a8 82 28 DIAGNosIs e.7 r Shell diameter 7.22-8.57 mm (mean 7.91 = mm), with 4 1/4+ to 45/8 (mean 4 3/8+), tightly z coiled whorls, last descending more rapidly. é z Apex and early spire (Fig. 45a) strongly con- z - cave, height of shell 3.95-4.87 mm (mean 4.52 mm). H/D ratio 0.53-0.63 (mean 0.57). be mn ee pe gs Protoconch of | 1/2 to 1 3/4 whorls, mean = = <= < <= . . , - & ite ot we diameter 896.7 xm at 1 1/2 whorls. Apical sculp- 8 2232 #2 43% ture (Fig. 45e) of fine, crowded spiral cords and numerous, weakly curved radial ribs, which are 2: at ce Uz Aes Re o* oF € raised al their intersection with the spiral cords = e e & e > 3 . “ Ze 2 F f§ 8 FoF creating a beaded effect. Post nuclear sculpture ae SHES 46 35 26 FS Se (Fig. 45f) of numerous, high, crowded, protrac- tively sinuated radial ribs, 106-153 (mean 118.8) ribs on the body whorl. Ribs/mm 4.22— = 5 z= 5.68 (mean 4.77). Microsculpture (Fig. 45f) of = a % ae fine radial riblets, 6-12 between each pair of = ba ana major ribs, continuing onto the sides of the major ribs, and numerous fine microspiral cords which x 2 € « are not raised at their junction with the 3 aS ef..ve microradials. Umbilicus (Fig. 45b) relatively = 232 75 Fz Te wide, U-shaped, diameter 1.79-2.26 mm (mean - 2.00 mm). D/U ratio 3.46-4.32 (mean 3,97). 33 Sutures (Fig. 45a) weakly impressed, becoming a2): wk = " ae’ @ deeper on the body whorl. Whorls rounded zz below and strongly shouldered above a weakly compressed periphery. Aperture ovately lunate. 3 Lip simple, columella not expanded. Parietal 5 callus developed. Colour yellow-brown to dark reddish-brown. Based on 6 measured specimens. 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Gyrocochlea convoluta Hedley, 1924. Along tributary of Iron Pot Ck, [km N of Toonumbar, NSW. QMMO10927. a, genitalia; b, penis interior; c, hermaphroditic duct; d, pallial cavity: e, ovotestis clump Scale lines as marked. FIG 48. Gyrocochlea convoluta Hedley, 1924. a—c, Beaudesert. Qld. AMC5815, holotype; d, g—h, Along tributary of Iron Pot Creek, 1km N of Toonumbar, NSW. QMMO10927; e-f, Mt Lindesay, Richmond R, NSW. AMC39750. a-c, entire shell; d, lateral teeth; e, apical sculpture; f, post nuclear sculpture; g, cusp angles on lateral teeth; h, lateromarginal and marginal teeth. Scale lines as marked. 85 ft MEMOIRS OF THE QUEENSLAND MUSEUM rizht angles to the plane of coiling, one clump on the apex and one on the base of the whorl and directed toward each other, Epiphallus (Fig. 46b) kinked and entering the penis subupically through a conical verge (Fig. 46b). Penis (Fig. 46b) a long, slender, muscularised tube covered with a thin penis sheath. Upper portion of the epiphallus connected (o penis sheath by short fibres (Fig. 46a). Internally the penis has several longitudinal spongy thickenings (Fig. 46b). Radula without unusual features, Based on 2 dissected adults (QMMO12735). RANGF AND HABITAT G. yinitineta occurs in the Big ‘Scrub area of northern NSW, the eastern McPherson Ranges, SEQ, and some adjacent mountains and valleys of the Seenic Rim. It prefers cool to warm sub- tropical notophyll vine forests and does not in- habit adjacent drier vine forests with Araucarta emergents, Specimens have been taken from under logs and under stones, often in pairs. REMARKS G. vinitincta and G. paucilamellata are sym- patric in purts of the Lamington Plateau, SEQ, and the structural differences in their penes probably provide species recognition features for the snails. Gyrocochlea conyoluta Hedley, 1924 (Figs 48-50; Tables 11. 12) Gyracuchlea convoluta Hedley, (924 (part), p. 216, pl. 29, fis. 79; Iredale, 1937a, p. 322. COMPARISONS G. convoluta differs from G. paucilamellata in being slightly larger (Table 11) and in having a greater number of ribs on the body whorl. Anatomically the differences are minor. G. can- voluta (Fig. 49b) has more irregular longitudinal pilasters in the lower part of the penial chamber and a larger pustular zone apicad of the epiphal- lic pore than G. pauetlamellata (Fig, 52b), G, convoluta is casily distinguished from G, vindtineta by its smaller size and more widely spaced radial ribs, and [rom Nauiiliropa omicron by its fewer radial ribs, larger umbilicus and monochrome brown shell on which the last whorl descends. PREVIOUS STUDIES Gyrocechlea convoluta Wis bused on the holotype and specimens from Camerunga (= Canungra), SEQ. The latter material has not been found and efforts to collect at Canungra yielded only G, paucilamellata; \Wedley probably had both species. TYPE MATERIAL HoLvoryre; AMCS5&815. Beaudesert, Qld. Height of shell 3.36.mm, diameter 5.88 mm, H/D ratio 0.57, B/U ratio 3,14. whorls 4 5/8. OTHER MATERIAL Mi. Lindesay, Upper Richmond River, NSW (3, AMC139750, 27 Apr 1958, L. Price), &. 1.7 km along MeIntosh’s Rd, nr Back Creek, E side of Richmond Range, NENSW (28"25'S, 152°44'E), broad- leaf/sclerophyll (1, OMMOLO970, 15 Mar 1981, AM/OM-ABRS); along tributary of Tron Pot Creek, | km N of Toonumbar, NSW (28°34’S, 152°45"E) (1, QMMO10927, Mar 1981, AM/QM-ABRS), DIAGNOSIS Shell large, diameter 5.46-6.05 mm (mean 5.78 mm), with 4+ 1/2- to 4 3/4 (mean 4 5/8+) tightly coiled whorls, last descending more rapidly. Apex and early spire (Fig. 48a) strongly coneave, height of shell 3,03-3,78 mm (mean 3,36 mm), H/D ratio 0.55-0.63 (mean 0.58). Protoconch of | 1/2 to | 5/8+ whorls, mean diameter 785.5 pm at 1 1/2 whorls. Apical sculp- ture (Fig. 48e) of fine spiral cords and fine low radial thickenings. raised at their junction creat- ing a beaded effect. Postnuclear sculpture (Fig. 48f) of numerous, widely spaced, high, strongly protractively sinuated radial ribs, 58-83 (mean 71.5) ribs on the body whorl. Ribs/mm 3.38— 4.36 (mean 3.92). Microsculpture (Fig. 48f) of fine radial riblets. 18-30 between each pair of major ribs, and very fine, crowded spiral cords, Umbilicus (Fig. 48b) U-shaped, diameter 1,62- 1.87 mm (mean 1.70 mm). D/U ratio 3.14-3.55 (mean. 3.40), Sutures impressed, becoming deep on the last whorl. Whorls rounded below and strongly shouldered above a slightly flattened periphery (Fig. 48c). Aperture ovately lunate, Lip simple, Parietal callus developed, radially sculptured. Colour golden brown. Based on 4 measured adults, Genitalia with epiphallus long, muscularised, hound for part of its length to the apical part of the penis (Mig. 49a), internally with longitudinal thickenings, entering penis subapically through a fleshy pore (Fig. 49b). Penis (Fig. 49h) long, tubular, without expanded apical portion, inter- nally with thin fleshy longitudinal pilasters (Pig. 49b) which become less regular in the lower part SYSTEMATICS OF AUSTRALIAN CHAROPIDAE a7 154P FIG 50, Distribution of Gpracochlea convoluta Hed- ley, 1924. of the chamber, anda small pustulose zone above the epiphallic pore, Penial retractor muscle in- serting apically on the penis. Radula (Fig. 48g, h) typical. Based on | dissected specimen (QMMO- 11927). RANGE AND HABITAT G. convaluta inhabits notophyll vine forests of the Upper Richmond Range, NSW, and gallery forests of creeks which flow from these high- lands. No data is available on microhabitat al- though shell form suggests that it lives under logs. REMARKS G. convoluia is a conchologically distinctive species and differs only slightly from GC. paucilamellata in genital anatomy, Gyrocochlea paucilamellata sp, nov. (Figs 51-53; Tables 11, 12) Gyrocochlea convolutq Hedley, 1924 (part), p. 216. ETYMOLOGY Latin pauci, few; lamella, plate; for the sparse number of radial ribs on the adult whorls, COMPARISONS The brown shell, widely separated radial ribs, biconcave shell and wide umbilicus distinguish G. paucilamellata from its congeners and other charopids in the border ranges area. G. vinitincta is much larger with more numerous tadial ribs (Table 11). G. canvolura has more than twice as many ribs on the body whorl. Anatomically G. paucilamellata (Pig, 52b) differs from G. vinitineta (Fig. 46b) by lacking a penial verge and in having an apical pustular zone in the penis, Nautiliropa omicron, which also is sym- patric with G. paucilamellata, has many more radial ribs and a flammulated shell. Anatomical- ly N. omicron has a penis which lacks a sheath, contains complex pilasters, and is pear-shaped (Fig, 65c). TYPE MATERIAL HoLotyPre: OMMO17321, Upper Pine Creek, Canungra, SEQ. NVF/Araucaria, under bark and logs. Collected by J. Stanisic, D, Potter, 1 Oct 1986. PARATYPES: OMMO16546, 21 specimens, same data as holotype; Kweebani Caves Walk, Binna Burra, Lamington N.P., SEQ, NVF, under logs (1, QOMMO16889, 30 Sept 1985, J. Stanisic, D. Potter, J. Chaseling); Mt Tamborine, SEQ, under bark in leaf litter (4, OMMO17320, VK 7299. 3 Nov 1979, V. Kessner); Upper Pine Creek, Canungra, SEQ, NVF/Araucaria, under Jog (1, QMMOQ16585, 29 Sept 1986, J. Stanisic, J, Chaseling). DIAGNOSIS Shell large for family, diameter 5.04-5.55 mm (mean 5.35 mm) with 4 1/4- to 4 5/8 (mean 4 1/2-) tightly coiled whorls, Apex and early spire (Fig. Sla) strongly concave. Height 3.19-3.45 mm (mean 3.31 mm), H/D ratio 0.58-0.65 (mean 0.62), Last whorl descending. Protoconch of | 1/2+ to 1 5/8+ whorls, flat, mean diameter 745.3 wm al | 1/2 whorls. Apical sculpture (Fig. 510 finely reticulate, consisting of numerous, crowded fine spiral cords crossing numerous crowded, weakly sinuated radial ribs. Post apical sculpture (Fig. 31f) of few, bold, protractively sinuated radial ribs, 27-40 (mean 32.7) ribs on the body whorl. Ribs/mm 1.57—2.52 (mean 1.96), Microsculpture (Fig, 51f) of fine, crowded radial riblets, 30-45 between each pair of major ribs, and very fine inconspicuous spiral cords, only visible between the microradials, Um- bilicus (Fig. 51b) wide, U-shaped, diameter 1.49-1.75 mm (mean 1.65 mm), D/U ratio 3.13— 3,38 (mean 3.25), Sutures impressed, forming a deep furrow as the body whorl descends. Whorls rounded below and strangly shouldered above a weakly rounded periphery (Fig. 51c). Aperture ovately lunate. Lip simple. Colour deep reddish- 88 MEMOIRS OF THE QUEENSLAND MUSEUM SKY t8eus SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 89 FIG. 52. Gyrocochlea paucilamellata sp. nov. Upper Pine Ck, Canungra, SEQ. QMMO16546, paratype. a, genitalia; b, penis interior; c, talon and hermaphroditic duct; d, spermatophore; c, ovotestis: f, pallial cavity. Scale lines as marked. FIG. 51. Gyrocochlea paucilamellata sp. nov. a—c, Upper Pine Creek, Canungra, SEQ. QMMO17321, holotype; d, g—h, same data as holotype. QMMO16546, paratype: e-f, Mt Tamborine, SEQ. QMMO17320, paratype. a—c, entire shell; d, jaw; e, apical sculpture; f, post nuclear sculpture; g, central and lateral teeth; h, marginal teeth. Scale lines as marked. 90 MEMOIRS OF THE QUEENSLAND MUSEUM ‘SS BLismore MS fait. SN > 500m = 153°E 154°E FIG 53. Distribution of Gyrecochlea paucilamellata Sp. nov. brown ta light-yellow horn. Based on 3 measured adults. Genitalia with epiphallus long, muscular, and strongly coiled, entering the penis subapically through a simple pore (Fig. 52b) surrounded by. a fleshy collar. Epiphallus partially bound (api- cally) ta the penis sheath. Penis (Fig. 52b) elon- gate, tubular, muscularised, internally with longitudinal pilasters (Fig. 52b) in a regular pat- tern, and smaller pilasters apicad of the epiphal- lic pore. Penial retractor muscle inserting apically on the penis. Spermatophore subclavate with a long recurved chitinous tail at one end (Fig. S2d). Radula typical. Based on 4 dissected specimens (QMMO- 16546). RANGE AND HABITAT G. paucilamellata occurs in the eastern part of the Lamington Plateau, SEQ, and in the Dar- lington-Canungra Range urea as far northas Mt Tamborine, SEQ in cool subtropical to warm subtropical netaphyll vine forest. sometimes with Araucaria emergents (Upper Pine Creek, SEQ). The species has been collected from under fallen bark and logs. In spite of relatively con- FIG 54. Gyrocochlea curtistana (Hedley. 1912). centrated collecting in the Lamington Plateau area, it has been found there only once. REMARKS G. paucilamellata has a distribution quite dis- tinct from that of G, convoluta and is geographi- cally isolated from it by the Albert River Valley. Gyrocochlea curtisiana (Hedley, 1912) (Figs 54-56; 64a-c: Tables 11, 12) Endodonia iuloidea var, curtisiana Hedley, 1912, p. 264, pl. 9, figs 49-5). Gyrocochlea curtrsiana (Hedley); Iredale, 1937a. p. 323. COMPARISONS G. curtisiana is distinguished in the genus by its almost flat spire, apical sculpture of regularly arranged pits, often flammulated shell and large number of ribs on the body whorl, Nautiliropa omicron has apical sculpture of regularly ar- ranged pits, flammulated shell and large number of ribs on the body whorl, but has a very depressed spire and a body whorl which does not descend. PREVIOUS STUDIES Hedley (1912) deseribed G. curtisiana as a variety of ‘Helix’ juloidea Forbes, 1851 from Port Molle. Central Queensland, following com- parison of specimens with the type of H. ivloidea by E.A. Smith(Hedley. 1912, p. 265). Hedley (in Hedley and Musson, 1892) had included this variety with ‘Charopa’ omicron (Pfeiffer, 1851). The two Species are similar in shel! appearance but are anatomically distinct. Hedley (1924) listed a large number of species under ‘Gyracochlea’ without including his new variety but recorded ‘Gyracochlea' omicron as having a distribution as. far north as Miriam Vale, SEQ, There is little doubt that these early published records of 'G*. omicron from the Miniam Vale area Were based On specimens of G. curtisiana. TYPE MATERIAL HoOLoryre: AMC32990, "found under log in forest just autside Springs Scrub". Mt Larcom, Port Curtis, SEQ. Collected by SW, Jackson, Aug 1908, Height a-c, Granite Creek, foothills of Bobby Range, SEQ. QMMO14716: d-h. Kroombit Tops. SEQ. QMMO165)7. OMMOQ16453. a-c. entire shell; d, central and lateral teeth: e, apical sculpture; {, post nuclear sculplure; e, central and lateral teeth: h, marginal teeth. Scale line a5 marked. 91 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE MEMOIRS OF THE QUEENSLAND MUSEUM ras? ae ee a = _ az. x an } ] aS ca f 7 i” COU FIG 55. Gyrocochlea curtisiana (Hedley, 1912). Kroombit Tops. SEQ. QMMO16517. a, genitalia: b, pemis interior: c, hermaphroditic duct; d, ovotestis: e, talon: f, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 43 SAC > 500m N ~ 151°E 152°E 153° Mo 56, Distribution of Gyrocochlea eurtisiane (Hed- ley. 1912), of shell 2.52 mm, diameter 5.29 mm, H/D ratio 0.48, D/U ratio 3.45, whorls 4 1/4. OTHER MATERIAL Summit. Mt Booroon Buoraon, SW of Miram Vale. SEQ. MVFiAraucaria, liner (4. QOQMMOl6860, 17 Sep! 1985. J. Stanisic, D. Potter): Granite Creek, foothills of Bobby Range, SEQ (24°30.7°S, 151°30.3°E), under branch on ground (1, QMMQ- 14716, 7 Dec 1983, T. Carless); ¢. L8 km Eof Builyan railway station, on Butlyan-Gladstone Rd, SEO (24°32'S, 151°24'E)raintorest next toriver bank, litter (12, OMMOION319, 15 Jul 1980. J. Stanisic. A. Green); Dawes Range, SEQ. MVP/Araucaria (2428'S, lS1U7°E (47, QMMO12642, QM- MQ12667, AMC136771, AMC136796, 4 Sep1 1982. QM/OM-ABRS): base, Mt Booroon Booroon, SW of Miriam Vale, SEQ. rainforest along creek, under logs (3. QMMO16863. OMMO16594, 17 Sept 1985, J. Stanisic, D. Potter); Kroombit Tops. c. 13 km W of "Chapmans" SEQ (24°25°S. 151°02"E) NVF (12, OMMO12223, QMMO12224, AMC136552, AMC- 136551, 6 Sept 1982, AM/QM-ABRS); Mr Fort Wil- liam. Kalpowar S.F., SEQ, NVF (2, QMMO12597. AMC136756. 4 Sept 1982, AM/OM-ABRS), ¢, 4 km below summil of Mt Fort William, Kalpowar &.F., SEQ, MVFiAraucaria, under logs and rocks (2, OMMO17011, 3 Jul 1984, J. Stanisic, D, Potter, K. Emberton): Mt Fort William, Kalpowar 8,F,, SEQ. CNVF, 833 m, under logs (2, OMMO16822, 3 Jul 1984, J. Stanisic, D, Potter, K. Emberton): Kroombit Tops, SEQ, open forest (3, OMMO16453, 30 Sept 1985, G.B. Monteith); Kroombit Tops, SEQ. MVF/NVF, under palm fronds and logs (30, OMMQ- 16517, 10 May 1984, J, Stanisic, D, Potter), DIAGNOSIS Shell large, diameter 4.71-5.97 mm (mean 5.38mm) with 4 1/8 to43/4 (mean 4 1/2-) tightly coiled whorls. Last whorl descending in front, Apex and carly spire (Fig. 54a) almost flat, SP/BWW ratio 0.04-0.05 (mean 0.04), to depressed. Height of shell 2.52—3.28 mm (mean 2.84 mm), H/D ratio 0.47-0.65 (mean 0.53). Protoconch of 1 3/8 to I 3/8 whorls, mean diameter 718.8 pmat 1] 1/2 whorls. Apical sculp- ture (Figs S4e, 64a) of curved radial ridges crossed by very low spiral cords which are par- tially fused and worn, producing a pitted ap- pearance. Post nuclear Sculpture (Fig, 54f) of numetous, crowded, protractively sinuated radial ribs, 93-159 (mean 125.5) on the body whorl, Ribs/mm 6.13—9.12 (mean 7.42), Micros- culpture (Fig. S4f) of fine thread-like radial riblets, 7-15 between each pair of majorribs and low crowded spiral cords which buttress the microradials on either side. Microspirals con- tinuous on the rajor radials. Umbilicus (Fig, 54b) wide, V-shaped, diameter 1.45-1.92 mm (mean 1.69 mm). D/U ratio 2.76-3.52 (mean 3,23), Sutures decply impressed. Whorls rounded below and shouldered above a weakly rounded periphery (Fig. 54c). Aperture broad, ovately lunate. Lip simple. Parietal callus present. Colour variable, light-yellow horn with red chevron-like markings. to dark-brown, Based on 17 measured adults. Epiphallus (Fig. 55a) long, muscular, entering penis subapically. Upper portion of epiphallus connected to penial sheath (Pig. 55a) by long fibres. Penis (Fig. 55b) long, muscular, internal- ly with several longitudinal pilasters, more slender and more crowded apically, Epiphallus entering through a pore (Fig. 55b) surrounded by 4 spongy thickening. Penial retractor muscle (Fig, 55a) Jong, twisted around the epiphallus. inserted on the penis. Vagina long, internally with fleshy longitudinal pilasters. Radula without unusual features. Based on two dissected specimens (QMMO- 16517). RANGE AND HABITAT G. curtisiana is confined to the Burnett-Cal- b/U RATIO 240 UMUILIC AL WIDTH (mm) tue RIGS/MM 747 RIBS 100 Sw RATIO WIDTH (ren) BODY WHORL 187 (even) (MEAN, SEM AND RANGE) PROTRUSION SPIRE H/D RATIO 0.56 DIAMETER (mm) 4.60 TAMLE 13 LOCAL VARIATION IN BIOMPHALOPA RICAVA (HEDLEY, 1912) AND HOMPUALOEA CONCINNA (HEDLEY, 1924) eicirr (mm) 2.68 WHORLS 38 NUMBER OF SPECIMENS NAME Biomphalopa recava AMC 32992 (Holutype) Finch Hatten MEMOIRS OF THE QUEENSLAND MUSEUM liope-Many Peaks-Dawes Ranges, SEQ, with an outlier population at Mt Larcom. As with other Gyrocochlea, G. curtisiana occurs outside of larger rainforest areas and has been collected in remnant rainforest at Builyan, SEQ, and ‘open forest’ at Kroombit Tops. G. curtisiana favours warm subtropical notophyll vine forests but may be found in adjacent microphyll vine forest (with Araucaria). ns nee 2.87 2a 4 be RS a iin 23 2 r} REMARKS The pitted apical sculpture, almost flat spire, rounded body whorls and flammulated shell of G. curtisiana (Fig. 54a,c,e) are departures from the average Gyrocochlea pattern. However geni- zB g % 83 tal anatomy, in particular the form and internal characteristics of the penis (Fig. 55b), is typical. l7 6.38 Wd 878 O05 TRA Biomphalopa gen. nov. ETYMOLOGY From the Greek omphalos, meaning um- bilicus; referring to biconcave appearance. iol ] aU 1ay 1,24 10 TYPE SPECIES Endodonta recava Hedley, 1912. PREVIOUS STUDIES Hedley (1924) introduced Gyrocochlea for charopids with biconcave shell and prominent radial sculpture on adult whorls, and included G. coneinna Hedley, 1924 and Endadonta recava Hedley. 1912. These species, characterised by boldly ribbed, tightly coiled biconcave shells on which the protoconch is macroscopically smooth and glossy, are here separated to a new genus. Ost 0.53 0.60 O54 058 os) 47 469 2d 345 DIAGNOSIS Shell diameter range 2.64-4.69 mm, with 4 1/8 to 5 3/8 tightly coiled whorls, last descending more rapidly. Apex and early spire deeply sunken. Apical sculpture of regularly arranged pits and vague radial ridges. Post nuclear sculp- ture of high, bold, crowded, protractively sinuated radial ribs. Microsculpture of fine radial tiblets and equally fine spiral cords, continuous on the surface of the major radials. Umbilicus wide and cup-shaped. Sutures becoming deeply Se furrowed as the last whorl descends. Whorls strongly shouldered above and rounded below a laterally compressed periphery, Aperture acute ovately lunate. Colour creamy-white with darker radial suffusions. Animal with long slender foot and varying degrees of black speckling on the visceral coil. 2.47 isa a7 16) 79 SW4 Ae Ans AMC 1412300 QMMO 11757 Hliomphalopa concinna AMC BSI2 (Holutype} Kennedy SF OMMU 6417 Mt Uellenden Ker OMMO 11345 OMMO 14877 Eungella Cardwell SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 95 Kidney with pericardial lobe long and slender, rectal lobe reduced. Male genitalia with epiphal- lus shorter than penis, muscular, entering penis sub-apically through a simple pore. Penis without a sheath, either tubular with a large central longitudinal pilaster and a group of smaller apical longitudinal pilasters (recava), or with a swollen apical section containing a large corrugated verge (concinna). Terminal female genitalia with very long vagina and free oviduct. Radula with tricuspid central and laterals in which mesocone is long, slender lanceolate. DISTRIBUTION AND ECOLOGY Biomphalopa recava is found in subtropical notophyll vine forests of the foothills and uplands of the Eungella Range whereas B. con- cinna occurs mainly in notophyll vine forests of the Cardwell Range and montane vine fern thick- ets and vine forests of the Bellenden Ker Range. Both species live under logs. COMPARISONS Biomphalopa is distinguished from Gyro- cochlea by its smaller size (Table 11), greater number of tightly coiled whorls, bolder radial ribs and wider, cup-shaped umbilicus. Nautiliropa also has a sunken spire with pitted apical sculpture (Fig. 63d) but is larger, has fewer, loosely coiled whorls, finer and struc- turally different radial ribbing (Fig. 63e) and a body whorl which does not descend (Fig. 63c). Anatomically the terminal male genitalia of Biomphalopa bear some similarities to those of Gyrocochlea but the lack of a penis sheath and presence of a single longitudinal penial pilaster contrast with Gyrocochlea (Fig. 46b). The super- ficially similar Nautiliropa has a pear-shaped penis with quite complex pilasters (Fig. 65c). Biomphalopa can be readily derived from Gyrocochlea and common ancestry is probable. Biomphalopa recava (Hedley, 1912) comb. nov. (Figs 57-59; Tables 11, 13) Endodonta recava Hedley, 1912, p. 267, pl. 10, figs 58-60. Gyrocochlea recava (Hedley); Hedley, 1924, p. 217; Iredale, 1937a, p. 323. COMPARISONS B. recava differs from B. concinna in its larger size, greater number of whorls, less crowded ribs and wider umbilicus. The penis of B, recava has a large central pilaster and group of smaller apical longitudinal thickenings in contrast to the slender central pilaster and large corrugated api- cal pilaster of B. concinna. TYPE MATERIAL HOLOTYPE:: AMC32992, Finch Hatton, W of Mack- ay, MEQ. Scrub. Collected by S.W. Jackson, 2 Sept 1908. Height of shell 2.68 mm, diameter 4.60 mm, H/D ratio 0.58, D/U ratio 2.40, whorls 5 3/8. OTHER MATERIAL Eungella Range, 50 miles W of Mackay, MEQ (2, AMC140230, Sept 1957, L. Price); Finch Hatton N.P., W of Mackay, MEQ (21°04’S, 148°38’E), 180 m, (2, AMC153726, 29 Apr 1975, J.B. Burch, W.F. Ponder, P.H. Colman); Digging’s Rd, Eungella N.P.. MEQ (21°09°S, 148°29"E) NVF (10, QMMO11757, QMMO0O13073, 8 Jul 1982, J. Stanisic); Palm Walk, Eungella N.P., MEQ (21°00°S, 148°30°E) 800 m, CNVF (1, QMMO6345, Nov 1976, M.J. Bishop); Dalrymple Heights, Eungella N.P., MEO, NVF, 1000 m (2, QMMO6367, Nov 1976, M.J. Bishop; 1, AMC153727, 28 Apr 1975, J.B. Burch, W.F. Ponder, P.H. Colman); Broken River, Eungella N.P., MEQ (21°10°S, 148°30°E) NVF (1, QMMO11715, 5 Jul 1982, J. Stanisic); Finch Hatton Gorge, c. 10 km N of Finch Hatton, MEQ (21°04’S, 148°38°E) NVF (1, QMMO0O11749, Jul 1982, J. Stanisic). DIAGNOSIS Shell diameter 4.17—4.69 mm (mean 4.49 mm) 5+ to 5 3/8 (mean 5 1/4-) very tightly coiled whorls, last descending very rapidly. Apex and early spire (Fig. 57a) deeply sunken, height 2.13—2.68 mm (mean 2.43 mm). H/D ratio 0.51- 0.58 (mean 0.54). Apical sculpture (Fig. 57e) of pits and vague radial ridges. Protoconch of 1 5/8 to 1 3/4+ whorls, mean diameter 671.0 pm at 1 1/2 whorls. Postnuclear sculpture (Fig. 57f) of high, crowded, strongly protractively sinuated radial ribs, 81-100 (mean 91.7) ribs on the body whorl. Ribs/mm 6.17—7.47 (mean 6.67). Micros- culpture (Fig. 57f) of fine radial riblets, 10-25 between each pair of major ribs and fine spiral cords which are continuous on the apices of the radial ribs. Umbilicus (Fig. 57b) very wide, cup- shaped, diameter 1.87-2.00 mm (mean 1.93 mm). D/U ratio 2.23-2.40 (mean 2.32). Sutures impressed becoming deeply furrowed as the last whorl descends. Whorls (Fig. 57c) shouldered above and below a laterally compressed periphery. Lip simple. Colour creamy-white with brown to reddish-brown radial flammula- 96 MEMOIRS OF THE QUEENSLAND MUSEUM 97 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE FIG. 58. Biomphalopa recava (Hedley, 1912). Digging’s Rd, Eungella N.P., MEQ. QMMO11757. a, genitalia; b, penis interior; c, ovotestis; d, hermaphroditic duct; e, ovotestis: f, talon and carrefour; g, pallial cavity. Scale lines as marked. FIG. 57, Biomphalopa recava (Hedley, 1912). a—c, Finch Hatton, W of Mackay, MEQ. AMC32992, holotype: d, h, Palm Walk, Eungella N.P., MEQ. QMM0O6346; e-g. Digging’s Rd, Eungella N.P., MEQ. QMMO11757. a-c, entire shell; d, central and lateral teeth; e, apical sculpture; , post nuclear sculpture; g. details of major radial rib; h, marginal teeth. Scale lines as marked. 98 MEMOIRS OF THE QUEENSLAND MUSEUM 149°E FIG. 59. Distribution of Biomphalopa recava (Hed- ley, 1912) in mid-east Queensland. (Pr = Proserpine; Ma = Mackay; Sa = Sarina). tions. Parietal callus strongly developed. Based on 3 measured adults. Genitalia with epiphallus (Fig. 58b) entering penis subapically through a simple pore (Fig. 58b). Penial retractor muscle inserting apically on penis head. Penis (Fig. 58b) swollen with a large central, longitudinal corrugated pilaster and shorter longitudinal pilasters apically. Free oviduct short, muscularised. Vagina long, inter- nally with longitudinal pilasters. Radula (Fig. 57d, h) typical. Based on 4 dissected specimens (QMMO- 11757, QMMO6345). RANGE AND HABITAT Under logs in warm subtropical notophyll vine forests of the Eungella area (southern extension of the Clarke Range) and Finch Hatton Gorge, MEQ. It has not been found in the drier rain- forests of Mt Dryander, near Proserpine, MEQ. REMARKS B. recava is distinguished by a biconcave shell with prominent radial ribs and tight coiling. ’Gyrocochlea’ iuloidea (Forbes, 1851), from Port Molle (=Airlie Beach), has a depressed spire and wide umbilicus but is larger with more loose- ly coiled whorls, apical sculpture of spirals and radials, broad V-shaped umbilicus, more rounded periphery, more prominent microspirals on the body whorl and monochrome, usually dark-brown, shell. Biomphalopa concinna (Hedley, 1924) comb. nov. (Figs 60-62; Tables 11, 13) Gyrocochlea concinna Hedley, 1924, p. 215, pl. 29, fig 1-3; Iredale, 1937a, p. 322. COMPARISONS See under B. recava. TYPE MATERIAL HoLotyPe: AMC8902, Cardwell, NEQ. Height of shell 1.58 mm, diameter 2.64 mm, H/D ratio 0.60, D/U ratio 2.57, whorls 4 1/8. OTHER MATERIAL Kirrama Range, via Kennedy, NEQ, 500 m, rainforest (1, QMMO17322, 2 Oct 1980, G.B. Monteith); Ken- nedy S.F., NEQ (18°13’S, 145°46’E), 900 m (1, QMM06417, Aug 1977, M.J. Bishop); Dunn Creek, on Kirrama S.F. road, Cardwell Range, NEQ, NVF, (1, AMC14296S, 27 Sept 1980, I. Loch); Mt Bellen- den Ker, NEQ, just below summit, c. 1500 m, under bark on rotting log, MVFT (2, QMMO14878, 4 Jul 1983, J. Stanisic, D. Potter); Mt Bellenden Ker, NEQ (base, 17, QMMO11297; 500 m, 2, QMMO11326; 1054 m, 5, QMMO11343; summit, 5, QMMO11255; Earthwatch/QM, 1981); Mt Bartle Frere, NEQ, walk- ing track below NW peak on western side, c. 1000 m, SNVF (2, QMMO11381, 8 Nov 1981, Earthwatch/- QM). DIAGNOSIS Shell diameter 2.64-3.45 mm (mean 3.08 mm), 4 1/8 to 5 1/8 (mean 4 5/8+) tightly coiled FIG. 60. Biomphalopa concinna (Hedley, 1924). a—c, Kennedy S.F., NEQ. QMMO6417; e-g, Mt Bellenden Ker, NEQ. QMMO14878, QMMO11297, QMMO11255; d, h, Cardwell, NEQ. AMC8902, holotype. a-c, d,h, entire shells; e, apicalsculpture; f, post nuclear sculpture; g, central and lateral teeth. Scale lines as marked. 99 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 100 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 61. Biomphalopa concinna (Hedley, 1924). Mt. Bellenden Ker, NEQ. QMMOI112SS5. a. genitalia; b, penis interior; c, hermaphroditic duct; d, talon; €, interior of free oviduct; f, g, ovotestis: h, pallial cavity. Scale lines as marked, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE T45°E i46°E FIG. 62. Distribution of Biomphalopa concinna (Hedley. 1924) in north-east Queensland. (Ca = Cairns: In = Ingham; Tu = Tully). whorls, last whorl descending more rapidly. Apex and early spire (Fig. 60a) strongly depressed. height 1.58—-1.87 mm (mean 1.71 mm) H/D ratio 0.53-0.60 (mean 0.56). Protoconch glossy, | 5/8 to 1 7/8 whorls, mean diameter 600.5 jum at 1 1/2 whorls. Apical sculp- ture (Fig. 60e) of radially arranged rows of pits. Post nuclear sculpture (Fig. 60f) of crowded, high, protractively sinuated radial ribs, 80-95 (mean 84.8) ribs on the body whorl, Ribs/mm 7,84-9.63 (mean 8.82). Microsculpture (Fig. 60f) of fine radial riblets, 6-20 between each pair of major ribs and fine crowded spiral cords, continuous on the apices of the radial ribs. Um- bilicus very wide, cup-shaped, diameter |.03— 1,27 mm (mean 1.19 mm), D/U ratio 2.28-2.80 (mean 2.58). Sutures impressed, deep on the last whorl. Whorls rounded below and shouldered above a strongly compressed periphery (Fig. 60c). Lip simple, columella not expanded. Parie- tal callus strongly developed. Colour creamy- white to light-brown, with darker brown radial suffusions. Based on 4 measured adults. Genitalia with Jarge, muscular epiphallus which reflexes before entering the penis subapi- cally (Fig. 61a). Penial retractor muscle inserted apically on the penis. Penis (Fig. 61b) with a large apical bulb, otherwise tubular, internally 101 with an apical, fleshy swollen pilaster (Fig. 61b) adjacent to the simple epiphallic entrance. Penis proper with a single. thin, almost lamellate, lon- gitudinal pilaster (Fig. 61b) and a few smaller thread-like longitudinal ridges, otherwise smooth. Free oviduct a short muscular tube. Vagina long, internally with very prominent cor- rugated longitudinal pilasters. Radula (Fig. 60g) typical. Based on two dissected specimens (QM- MO11255). RANGE AND HABITAT Under logs and under bark on rotting logs in the Kirrama Range, NE of Ingham, and the Bel- lenden Ker Range, SE of Cairns, NEQ; it prefers montane notophyll vine forests. REMARKS The verge in the penis indicates the possibility of sympatric relatives. Nautiliropa gen. nov ETYMOLOGY Referring to the nautiloid-like appearance of the shell. TYPE SPECIES Helix omicron Pfeiffer, 1551. DIAGNOSIS Moderately large size, c. 4 1/2 tightly coiled whorls, the last inflated and not descending, Apex and early spire (Fig. 63a) strongly depressed. Apical sculpture of Jow radial ribs and low spiral cords, modified to produce a pattern of regularly arranged pits (Fig. 63d). Post nuclear sculpture (Fig. 63e) of numerous, protractively sinuated radial ribs formed by the periostracal extension of a single underlying radial thickening. Microsculpture of crowded microspirals and microradials. Umbilicus wide, U-shaped. Sutures impressed, whorls rounded below and shouldered above a rounded periphery (Fig. 64f). Aperture ovately lunate, lip simple. Animal with long. slender foot, weakly speckled with darker pigmentation on the lower spire. Pallial cavity elongate with complete sig- murethrous ureter. Kidney with long pericardial lobe and vestigial rectal lobe, Epiphallus (Fig. 65b) not bound to the penial sheath, entering penis apically through a fleshy pilaster. Penis (Fig. 65c) pear-shaped with a muscular collar wo and a complex pilaster pattern. Penial bulb con- taining epiphallic entrance and horseshoe- shaped pilaster. Lower penial chamber with api- cal sumulator (Fig. 65c) and longitudinal thick- enings. Terminal female genitalia short. Radula (Fig. 63c.f) with hook-like tricuspid central, bicuspid claw-like laterals without ectacone, and bicuspid marginals with long basal shafts and short endo- and ectocones. Teeth raws in a V- shape. COMPARISONS Nautiliropa emicron is removed from Gyra- coehlea because of basic differences in majortib structure and coiling pattern. Gvrecochlea has mujor ribs composed of several small ribs, and a final whorl which descends (Fig. 45c), whereas Nautiliropa has a last whorl which coils more tightly without descending. and major ribs com- posed of a single rib with a large periostracal blade (Fig. 63e). Although the modified apical sculpture of Nautilirapa is also present in Gyrocochlea curtisiana (Hedley, 1912) this fea- ture has evolved independently in several lineages of Charopidae. Nauliliropa omicron (Pleiffer, 1851) comb. nov. (Figs 63, 64d-f, 65, 66; Tables 11, 14) Helix orticron Peiffer, 1851, p. 128; 1854, p.457, pl. 155, figs 13-17; Cox, 1868 p.18, pl. 10, fig.. Helix ammanitoides Reeve, 1854, pl. 181, sp. 1246. Gyracachlea emicron (Pfeiffer); Medley, 1924, p. 216, pl. 30, figs 16-18; Iredale, 19374, p. 323: Tredale, 1941a, p, 268, COMPARISONS. N. emicron is most liable to be confused with species of Gyrocochlea which are sympatric with it in castern parts of the McPherson Ranges, SEO, The flammulated shell is good for field identification. However, rare monochrome brawn specimens of N. emicron need to be dis- tinguished by the more crowded radial ribs, smaller size, pitted apical sculpture and non-des- cending body whorl. G. curtistana has a flammu- lated shell with crowded radial ribs (Fig. 54a) but lacks the strongly depressed spire of W, oneron, PREVIOUS STUDIES Early records of Nautiliropa omicron (Pfeit- fer, 1851) cited by Cox (1868), Hedley and Mus- son (1892) and Hedley (1924), included records of Gyracechlea curtisiana which is restricted ta MEMOIRS OF THE QUEENSLAND MUSEUM a small area from south of Miriam Vale to Mt Larcom, SEQ. Attempts to locate the type of N. omicron have been unsuccessful. Most probably it was housed in the Stettin Museum and destroyed during the World War II, The initial deseription and subsequent figures (Pfeiffer, 1854) leave no doubts as to the identity of the species. Tyre MATERIAL Neo TYPF: QMMO2729 (ex OMMOLOSOL) Mt Warning N.P., NSW (24°24°S, 153°17°B), top of rowd in rainforest, Heizh! 3.03 mm, diameter 5,71 mm, H/D rato 0.53, D/U ratio 3.98, whorls 4 3/B-. OTHER MATERIAL W side of Somerset Dam, NW of Brisbane, NE of Esk. SEQ. open sclerophyll an hillside (2, AMCS3175, 6 Sept 1970, W.F. Ponder & P. Marsh); NW of Brisbane, SEQ, Laceys Ck just § of Raynbird Ck,, open dry sclerophy!l and Lantana along creek, (27°14°S. 152°43°E) (2. AMC 137792, OMMO12976, 30 Aug 1982, AM/QM-ABRS); ¢ 15 km from Leguime on Acacia Plateau Rd, Noreelah Ck S.F., NE NSW... 800 m, NVF/MVF (10, AMC136824, QMMO12721, 8 Dec 1981, AM/OM - ABRS); Couloula Nat. Park, SEQ (25°57'S, 153°06°E) SNVF on sand (7. AMC136487, AMC136490, OMMO12111, QMMO12113, 6 Sept 19K2, AM/QM ABRS); Coolomong Sanctuary, upper Currumbin Valley W of Tweed Hus, SEQ, rainforest reserve, litter (1, AMC137778, 30 Aug 1982. AM/QM - ABRS): Macpherson Ra. SEO (1, AMC139775, Lower, ex Cox): Mt Tamborine, SEQ (2. AMC63795, J. Sim- monds): Bnsbane. SEQ (2. AMC 139767, ex Cox); off trib. lo Back Ck., Unumegar S.F., NE NSW (28°26'S, 152°42°E) (1, AMC128584, 15 Mar 1981, AM/OM - ABRS); Mt Nebo. W of Brisbane, SEO (1, AMC 140436, 23 Mar [943, G.P. Whitley); Natural Bridge N.P., SEQ, NVF (28°13°S, 153°14'E) (8, AMC129269, AMC129271. QMMO10457, QMMO10460, 1S Mar 1981, AM/QM - ABRS); Mt Tamborine, SEQ. (1, AMC140439, JA. Simmonds); Nth Pine River, SEQ, in scrubs (10, AMC103605, ex Cox); Little Yabba Ck., Imbil S.F.. SEQ, NVF (26°28"S, 152°38'E) (2, AMC136448, OMMO12041, 7 Sept 1982, AM/OM - ABRS); sidetrack off Mt Archer Rd, Mt Mee 8.F., SEQ. NVF/Arauecaria (27°04'°S, 152°41'E) (8, AMC136523, QMMQ12181, 31 Sept 1982, AM/QM - ABRS); Dandabah, Bunya Mis N.P., SEQ, behind pieni¢ urea (26°53'S, 151°36°E) (83. AMC136607, OMMO 1] 2283, 31 Sept 1982, AM/OM - ABRS); Mur- willumbah, NSW (2, AMC139765, Lower, ex Cox); Mt Lingdesay, Upper Richmond R.. NSW (2, 103 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE (c0'e-zP'z) SO0FLIT PRT (co'r-ze'e) TOVOFLS E (o9'e-z9'2) 6POOF LTE f£e (once) ISQUFESE 60 (zz'b- 16°¢) 6110766 (36°E15't) LEVORILE OlLVa 1/d (98°0-99'0) B100-8L-0 90 (eo) 1POOF0S'T (09° 1-961) OZ VOF8E I te $s (p9' 1-981) B10 UFRSI tz (9 1-1z'1) LEOOFIE orl zl ort (pp I-86°1) RIOUFITL (wu) HLGIA qvormawn (zR'9-ze'S) OPTOFSTD (peu-te'9) SOS'OF 189 eS (6z'L-£r'S) p9OLOFZLY RO'6 (66-278) ZISOFHZT'S LoL rae SLL (so9-zirs) POL OFLES WW/SHIY (pt t-i6) Is EFS P01 Lo wom (1e1-68) OBER LIL trl (.91-971) SUEIF0'7S1 (96-LL) ELOFL OR saa OLLVa MAES (910-490) 80007690 oan £90 _ (61'7-@0'2) HE OFIUT — (z0'z-S8' 1) SRU'0F b6'T PLO UFOET “2 Lez = (172-012) 6P0'0761'Z =A raid 7 ra ra ae 6lz ce (SeZ-61'2) £SOOFPTT ~ (wu) (wu) HLGIA NOISNU.LOUd TYOHM AGO aulds (AONVH GNV WS ‘NVAN) (is-or'0) (pr'z-SR'1) 1007560 SEO'0FLU'T rro 18'1 (9s0-8F'0) (or s-€1's) Hourzs0 9p U-ZE'S (9s'0-S5'0) (z8°t-02'r) SNOUFISY) SEC OFRS'F rsd Ler 1s0 ers (ope-0re) (L6'S-12'S) M0'OFESO OLUUFIS'S is0 F's (s°0-+5'0) (9p'S-L8"+) 600°0755'0 zs sso Ine 9s0 wp rs0 ers (ss'0-€5°0) (1L'S-18't) LODOFPSO 952 UF1Z'S (um) OLLVY G/H YALAWVIC (ca 1-zH#'0) 9100-260 ORO (982-192) SHY UFOLT (697-St7) OLVOFTSZ (LVe-2L'2) PEO OFLOT O8'Z (g0'g-Lt Z) 9LOUF ORT Let (c0"e-69'Z) COL OF LRT (um) LHDIH (261 ‘ATTOIH) SALYOLNOD VIOWOLTT ONY (S81 “Wass ad) NOXIINO VdONTILLOYN NI NOLLWISVA 1V907- +1 STV (+-2/1 €) “WLE RSE (42/1 t-8/1 p) “WEP (r/L ep) “WLP +8/E F “HED (48/5 t-A/E b) +UIP REP (R/S bF/ IF) +H/E b wer wie Wi Cue P-8/L P) P/LP STHOHM ra] 6tf6L] OWV SOAB-) YPQRSST A (adAyoa"{) i poOreT OWV poosuays smyso uo) DjOWOII'] 9 cR7Z1 ONWO surrUnoW, RAUNg] S601 ONWO “4S YR|ITIOy a 1 74971 ONWO “4S BunquivoD, 1 ¥eSkL ONWO dry s,weysuiuuny i St7Z1 ONWO aury nngyonig i gf£4 ONWO a7 IW £ 17191 ONWO snouo] IW 1 92891 ONWO SATIAWO 1 RELYL OWWO euing] EUUIE) l Ls¢01 ONWO ‘d'N 28pug [eMeN € lWsdt ONWO SuWer A, IW uosonuo odonjun0N SNSWIDAdS: dO WaHWNN AWVN 104 MEMOIRS OF THE QUEENSLAND MUSEUM _ \ ; W GWA ARAL 1 FIG. 63. Naurilirepa omicron (Pfeiffer, 1851). a-b, Mt Warning N.P.. NSW. QMMO27290, neotype; c, f, Mt Glorious, SEQ. QMMQ16127; d-e, Kenilworth S.F., SEQ. QMMO9688, a-b, entire shell: c, marginal teeth; d, apical sculpture; ¢, post nuclear sculpture; f, central and lateral teeth, Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 105 c f FIG. 64. a-c, Gyrocochlea curtisiana (Hedley, 1912). a, Kroombit Tops, SEQ. QMMO16453; b—c, Springs Scurb, Mt Larcom, SEQ. AMC32990, holotype. a, terminal part of protoconch; b, c, entire shell. d-f, Nautiliropa omicron (Pfeiffer, 1851). d-e, Mt Glorious, SEQ. QMMO16127; f, Dandabah, Bunya Mts, SEQ, QMMO12283. d-f, entire shell. Scale lines as marked. 106 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 65. Nautiliropa omicron (Pfeiffer, 1851). a-g, h, Mt. Glorious, SEQ. QMMO16127. f, Dandabah, Bunya Mts, SEQ. QMMO12283. a, genitalia; b, details of penial complex; c, penis interior; d, spermatophore; e, hermaphroditic duct; f, talon and carrefour; g, ovotestis; h, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE AMC139761, 27 Apr 1958, L. Price); North Pine R,, SEQ (2. AMC139762, Lower. ex Cox); Tweed R., NSW (6, AMC139768, Petterd, ex Cox): Canungra, SEQ (1, AMC32989, S.W. Jackson); Pine R., SEO (2, AMC264. 1887, Musson); Goomburra S.F., Mistake Mts SEQ (27°S9’S, 152721 E) (f, AMC 128619, OM- MO1/1122, 16 Mar 1981, AM/OM - ABRS): c. 28.5 km from Goomburra, Goomburra S.F.. SEQ 600 m. CNVF/Palms (27°59'S, [52°21 E) (17. AMC136814, QOMMO12692. OMMO12698, 7 Dec 1981. AM/OM ~ ABRS). Condamine R. Valley, ¢ 9 km NE of Killarney, NSW, along stream (28"18°S, 152°22°E) (3, AMC128907, OMMOLN415, If Mar 1981. QM/QM ~ ABRS): Montville Rd. near Kondalilla Falls N.P. SEQ. vine scrub on roadside (26°40'S, 152°52°E) (5, AMC 136414. QMMO11983, 8 Sept 1982, AM/OM - ABRS); Clarence R,, NSW (3, AMC139758, ex Pel- ierd) ¢ 3.5 km NW Mt Pleasant,.on Dayboro - Mt Pleasant Rd, SEQ, dry and wet selerophyll (27°0S8'S, 152"44°E) (4. AMC136497, OMMO 12124, 3U Sept 1982, AM/QM - ABRS). ¢. 1 ko S Raynbird Ck, Ra, on Laceys Ck, Re. Laceys Ck. SEQ. open forest (277148. 152°43°F) (10. AMC136476, OMMO- 12090; AMC 136462, OMMOI121NO, 30 Sept 1982, AM/QM - ABRS): Benarkin S.F., neat Blackbolt. SEO (8, AMC 139757. Feb 1961, D.F, MeMichael); Dandabah. Bunya Mis N.P.. SEQ, behind picnic area (26°53°S, 151°36°E) (4, AMC136606, QMMOI122832. 31 Sept 1982. AM/QM ~ ABRS); Clarence R.. NSW (6, AMC 1397h6, ex Cox}; Tooloom Scrub, Beaury S.F.c. 8.3 km from Woodenbong, Legume Rd.. NSW, MVF (28°29°S, 192°24°E), (1, AMC128534. 15 Mar 1981, AM/QM. - ABRS); Cunninghams Gap N.P., SEO. near monument. MVF (2804'S, 152°24'E) (6, AMC128608, OMMO10991, 14 Mar 1981. AM/QM - ABRS): SW of Gympie, SEQ, NE slopes of Mt. Moorooreerai. remnant thickel amongst hoop pine plantation (26°16°S, 152°33°E) (7, AMC138 105, OMMO13354, QMMO13356, 7 Sept 1982, AM/OM - ABRS); S of Gympie, SEO, entrance fo 5,F, of Yabby Ck Rd near Dwyer Ck., reniiant vine thicket along gully (26°28°S, 152°38'"B) (4, AMCIAAITL, AMCI38112, OMMO13367, OMMO} 3369, 7 Sept 1982, AM/OM - ABRS); Richmund R.. NSW (1. AMC 139764, 1900.S.W. Jackson): Mt Warning NP, NSW, top of road in rainforest (28°24°S, 153°17°B) 7, AMC129304, OMMO10501, 19 Mar 1981) AM/OM - ABRS): Blackbull Range, ¢. 7.4 kat 5 of Benarkin. SEQ, MVF/Araucaria(26°53'S, 152° 11'B) (67, AMC!36564, AMCI36571, OMMO 12238, QMMO12245, 31 Sept 1981, AM/QM - ABRS); ORvilly’s, Lamington N.P.. SEQ (28°14°S, 153715 E) (6. O6MMO10439, 17 Mar 198). AM/OM - ABRS): Mt Warning N.P.. NSW. near base of Mt 107 (2824'S, [53° lo'E) (2. QMMOL0486, Mar 1981, AMIOM - ABRS): Toonumbar Forest Rd., Toonum- bur SB. ¢. 34 km from Kyogle NE NSW, CNVF (28°33°S. 152°45°E) (1. OQMMQO 10912. 14 Mar 1981, AM/OM ABRS); Koreelah S.F., NE NSW, on Acacia Plateau Kd e, 12,2 km trom Acacia Ck - Killarney Rd. NSW. 8350 m (28°217°S. 152°24'E}. SEVT/MVE/Arauearia (3. OMMOLOA94S, AMC (28561, 15 Mar 1981, AM/OM - ABRS); Mt Glorious. ¢, 3 km SE of summil, SEQ (27°20°S, 15246 EB) (3. OMMO1L991. OMMO11996, AMC1I36521, 2 Nec 1982, AM/QM - ABRS): Ponona 5.F., ‘SE of Pomona, SEQ (26°24°S. 152°54°R) (1, OMMO12006. 8 Sept 1982, AM/OM - ABRS 1982); MI Guyra N.P.. SEQ (25°49°S, 152°35°F) MVPINVF/Arancaria. (2, OMMO12074 7 Sept 1962, AM/OM - ABRS), Byron Ck crossing. Byron $.F., ¢, 8.3 km NW of Mt Pleasant. SEO (27°07 °S, 152°43'E) (2, OMMO1 2159, AM(C136500, 30) Sept 1982, AM/QM - ABRS), Cunningham s Gup N.P.. SEQ. ur monument (28°04°S. 152°24°F) (13, OMMO12677, AMCI30806. 7 Dee LOSL. AM/OM - ABRS). Mr Warning N.P., NSW (28°24°8. 153° 16° E), CNVF (1, OMM0112736, 135 Dee 19S. AM/OM + ARRS): S of Gympie, SEQ. entrance to Broploo S.F. on Gibber Road. MVF (232° 30"S, JA2"41°30"B) (1, OMMO13377. 7 Sept 1982. AM/OM -ABRS): Upper Booloomba Ck, Conandale S.F., SEQ (26°41'S, T5237 b) (1. OMMO 13386. & Sept 1982. AM/QM » ABRS): Boombona N,P.. Mt Nebo, SEO, (1, OMMO6071, 14 Nov 1976, MJ. Bishop); Dandabih, Bunya Mis N.P.. SEQ (27°53°8, 151°35°E) Araucuria/N VF, (20, OMMO6082, 5 Mar 1976, M.T. Bishop); Binna Burra. Lamington N.P.. SEQ, 850 m CNVE (2. OMMOs0S3, Apr 1976, M.J. Bishop): Manoring N.P., Mt Glorious, SEQ, wetsclerophyll (2, QMMO6263, Aug (976, MJ. Bishop); Mt Byron S.F.. SEO (27°UIS. 152"30"E) (2, O€MM06277, Aue 1976, MJ. Bishop), Mt Glorious, SEQ (27°2U'S, 152°4tV FE) (1, OMMO6293, Feb 1976, MJ. Bishop); Macdonald Park. Tumbotine, SEQ, NYP (3. OMMO6a06. 31 Aug 1976.M.J. Bishop); Boombana NP, Mt Nebo, SEQ, wet sclerophyll (27°23'S, 152°47'E) (1, OMMO6344. Aug 1976. MJ. Bistup): Fred's Rd, Mt. Mee SEQ. (27°05°S, 152°43'E) (LL. OMMO7335, 14 Apr 1980. J. Stanisic, N. Pall, A. Green); Kenilworth §.F., SEQ (26°40°S, 157°36°E) Arauwearia, (7, OM MO%6S5, 22 May 1980.1. Stinisic, A. Green}: Matala N.P., Mt Glorious. SEQ, CNVE (22. OMMO114604, OMMOTIAd7, 20 Jun LY&2, MSA party); Spicer’s Gap, SEO (1, OMMO11838, 7 Aug 1982, A, Rozetelds). Mit Nebo. SEQ. nr Jolly s Lookout, NVF (I OMM(Q12771, 2 Dec J982. J. Stunmic): Renilworth 27"S 29°S 152°E 151°E 153°E 154°E FIG, 66, Distribution of Nauriltropa omicron (Pteit- fer, (S51). S.F, SEQ, ¢, 0.3 km N of Little Yabba Ck on Maleny to Kenilworth Rd, SEQ (26°37'S, 152°42°E) (3, QMMQ 3435, 19 May 1983, K. Collins, J. Stanmsic): MI Glorious, SEQ, NVF (6, QMMO14165, 28 Nav 1983, J. Stanisic); Boombana N,P., M) Nebo, SEQ, NVF (1, QMMO1S5060, 5 Apr 1984, J. Stanisie); sietrack off Mt Archer Rd, Me Mee S.F., SEQ (27°04'S, 15241 E) (10, QMMO150I74, 6 Sept 1983, J.Stanisic. Db. Power); Fred’s Rd, Mt Mee, SEQ,, NVF (27°0S°S, 152°43"E) (2. OMMO15985, 28 Oct 1984, J, Stamsie, G, Annabell); Mt Glorious, SEQ., NVF (3, OMMO15984, 28 Oct 1984, J. Stanisic,G. Annabell); MI Glorious, SEQ, NVF (11, QMMO16127, 26 Jan 1986. J, Stanisic, J. Chaseling). Mt Tamborine, SEQ, Curtis Falls Circuit, CNVF, (1. QMM()16437, 23 Feb 1986, J. Stanisic, J. Chaseling): Upper Pine Ck,, Canungra, SEQ, NVF/Araucarta, (1, OMMO16547. | Oct 1986, J, Stanisic, D. Potter), Redwood Park, Toowoomba, SEQ, NVF (7. QMMO16572, 10 Sept 1986, J. Stamisic); Araucaria Track, Binns Burra, Laminglon N.P., SEQ. NVF (2, OMMO16578. OMMO16580. 27 Sept 1986, J, Stanisic); Tulluwallal Circuit. Binna Burra. Lamington N.P., SEO, NVF (7, OMMO16745. 2 Oct (985. J, Stanisic, D. Poller, J. Chaseling): Natural Bridge N.P., SEQ, on roudside c. 300 m N of entrance. SNVF (5, OMMOLO818, 29 Mar 1984, 1. Stinisic): Blackbull Range, SEQ., on side of Bruce Highway, MEMOIRS OF THE QUEENSLAND MUSEUM MVEA rauearia (31. OMMO 16826, 12 May 1984, J, Stunisic. D. Potter). Mt Glorious, SEQ. CNVF (4, QMMO 16832, 3 Apr 1984, J. Stanisic); Dundabah, Bunya Mis N.P.. SEQ. SVE (14, OMMO16847, 12 May 154. J. Stanisic. D, Potter): Big Falls Circuit, Bunya Mis.N.P.. SEO. SNVF (19, OMMQ16850, 5 Nov J985. J Stanisie, D. Potter); Cherry Plain - Wesicott Plain Circuit, Bunya Mts N.P., SEQ, NVE/MVF (16, OMMO16S856, 6 Nov 1985, J, Stanisic, D. Potter): Mt Hobwee Circuit, Binna Burra, Lamington NP. SEQ. NVF (2, O€MMO16865, 1 Oct 1985, J, Slunisig, DQ. Potter, J. Chaseling); Border Track, Binna Burra, Lamington N.P., SEQ, NVF (8, QMM(16870, 10 Mar 1984, J. Stamisic). Tullawallal Circuit. nr antarctic beeches, Binna Burra, Lumington N.P.. SEQ. NVF (1, OMMO16874, 11 Mar 1984, J. Stanisic); Border track, nr O' Reillys, Lamington N.P., SEQ, NVF (7. OMMO16876, 31 Mar (984, J. Stanisic); initial parLof Kweebini Caves Walk. Binna Burra, Lamington N.P., SEQ. NVF (4, QMMO16885, 9 Mar 1984, J. Stanisic); Kenilworth S.F.. SEQ (26°37'S, 152°42°E) (1. OMMO18530, 22 May 1980. J, Stanisic, A. Green); Mudgeeraba, SEQ, (3, QMMOT&S31, C.J, Wild); Cunningham's Gap, SEO. (4. Q€MMO18532. OMMO18534, 18 Jun 1979, G, Annabell); Mt Nebo, c. 2km east of summit, SEQ, (152"47 E, 27°24°S), NVEF/Araucaria, (1, OMMO1S533, 13 Aug 1980, J. Stanisic); Koondai-1 Lookout. Bunya Mts N.P.. SEO (26°53'S, 151°33'B), (1, OMMO18535, 2 Mur 1976, M.J. Bishop): Mary Cuirneross Reserve, Maleny, SEQ, (26°46°S, 52°51 EB). (1. OMMOTISS49, 22 May 1980, J. Siunisic, A. Green); c. 1.6 km N of Numinbah valley jurmoff on Burleigh to Springbrook Rd, SEQ, NVF with palms (1. QMMOI&SS0, 17 Dee 1980, J. Stanisic). Diagnosis Shell large, diameter 4,20-5.97 mm (mean 3.24 mm) with 4 to 4 5/8+ (mean 4 3/8+) tightly coiled whorls, the last very inflated and not des- cending in front (Fig. 644). Apex and carly spire (Figs 63a, 64d) strongly depressed, height 2.35- 3.11 mm (mean 2.82 mm). H/D ratio 0.48-0.57 (mean 0,54). Protoconch of | 3/8 to 1.5/8 whorls (mean | 3/8+) with mean diameter 597.5 xm at 1 1/2 whorls. Apical sculpture (Fig. 63d) of low, curved radial ridges and low spiral cords which are partially fused and worn producinga strongly pitted appearance, Post nuclear sculpture (Fig, 63e) of numerous. crowded, thin, protractively sinuated radial ribs. 72-167 (mean 113.6) ribs on the body whorl, Ribs/mm 5.02-9.79 (mean 6,90), Ribs with a high periostracal blade, often lost in dead or slightly worn specimens. Micros- SYSTEMATICS OF AUSTRALIAN CHAROPIDAF culpture (Fig. 63c) strongly reticulate, with fine microradials. 6-15 between each pair of major ribs and lower, crowded spiral cords which but- tress the microradials on either side, Umbilicus |Figs 63b, 64e) wide, U-shaped, diameter 1.23- 1.64 mm (mean 1.47 mm), D/U ratio 2.62—4.22 (mean 3.58). Sutures impressed, whorls rounded below and shouldered above a rounded periphery (Fig. 64f). Last whorl coiled more tightly. Aperture ovately lunate only slightly in- clined from the shell axis. Lip simple, columella not expanded but slightly deflected toward um- bilicus. Parietal callus well developed. Colour light-horn to light-brown, often with darker radial flammulations arranged in a zig-zag pat- tern. Based on 31 measured adults. Genitalia with epiphallus (Fig, 65b) mus- cularised. looped around the pemial retractor muscle before entering penis apically. Epiphallic pore (Fig. 65c) between two swollen pilasters. Penis (Fig. 65c) pear-shaped with a distinet api- cal bulb, separated from the lower penial cham- ber by a constricting muscular collar, Penial bulb internally with apical to slightly lateral epiphal- lic entrance and horseshoe-shaped pilaster, Lower penial chamber with a large apical pocket stimulator (Fig. 65c), and anastomosing spongy thickenings. Penial retractor muscle shart, inserting on penis apex adjacent to epiphallus. Vagina long internally with fleshy longitudinal thickenings. Spermatophore, simple, clavate. Radula as for genus. Based on five dissected adults (QMMO161 27. QMMO12283). RANGE AND HABITAT Highlands. of the Great Dividing Range be- tween Tooloom, NE NSW and the Bunya Mts, SEQ; the northern end of the Richmond Range; the McPherson Ranges south to the Tweed River, in the cast; and the coastal and adjacent ranges north to Mt Guyra, SEQ. N. omicron shows preference for notophyll vine forests but also occurs in adjacent microphyll vine forests, wet sclerophyll forests, and tn gallery rainforest along watercourses. NV. omicran lives.under logs, REMARKS N, omicron is one of the most conspicuous, widespread and frequently encountered of the subtropical microsnails. However most of the 637 specimens available were subadult reflect- ing the high proportion of mid-year, litter- collected matertal. 109 The modified radula of NV. omicron is unparal- Jeled in Australian charopids. Radulae of the large southern charopids Mulatena fordei (Brazier, 1871), Stenacapha hanultant (Cox, 1868) and Thryasona diemenensis (Cox, 1868) have typical central and lateral teeth (Smith and Kershaw, 1985) bul marginals show modilica- tions broadly similar to those in N. amicron. M. fordei and S, hamilton? have strongly slanted, slender. unicuspid marginals while 7. diemenen- sis has bicuspid marginals. In N. amicran the change from average charopid patterns may be the result of character displacement under con- ditions of microsympatry with the conchologi- cally similar Gyrocochlea. Specimens from eastern localities, in par- ticular those from the D’Aguilar Range, SEQ, have more crowded ribs than those from the Great Dividing Range (Table 14). These dif- ferences may be related to local environmental conditions. Letomola Iredale, 1941 Letomola Iredale, 1941a, p, 267. Letomala (error) Kershaw, L95Hb, p. 9. TYPE SPECIES Rhephodon contortus Hedley. 1924; by original designation. PREVIOUS STUDIES Iredale (19d1a) introduced Letomola as monotypic. Smith and Kershaw (1979) included the Tasmanian species ‘Helix’ barrenensis Pet- terd, 1879 without giving reasons. and it is here excluded. DIAGNOSIS Small shell (mean diameter 2.05 mm), with few (mean 3 7/8-) tightly coiled whorls. Spire (Fig. 67a) depressed. Apical sculpture (Fig. 67d) of irregular malleations and ridgelets. Postnuclear sculpture of weak growth ridges and vague spiral cords. Umbilicus (Fig. 67b) very wide, cup-shaped (mean D/U ratio 2.68). Sutures impressed. Whorls rounded below the periphery, flattened at the basal margin with a strong supraperipheral sulcus above (Fig. 67c). Aper- ture ovately lunate with one parictal and one basal barrier (Fig. 68a, b), Lip simple, margins convergent. Colour brown with lighter streaks. Animal with foot and tail broad, bluntly rounded posteriorly, Live animal dark-grey, particularly on the sides of the foot above the pedal grooves, 110 neck and mantle collar. Pallial cavity elongate with bilobed kidney, pericardial lobe reflexed. Genitalia with unusual features. Ovotestis a single, bilobed clump of follicles (Fig. 69c, d) embedded in the digestive gland. Terminal genitalia elongate. Epiphallus (Fig. 69a) a long slender tube, entering penis apically adjacent to an apical pilaster. Penis (Fig. 69b) slender with longitudinal pilasters. Penial retractor muscle inserted at the penis/epiphallus junction. Radula (Fig. 68d-f) narrow with few lateral and marginal teeth rows; central tooth very small, mesocone reduced; laterals tricuspid with much enlarged mesocone; marginals becoming multicuspid, serrate, with all cusps reduced in size. COMPARISONS L. contortus 1s conchologically distinctive with wide umbilicus (Fig. 67b), large, finely malleate protoconch (Fig. 67d), reduced adult sculpture (Fig. 67e, f), strong sulcus on the body whorl (Fig. 67c), and two apertural barriers (Fig. 68a, b). It is sympatric with Rhophodon kempseyensis (Fig. 77) which has a wide um- bilicus and brown shell with white streaks but differs in having a radially lirate protoconch, extremely crowded ribs on the body whorl (mean tibs/mm 33.34), and more numerous apertural lamellae. Letomola contortus (Hedley, 1924) (Figs 67-70; Tables 11, 14) Rhophodon contortus Hedley, 1924, p. 220, pl. 32, figs 35-37; Kershaw, 1955, p. 30. Letomola contortus (Hedley); Iredale, 1941b, p. 2; Iredale, 194 1a, fig. 6. Letomala (sic) contortus (Hedley); Kershaw, 1956b, p. 9. TYPE MATERIAL Lectotyre: AMC28494, Sherwood, Macleay River NSW. ex Technological Museum (= collected by C, Laseron) Height of shell 0.80 mm, diameter 1.81 mm, H/D ratio 0.44. D/U ratio 2.84, whorls 3 5/8-. PARALECTOTYPE: AMC150095, same data as lec- totvpe. OTHER MATERIAL Yessabah Cave, W of Kempsey, NSW (31°06'S, 152°42°E). on roof and walls of limestone cave and in floor litter (200+, AMC119349, 9 Oct 1979, P.H. Colman, J. Stanisic); Yessabah Cave, via Kempsey, NSW (31°05°30"S, 152°41°E) vine thicket, on lime- MEMOIRS OF THE QUEENSLAND MUSEUM stone rocks and in litter (54, QMMO17014, 3 Mar 1987, J. Stanisic, D, Potter, P.H. Colman), DIAGNOSIS Shell glossy, minute, diameter 1.81—2.24 mm (mean 2.05 mm) with 3 1/2 to 4+ (mean 3 7/8-) tightly coiled whorls. Apex and spire depressed with each whorl higher than the preceding whorl, except for the body whorl which descends in front, Height 0,80-1.03 (mean 0.91 mm). H/D ratio 0.40—0.51 (mean 0.45). Protoconch of 1 3/8 to 1 1/2- whorls, large, bulbous. Apical sculpture (Fig. 67d) of irregular malleations and ridges. Post nuclear sculpture (Fig. 67e) of broad radial ridges crossed by weak spiral cords, especially noticeable near the sutures. No additional microsculpture. Umbilicus (Fig. 67b) wide, shal- low, cup-shaped, diameter 0.64—-0.86 mm (mean 0.77 mm). D/U ratio 2.42—3.03 (mean 2.68). Sutures impressed, becoming a deep furrow as the last whorl descends. Whorls rounded below the periphery with a prominent supra-peripheral sulcus. (Fig. 67c). Aperture irregular ovately lunate with one parietal and one basal barrier. Basal barrier more deep-seated than parietal. Parietal barrier (Fig. 68a) extending 1/8 whorl, horizontal anteriorly, becoming angled downward posteriorly. Basal barrier (Fig. 68b) 1/8 whorl in length, following angle of coiling, tapered at either end. Lip simple, columella only slightly dilated, margins convergent. Parietal callus weakly developed. Colour deep reddish- brown with irregular white radial streaks, brown on the base. Based on 13 measured adults. Epiphallus-vas deferens junction valvular. Epiphallus (Fig. 69b) entering penis apically. Penial retractor muscle inserted on the epiphal- lus-penis junction. Penis (Fig. 69b) a long tube with a slightly expanded apex. Epiphallic entrance a simple pore adjacent to a short tongue-like pilaster. Penis proper sculptured with longitudinal pilasters. Free oviduct and vagina long tubes without unusual features. Radular characteristics as given in generic diagnosis. Based on 6 dissected specimens (AMC- 119349, QMMO17014). RANGE AND HABITAT From the limestone outcrop at Yessabah, west of Kempsey, NSW where the vegetation is semi- evergreen vine thicket. In spite of several collect- ing efforts it has not been found further westward in the limestones of the Carrai State Forest. L. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 111 FIG. 67. Letomola contortus (Hedley, 1924), a—c, Sherwood, Macleay R, NSW. AMC119349. a-c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, sculpture on base. Scale lines as marked. 112 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 68. Letomola contortus (Hedley, 1924), Yessabah Cave, W of Kempsey, NSW, AMC119349, QMMO17014. a, parietal barrier; b, basal barrier; c, microdenticulation on basal barrier: d, radulay e, central and lateral teeth; f, marginal teeth, Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 113 SS FIG. 69. Letomola contortus (Hedley, 1924). Yessabah Cave, W of Kempsey, NSW. AMC119349. a, genitalia; b, penis interior; c, d, details of ovotestis; e, hermaphroditic duct; f, talon and carrefour; g, pallial cavity. Scales lines as marked. lid ST#e / Soccer eB ~ 9 7 a ; 2S . | SQN Safe FIG, 70, Distribution of Letomola contortus (Hedley, 1924), contorts has been collected live on limestone tocks after rain and presumably feeds on the microflora growing on the rock surface. REMARKS L. contortus displays a number of departures from typical charopid patterns. The malleated protoconch (Fig, 67d) and supraperipheral sul- cus (Fig. 67c) are unusual features in the context of the east-coast species reviewed herein. Sulci are seen in some Pacific [sland Endodontidae and Charopidae, and according fo Solem (1983, p. 9) they are usually associated with keel forma- tion. In “. contortus the sulcus indirectly con- stricts the size of the aperture. Anatomically the slender reproductive organs are more lypical of the Punctidae. However, the radula is typically charopid even though the strong reduction in size of the central tooth and serrate appearance of the marginals are unusual. Omphalorepa varicosa has a grossly similar radula ~ few (eth per row and reduced central tooth. Rhophodon Hedley, 1924 Rhophodon Hedley, 1924, p. 219; Iredale, 1937a, p. 329; Tredale, 1941b, p. 2; Kershaw, 1955, p,30: Bureh, 1976, p.133, Egilodonta \redale, 1937a, p, 328; Kershaw, 195ha, p. 142; Burch, 1976, p.132. MEMOIRS OF THE QUEENSLAND MUSELIM TY?PF SPRCIES Rhophedon peregrinus Hedley. 1924; by onginal designation. PREVIOUS STUDIES Hedley (1924, p, 219) introduced Rhaphadon for charopids with apertural barriers. Apertural barriers among Australian Charopidae are rela- tively rare but several generic groupings have been proposed including Dentherona Iredale, 1933 (\vpe species: Helix dispar Brazier, 1871); Bischoffena Iredale, 1937 (type species: [Helix hischoffensts Petterd, 1879); Egilodonta Iredale, 1937 (type species: Charopa bairnsdalensis Gabriel, 1930); and Letomola Iredale 1941 (type species: Rhophaden contortus Hedley, 1924). Rhophodon is expanded to include seven species of very small to minute charopids with apertural barriers and shells which have a predominantly radially lirate protoconch, post- nuclear sculpture of prominent radial ribs, and a wide saucer-shaped umbilicus. DIAGNOSIS Very small to minute Charopidac, diameter tange 1.29-2.98 mm. Whorls 4 1/4 to 5 1/2-, tightly to very tightly coiled, Jast descending more rapidly, Apex and early spite depressed (Kempsevensis, peregrinus) to very slightly elevated, Protoconch exsert of 1 1/2 to 1 3/4 whorls. Apical sculpture of numerous moderately crowded to very crowded (more so toward the protoconch - adult whorl boundary) weakly curved radial ribs and very weak thread- like spiral cords and cordlets, Post-nuclear sculpture of few, widely spaced (colmani, bairnsdalensis) to many, crowded (kemipseyen- sis), thin, protractively sinuated to straight radial ribs, Micrasculpture of numerous, crowded radial riblets, and widely spaced (kempsevensis) to crowded, low rounded spiral cords which are raised at their intersection with the radial riblets to form an elongate bead. Umbilicus wide. cup- shaped. Aperture ovately Junate, with few (bairnsdalensis) to many (consobrinus) barriers. Parietal barriers consisting of one to four (generally three) eresventic lamellae. with or without additional thread-like traces. Palatal barriers three to six (usually four) crescentic lamellae, with or withoul a Supertor trace, Columellar lamellae sometimes present. Pallial region elongate with almost unilobed kidney in which the pericardial lobe is well developed, reflexed ut the tip, and the rectal lobe is reduced to a vestige abutting the hindgut. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE Ovotestis two small clumps of finger-like lobes embedded. in the subapical whorls of the digestive gland. Terminal male genitalia with epiphallus entering the penis apically or subapi- cally (consebrinus) through two fleshy lips sur- rounded by a museular collar. Penis with an apical bulb containing epiphallic pore, some- Imes expanded and with longitudinal pilasters (consabrinus). Penis with (peregrinus, con- sobrinus, kempsevensis) or without a sheath, ins ternully with small, fleshy, longitudinal pilasters and basal, circular, stimulatory pads (con- sobrinus, peregrinus), Penial retractor muscle inserting at the penis/epiphallus junction. Female reproductive system elongate with long vagina and free oviduct, Radula without unusual features except in kewtpseyensis. DISTRIBUTION AND ECOLOGY Rhophodon is lound from Bairnsdale, Vic- toria, to the Bulburin State Forest, near Miriam Vale, SEQ. Species have very small tanges, Some are known only from closed forest but uthers live in fringe rainforest areas and lime- stone refugia. R. peregrinus and R. cansabrinus are sympatric in the eastern McPherson Ranges, SEQ, and some adjacent areas including the coastal vine thicket at Burleigh Heads, SEQ, Differences in penial morphology between the two species are probably species recognition features, Species live in a Variety of microhabitats and have been collected from leaves in litter, under logs, atid from moss and Jitter accummulated on rock surfaces. R peregrinus has also been found in an Aspledivm fern on a tree. PATTERNS OF SHELL VARIATION Size and Shape - Rhophaden includes some af the smallest charopids found in castern Australia with mean shell diameters 1.45-2.66 mm. The largest species, R. kempsevensis, is confined to the limestone oulcrop between Yessabah and Carrai, westot Kempsey, NSW, where the readi- ly available calcium probably enables greater size, Otherwise size variation appears to be mosaic in nature, Height is correlated with diameter (mean H/D ratio for the seven species ranges U.40-).46) and is reflected in conservative shell shape, Spire protrusion ts mimmal. Body whorl width shows some reduction in R, calmani and R. minutis- simas correlating with small size. Mean whorl counts range from 4 1/2-105-. Species with the Jargest whorl counts are K. kempseyensis; R. bairnsdalensis {rom limestone habitat in southern NSW; and R, elizabethae from ealeium tich araucarian Vine forests of SEQ. Sculpture ~ Shell sculpture is complex but variation is relatively simple, The most sig- nificant differences are in the number and spac- ing of the radial ribs and it is possible to divide the species into three groups on this basis - few, bnid, widely spaced ribs (R. bairnsdalensis, R. colmant, Figs 87a, 89a): few, narrower, moderately spaced ribs (R. minulissimus, R. el:zabeiliae. Figs 81a, $4a); and very many. fine, extremely crowded ribs (R. peregrisnus, R. con- sobrinus, R, kempseyensis, Figs 71a, 74a, 77a). Dillerences in microsculpture are minor with the more widely spaced microspiral cords of R- kempsevensis (Fig. 77e) being an extreme modification which may relate (o the larger whorl size of that species. Minor differences occur in number of apical ribs and intensity of apical cording. Aperturdl Barriers - Variation in the number of parietal barriers ranges from a single elongale barrier (Fig, 90a) to six barriers and traces (Fig. Tig). The average pattern is for three elongate barriers, expanded posteriorly and deflected downwards. The average pattern of palatal barriers is for four major harriers to which a superior (race may be added, Major deviations-are a single barrier and trace (Fig, 90b, c), three barriers (Fig, &4h) and six barricrs (Fig. 87a). Jo general the bar- riers are short crescent-shaped lamellae. In &, colmenit (Fig. 87g) the barriers are expanded with greatly clongate anterior blades. A columetlar barner may be present and is usually short crescent-shaped and may be sup- plemented with an accessory trace (Fig, 747), PATTERNS OF AN ATIUMICAL VARIATION Stability of the Rhaphoden morphotype, which is evident in the conchological patterns. extends lo the analomy, Apart from differences related to species size e.g. relative lengths of the genital and pallial structures, the most notable variation occurs in the male terminal genitalia, Male genitalia - Typically the penis has an apical bulb which contains the epiphallic entrance and a main penial chamber with few to several fusiform, longitudinal, fleshy pilasters. Exceptions are R. consobrinus (Fig, 75e) and R. peregrinus (Fig. 72b) which have altered penial surfaces, These nwo species are sympatric and ‘the differences in penial morphology are probab- ly species recognition features. The presence of Llo ‘TAULE 1S - RANGE OF VARIATION IN RHOLMODON AND DISCOCHAROPA (MEAN AND RANGE) D/U RATIO UMBILICAL. WIDTH RIDSIMM RIBS SrBWW RANO WIDTH HODY WHORL (mm) SHIRE HELUNT DIAMETER HD RATIO PROTRUSION WHORLS (mm) NUMBER OF SUECIMENS NAME (mm) (mm) (mm) MEMOIRS OF THE QUEENSLAND MUSEUM > a= = = 4228 & EG nN iad ny m ia moN Loe) echepotedetsotndre BYRAARS SRE ARoS= NAAN ES ren og ree rd SNS rN a edn — mt om = = a2 = e233 Ee sess - & «s = ; = = g z oe, Oe = x Bs a ae zi oBehes 208 SSRI ert eoasnx G2 es ASASRS, ime os = = Bere Sse ez SUBIETOTSISIUIes Sends betetotce gdgacseredensass Sota Tor atardss = = | api =-s a= scene 5 = = cans = ZSre secs s 2 = = aS 2 22:3 i wei 22 iss Ce (SS=sH5 (1,792.22) 224 (1.95-2,42) is T4095) fh 0. A S/R (4 VBS Vide) 4172+ (4b 57m) 312 4-3/4) 4/2. (4 Vid-tod 5/B) 4a. (484105 1/-) 5. (4 f/B-5 44) (4 5/H-S T/R4) 5 (4 3/tH05 1/2-) Rhophodon peregrinus turtrrescdelervis Miscovkarepa aperia a penis sheath correlates with large size and greater intensity of radial ribbing. The epiphallus is normally long and muscular but in R. kemp- seyensis (Pig. 79a) it is very long, correlating with increased whorl count, Radula - The Rhophodon radula shows little interspecific variation reflecting overall similarity in species microhabitats. Size dif- ference between the central tooth and laterals is exaggerated in R. kempseyensis (Fig. 78e). The large broad lanceolate mesocone of the laterals is a feature seen in other charopids (Figs 68e, 123h) sympatric with R. kempseyensis and ap- pears to be correlated with life on limestone. COMPARISONS Rhophodon differs from the conchologically similar Egilomen Iredale, 1937 in having aper- tural barriers and in details of microsculpture, The microspiral cords in Egilomen are more prominent than those of Rhophedon and are crossed by fine thread-like radials contrasting with the high microradial ribs of Rhophodan. Anatomically the pallial configurations of Rhophodon and Egilomen are distinct. The strongly bilobed condition of the Egilomen kid- ney contrasts with the almost unilobed kidney of Rhophedon. Although the penial complexes of the two genera are grossly similar, the tubular penis and stout muscular epiphallus of Egilomen are important differences from Rhophadon. KEY TO SPECIES OF RHOPHODON 1.Ribs on body whorl very widely spaced (Figs Ribs on body whorl more crowded ........., 3 2.Aperture with three swollen parietal barriers (BATT csecgecapice top ge petaees tense nate colmani Aperture witha single lamellate parietal bar- rier(Fig. 90a) batrnsdalensis 3,Major radial ribs bold, moderately spaced ..... 4 Major radial! ribs finer, very crowded ,..,.......5 4.Shell very small (mean diameter 1.52 mm); second parietal barrier much shorter than first DEVI T Fai. e'acsiestenlsaigs vs tplgesieeslaslen MUNUUSSTINIUS Shell larger (mean diameter 2.47 mm). flam- mulated: Second parietal barrier elongate 117 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE (S9'2-ZE°7) (eri-zon) OSWUFOP'E BZOOFLO'L rasra so LOT 601 9LZ 660 (pp 2-177, (gz I-e0"1) ECUURTET Sz0'0=F I'L “ez aa (rR z-£0°2) NUOUFLE TS prUOFi6 Suz ZK) Lz SRO Tez aL0 Ls@ v0 652 red Ltt 060 (itera ze0 SZ oso (9p'z-REZ) (06°0-08'0) oroorzre osoursyO (Os‘z-zz'z) (98'0-FL'0) OLvUFPeZ eszorOR'0 EP'z zr’ (ui) Ola va HLGIM AVON (12'01-60'8) ETEUFLYR £679 Oy L £98 (6 REFEY'NZ) OLY FCP TE l@le (eR rZ-90' 12) OA LFER'TZ Ire It £691 Leu 641 LR'SZ SB'RL fF 07 Sa'pz (op OL-1F'b2) TOS OFZELZ 86'9Z WWSfla “AON ‘dS FVLAA VZITA NOGOMAOHY ONY (7-29) (6V'0-010) SUEFSLO SIV OFF zs Wo 9S 900 7a so (zig-nE7z) SUSIFS'ELZ a oe <— (9RI-SS1) Hv AFR LY SOL £00 stl sv vil 900. 601 ) vt ev £91 a zu oe 11 = Pal = (6gt-spi) £s'pep'091 = 641 ces Sau OLLVa MMEVdS (gR'0-#9°)) £TVUFRLO HOOF O “Ly Aud LL0 roo SRO P00 (880-480) SOVUFSH “_ ran) <= (Lt) (row) TIVOFTLE 7 we0 ao 0s) rod ROD roo rot rate) L4) m0 #90 -— yo" ~es aL = (ov0-49'0) O10FS9'0 _ (0L°0-29'0) LO00F99) — 20 ~~ (uuu) (ww) HLCIAN NOISNY.LOUd MOH ACOs AMS (AONVU ONV WAS 'NVAA) (apt) HOW OFSEO orn zr 1rO (ep O-ke') oro raal) tru wo PO tra tro (Lp‘0-1¥'0) 900 0F EPO fro OLLYY C/H (p9'2-6E°7) OS) UF REZ 6ET Cara YLT (ao C-LE2) ERrRso Te “LT (we t-92'C) SEOUFEDT Lira RZ SOT 6k" 02 (277-16 D SST OFLOT (£61611) OO UFLEL 66'1 (wu) MALLAW VIC (pz"1-86'0) REMOFZUT wh (LV 1-€0'1) FIO UF LOT £ (ee'0-So'0) LU UF YEU t60 401 BRO THO peu oro PRO (sa'0-zR'0) SOU'0F68'0 (98'0-4L'0) ZO UFOs'O ORO (wi) LUOJTH (B/L S—p/E b) s WS tb +P/1S CH/L S-H/L PD s WS b WLP Uir +WEP WSb HST tHE P WS (+p/t S-A/L ») aa (p/E p-B/E p) HSE “WL STYOHM 9 fasyt OWWO i 85891 ONWO if LILylLOWWO {ad AinjoH) ' SIEL| OWWNO aduey Aqqogy amyjaynzya uypoydoyay ar S10L1 ONWO (ad oj) L 9LOLLOWWO yay [RAMEN sisuadavdway uapoydoyy c #rS91 OWWO yaa aug aaddyy 1 fort OWWNO “Wa qos Poy i SShOLOWWO i} TarOL OWNO TUE A 1 1X69 OWWO spray Yalapng (adAiojaa7q) 1 77661 OWY JAALY PUOWYDIY, snuiagosuos uopoydoyy i E601 ONNO qriag WOKTOO], 1 95601 ONWO. “¥°S yRpaUOY 1 S6s9l ONWO SAY.O. z 5919 OWWO Spray ysiapng a a8fks6 WV (edAjoan) if 6ILESl WY JIALY PIOM I, snundasad wopaydoyy SNIEWIDAdS AO Wag WAN AWVN ‘AON ‘dS SISNTANSU WAN NOCOHAOIA $76 “AT VOIH SONRIGOSNOD NOUONAOIY "e761 ATIOIH SON OAAAd NOCONMdOLR NI NOLLY PAYA W207 - 91 TTYL 118 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 119 FIG. 72. Rhophodon peregrinus Hedley, 1924. On border track c. 6km from O’Reillys, Lamington N.P., SEQ. QMMO16595. a. genitalia; b, penis interior; c, ovotestis: d, hermaphroditic duct: e, talon and carrefour region; f, pallial cavity. Scale lines as marked. FIG. 71. Rhophodon peregrinus Hedley, 1924. a—c, Tweed River, NSW. AMC153719, neotype; d, c. 6km from O’Reilly’s on border track, Lamington N.P., SEQ. QMMO16595; e-h, Tweed River, NSW. AMC- 95838. paratype. a—c, entire shell: d, marginal teeth; e. apical sculpture: f, post nuclear sculpture; g, parietal barrier: h, palatal barriers. Scale lines as marked. 121) with an anterior taper: Bobby Range. SEQ otiokeetosnelionenrepaserereabreneendecnesnenstiett elizabethae S.Mayor radial ribs extremely crowded (mean ribs/mm 33.34); shell larger (mean diameter DGG MM) wees senses eeseneeseee cts REMPSEVONSTS Major tadial ribs less crowded (mean Tibs/mm < 30.00); shell smaller (mean Ciameter< 2,40 MM). ...cccsesceeeersersererpenvenees 2 6.Parietal barriers consisting of four lamellae and two traces (Fig. 71g); three palatal barriers (Fig. TUM). psscccsccscenenessteecesteeesrses PEPE RFINUS Parietal barriers consisting of three lamellae, and rarely, two traces (Fig. 74h); five palatal Jamel lac (Fig. T41)...cc.csee seen, CONSODINUS Rhophodon peregrinus Hedley, 1924 (Figs 71—73; Tables 15, 16) Rhoaphodon peregrinus Hedley, 1924, p. 220, pl, 32. figs 38-40; Iredale, 1937a, p, 329; Iredale, 194 1a, p. 268, fig. 6: Iredale, 1941b, p. 2. COMPARISONS R. peregrinus is conchologically similar to R. consobrinus but is distinguished by the more crowded ribs and slightly depressed spire and greater number of parietal barriers. The two species are sympatric and have quite different penial morphology. These penial surface dif- ferences are effective species recognition fea- tures. R. kempseyensis is larger with relatively greater number of whorls and almost twice as many ribs on the body whorl (Fig. 77a). Other Rhaphodon species have markedly fewer ribs on the body whorl than R. peregrinus (Table 15). PREVIOUS STUDIES Although Hedley (1924) referred to a type specimen collected by W. Petterd in the Cox collection, the only “type” material of RK. peregrinus located in the Australian Museum was alotof 38 specimens (AMC95838) from the type locality, labelled paralectotypes although this designation has no basis in literature. Also in the Australian Museum type collection was a vial labelled "Rhophodon peregrinus ... Type lost ...". TYPE MATERIAL NEOTYPE AMCIS53719 (ex AMC95838), Tweed MEMOIRS OF THE QUEENSLAND MUSEUM 154° FIG. 73. Distribution of Rhophodon peregrinus Hed- ley, 1924, River, NSW. Collected W. Pellerd, ex Cox Coll. Heightof shell 0.86 mm, diameter 1.99 mm_H/D ratio 0.44, 0/U ratro 3.43, whorls 4 7/8-, OTHER MATERIAL AMC95838%, 37 specimens, same collection data as neolype: Burleigh Heads N.P., SEQ, vine thicket, litter (1, QMMOLI41I6, 6 Jan 1984, J. Stanisic; 25, QM- MO6165, Oct 1976, MJ. Bishop): ¢ 6 km from O'Reilly's, on border track, Lamington N.P., SEQ. NVF, on racks (7, QMMO16595, 31 Mar 1984, J, Stanisic); Tooloom Scrub, Beaury 5,F., NE NSW (28°29'S, 152°24E) NVF (2. OMMO10938. 15 Mar 1981, AM/QM - ABRS); Burleigh Heads (5, AM- C55506, ex J. Brazier); Byron Bay scrub, NSW (2, AMC63847, Lower, ex Cax); Beaury S.F., c. 15 km W of Urbenville, NSW 810 m (28°21°S, 152°24°E) (27, in litter and in Asplenium on tree, AMC152160, AMC152161, 18 May 1976, P.H. Colman, [. Loch); Koreelah S\F., c. 12,2 km from Acacia Ck - Killarney Rd, NSW SEVT/MVF (28°217S, {52°24'E) (4, QM- MO10956, AMC128559, 15 Mar 1981, AM/QM - ABRS); Richmond Ry. Lismore, NSW, in scrub under decaved leaves and dead timber (3, AMC63872, ex Brazier). DIAGNOSIS Shell minute, diameter 1,.79-2.22 mm (mean 1.93 mm) with 4 3/8 to 5 1/4+ (mean 4 5/8+) tightly coiled whorls, the last descending more rapidly. Apex and early spire depressed to strongly depressed. Height of shell 0.744+).95 mm (mean 0.85 mm). H/D ratio 0.41-0,47 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE (mean 0.43). Protoconch exsert, 1 1/2 to 1 3/4 whorls, mean diameter 410.9 1m at 1 1/2 whorls. Apical sculpture (Fig. 7le) of curved, broad radial ribs and weak spiral wrinkles, ribs becom- ing more crowded toward the nuclear/post nuclear junction. Postnuclear sculpture (Fig. 71f) of numerous, very crowded, high, slightly protractively sinuated radial ribs, 112-189 (mean 156.9) ribs on the body whorl. Ribs/mm 18.85-30.44 (mean 25.93). Apices of ribs with spiral sculpture. Microsculpture (Fig. 71f) of fine radial riblets and conspicuous low spiral cords which are raised at their junction with the radial riblets producing a beaded effect. Um- bilicus (Fig. 71b) wide, cup-shaped, diameter 0.74-0.90 mm (mean 0.82 mm). D/U ratio 2.22- 2.54 (mean 2.36). Sutures impressed; whorls shouldered above and rounded below a weakly rounded periphery. Aperture ovately lunate. Numerous barriers present. Parietal barriers (Fig. 71g) consisting of four crescent-shaped lamellae and two thread-like traces. First barrier large, deflected upwards; second, third and fourth without marked anterior taper, directed downwards. Superior and inferior traces may be present. Palatal barriers consisting of three cres- cent shaped lamellae (Fig. 71h) and a superior trace (Fig. 71g) high up on the palatal margin. Columellar barrier small, thread-like. Lip simple, columella not expanded or reflected. Parietal callus developed. Colour white to straw-yellow with brown radial streaks and suf- fusions. Based on 16 measured adults. Genitalia with vas deferens/epiphallus junc- tion (Fig. 72a) complex, valvular. Epiphallus (Fig. 72a) a muscular tube continuing to the penis head where its coils before entering the penis apically. Penis (Fig. 72a) an almost cylindrical, muscularised tube with a thin sheath and head slightly expanded, tapering as it nears the atrium, internally with a simple epiphallic entrance (Fig. 71b) surrounded by a muscular collar. Interior of the penis with large, apical, longitudinal pilaster, a basal doughnut-shaped pilaster, and general wall sculpture of spongy pustulations. Radula (Fig. 71d) typical. Based on 5 dissected adults (QMMO16595). RANGE AND HABITAT Cool subtropical notophyll vine forests of the McPherson Ranges, SEQ, westward to the Great Dividing Range on the NSW/QLD border; coas- tal rainforests at Byron Bay, NSW and Burleigh Heads, SEQ. An early record at Lismore, NSW, needs to be confirmed. R. peregrinus has been collected in litter, among moss on rocks, among accumulated debris, and unusually, in an epiphytic fern growing on a tree. REMARKS Lack of material makes a study of interpopula- tional variation impossible at this stage. How- ever, a single specimen (Table 16) from the Koreelah State Forest, NSW, in the western part of its range, differs in rib spacing from speci- mens further east. Rhophodon consobrinus Hedley, 1924 (Figs 74-76; Tables 15, 16) Rhophodon consobrinus Hedley, 1924, p, 220, pl. 31, figs 32-34; Iredale, 1937a, p. 329; Iredale, 1941b, p. 2. COMPARISONS R. consobrinus can be distinguished from R. peregrinus by its modified penis which has the apical bulb reflexed and expanded to include longitudinal pilasters (Fig. 75e). Conchological differences are minor except for the disposition of apertural barriers. R. consobrinus (Fig. 74h) has fewer parietal barriers than R. peregrinus (Fig 71g). R. kempseyvensis (Fig. 77a), has al- most twice as many ribs on the body whorl, is larger and has a higher whorl count than R. peregrinus. Other species can be differentiated by their smaller rib counts (Table 15). TyPe MATERIAL LECTOTYPE: AMC19922, Richmond River, NSW. Presented W.F. Petterd. Height 0.95 mm., diameter 2.18 mm., H/D ratio 9.43, D/U ratio 2.59, whorls 4 5/8. OTHER MATERIAL Burleigh Heads N.P., SEQ, vine thicket (7, QMMO16581, 29 Sept 1986, J. Stanisic, J. Chaseling; 2, QMMO6160, Nov 1976, M.J. Bishop; under logs, 7, QMMO16922, 2} Jan 1987, J. Stanisic, D. Potter): Upper Pine Creek. Canungra, SEQ. NVF/Araucaria (on rocks, 20, QMMO16548, 1 Oct 1986, J. Stanisic, D. Potter: on moss covered rock, 7, QMMO16906, 21 Jan 1987, J. Stanisic, D. Potter: under logs, 7, QM- MO16586, 29 Sept 1986, J. Chaseling, J. Stanisic); Binna Burra, Lamington N.P., SEQ. (28°12'S, 153°11°E) NVF/Palms (1. QMMO10446, 18 Mar 1981, AM/QM - ABRS); Red Scrub Flora Reserve, Whian Whian S.F., NSW, 210 m (28°38’S, 153°19°E) (1, AMC152157, 15 May 1976, P.H. Colman, T. 122 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 123 FIG. 75. Rhophodon consobrinus Hedley, 1924, a—e, g, h, Upper Pine Ck, Canungra. SEQ, QMMO16548, QMM0O16586, QMMO16906. f, Burleigh Heads N.P.. SEQ. QMMO16581. a, genitalia; b, vas deferens - epiphallus junction; c, details of penis - epiphallus junction: d, ovotestis; e, penis interior; f. spermatophore; g, talon, hermaphroditic duct and carrefour region; h, pallial cavity. Scale lines as marked. FIG. 74, Rhophodon consobrinus Hedley, 1924. a-c, Richmond River, NSW. AMC19922. lectotype; d-e, h-i, Upper Pine Creek, Canungra, SEQ. QMMO16906, QMMO16548; f-g. Natural Bridge N.P., SEQ. QMMO0O10455. a-c, entire shell; d, central and lateral teeth: e. marginal teeth; f, apical sculpture; g, post nuclear sculpture; h, parietal barriers; i, palatal barriers. Scale lines as marked. In 154°r FIG, 76. Distribution of Rhophodon consebrinus Hedley, 1924, Loch); Rocky Ck., Red Scrub Flora Reserve. NSW (2838S, [53° L9°B) rainforest (5, OMMO10403, AMC128864, AM/QM - ABRS): Tyagarah (1, AM- C152158, Lower. ex Cox); Tweed River, NSW (3, AMC5310, ex Cox): Cape Bryon, NSW (1, AMC 140480, 1 May 1958, L. Price), upper Tweed River, NSW (1, AMCIS52159, Petlerd, ex Cox); Natural Bridge N.P..NSW, NVF (28°13'S, 153°14'B) (QMMO1045S, AMC 129268, 18 Mar 1981, AM/QM - ABRS): Richmond Ry, Lismore, NSW, in serub under devuyed leaves and dead timber (1, AMC63872, ex Brazier), DIAGNOSIS Shell mimute, diameter 1.95—2.43 mm (mean 2.35 mm) with 4 3/8+ to 5 1/8- (mean 4 3/4-) tightly coiled whorls, the last descending more rapidly. Early spire and apex weakly to moderately depressed (Fig. 74a). Height of shell 0.82—1,07 mm (mean 0.94 mm), H/D ratio 0.40- 0.44 (mean 0.42). Protoconch exsert, of 1 1/2 to 1 5/8 whorls, mean diameter 457.0 jvm at 1 1/2 whorls, Apical sculpture (Fig. 74f) of curved, broad radial ribs, becoming more crowded toward the nuclear/postnuclear shell junction, and weak spiral wrinkles. Postnuclear sculpture (Fig. 75g) of numerous, crowded, broad, slightly protractively sinuate radial ribs, 109-186 (mean 143.4) ribs on the body whorl. Ribs/mnt 16.93— 24,83 (mean 20.30). Apices of ribs rounded and sculptured with radial lines. Microsculpture (Fig. 74) of fine radial riblets, 5—9 between cach pair of major ribs, and very low spiral thicken- MEMOIRS OF THE QUEENSLAND MUSEUM ings, Which are raised at their junction with the radial riblets to form elongate beads. Umbilicus (Fig. 74b) wide, cup-shaped, diameter 0.76-1.07 min (mean 0.88mm), D/U ratio 2,23-2.82 (mean 2.56). Sutures impressed. Whorls (Fig, 74c) rounded below and shouldered above a slighily compressed periphery. Aperture ovately. lunate. Numerous barners present, extending posterior- ly for about 1/8 whorl. Parietal barriers (Fig. 74h) three. long, crescent-shaped lamellae, and several, variable, thread-like traces. First and second parietals large. crescent-shaped, second deflected downwards; third low, directed downwards. Parietal traces may occur above the lirst parietal lamella and between the first and second purietal. Palatal barriers (Fig. 74c) five, the fifth crescent-shaped almost touching the first parietal: fourth, almost equal in size to the fifth, also crescent shaped: third smaller, eres- cent-shaped, sometimes bifid apically; second and first basal, thin, small. A superior palatal trace may be present at the parieto-palaltal mar- gin (Fig. 74h), Columellar lamella small, Lip simple, columella not expanded or deflected. Parictal callus developed.. Colour light straw- yellow with reddish-brown radial streaks and suffusions. Based on & measured adults, Genitalia with penial retractor muscle inserted at ecpiphallus/penis junction (Fig. 75a). Penis (Fig. 75e) tubular. reflexed c. 1/3 way along its length, with a thin sheath, and complex internal sculpture. Epiphallus entering penis through wo thickened lips (Fig. 75) adjacent ta several high longitudinal pilasters. Remainder of penis chamber wall with broad low pustules giving way to low thinyidges basally. Base ot the lower chamber with a stimulatory doughnut-like pad, Spermatophore (Fig. 75f) hook-shaped, Radula (Fig. 74d.c) similar to that of R. peregrinus, Based on 13 dissected specimens (QMMO- 16548, OMMO16581, QMMO16906, OMMO- 16922, QMMO16586). RANGP AND HABITAT Moist, warm lowland notophy!l vine forests of the Big Serub area, northern NSW and eastern McPherson Ranges and the coasial vine scrubs at Byron Bay, NSW, and Burleigh Heads, SEQ. R, consobrinus has been collected in litter, under logs and on moss covered rocks, REMARKS Although R. peregrinus and R, consebrinus have not been found microsympatrically, their SYSTEMATICS OF AUSTRALIAN CHAROPIDAE overlapping ranges and convergent microhabitat preferences make this a distinct possibility. Cer- tainly, penial differences suggest microsym- pat ry. Rhophodon kempseyensis sp. nov. (Fig. 77-80; Tables 15, 14) ETYMOLOGY For the Kempsey region, COMPARISONS This. species is recognizable by its lustrous golden brown shell which has numerous, ¢x- tremely crowded radial ribs. It has the highest rib count of any east Australian charopid. The species closest to R. kempseyensis are RK. peregrinus and R. consobrinus but they have only c. half the number of ribs on the body whorl,, are smaller, and have relatively fewer whorls (Table 15). Letemola contortus which is sym- patric with R. kempseyensis, has a wide um- bilicus and two apertural barricrs - one basal and one parjetal (Fig. 68a, b), Sculpturally /., con- rortus differs from R. kempeverisis in having a more bulbous protoconch (Fig. 67d) with ir regular malleations and ridges, and adult whorls (Fig. 67e, f) with low, broad radial ridges. and weuk spiral cords. TYPE MATERIAL HoLoryre: QMMO17016, Natural Arch, Carrai S.F., NSW. 30°54'S, 152°21'E, limestone, on mossy rocks. Collected by J. Stanisic, D. Powter, P,H, Col- man. 3 Mar 1987. Height of shell 1,1 3mm, diameter 2.77. H/D ratio 0.41, D/U ratio 2.37, whorls 5 1/&+ PARATYPES, QMMO17015, 106 specimens, same collection data as holotype; Limestone caves wt Natural Arch. W. of Kempsey, NSW (30°59°S, b52"20'E) (150+, AMC153720, 10 Mar 1981. W. Ponder, O, Griffiths); Haydonville, W. of Kempsey. §. side of ‘Natural Arch’, under rock over- hang (4 specimens, AMC152156, 20 Jan 1982, ex A.B. Rose, M. Dodkin); Yessabah Cave, W. of Kempsey, in leaf and soil litter (31°06'S, 152°42°E) (50+ specimens. AMC119351,9 Oct 1979, J, Stanisic, PH, Colman), Natural Arch, Carrai Caves, litter (70+ specimens, AMC 152155, 22 Feb 1980, P.-E. Calman). DIAGNOSIS Shell minute, diameter 2.47-2.98 mm (mean 2.66 mm) with 4 3/4- ta 5 1/2- (mean 5) hghtly voiled whorls. Spire and apex depressed, last whorl descending more rapidly, Height 1,03— 1.17 mm (mean 1.07 mm), H/D ratio 0,380.43 125 mm (mean 0.40 mm). Protoconch of 1 1/2 to | 5/8 whorls, exsert, mean diameter 449.8 jm at I 1/2 whorls. Apical sculpture (Fig. 77d) of curved radial riblets, more crowded at nuclear- post nuclear boundary, and fine spiral wrinkles. Post-nuclear sculpture (Fig. 77e) of numerous. crowded, protractively sinuated radial ribs, 23(+ 330 (mean 297.7) ribs on the body whorl. Ribsimm 28.63-38.19 (mean 33,34), Micros- culpture (Fig, 77e) of crenulate radial riblets, 2-3 between cach pair of major ribs and very low, widely spaced, rather inconspicuous spiral cords, raised at their junction with the microradivls to produce a weak beaded effect, continuous on the apices of the radial ribs. Um- bilicus (Pig. 77b) wide, cup-shaped, diameter 1,03-1.29 mm (mean 1.14 mm). D/U ratio 2.2.1- 2.44 mm (mean 2.33 mm), Sutures impressed. Whorls rounded ubove and below the periphery (Fig. 77e), shouldered at the upper margin, Mat- tened below, Aperture ovately lunate with seven barriers. Four palatal barriers (Pig. 78a), fourth and third large crescent-shaped; second smaller, crescent-shaped at the basal margin; first similar in size to second, Situated at the baso-columellar margin. Three parietal barriers (Fig, 78b). First and second clongate crescent-shaped lamellac: third a low elongate blade. Lip simple, columells not thickened, almost vertical, not reflected. Parietal callus present. Colour light straw-yel- Jow with broad reddish-brown radial streaks. and suffusions, Based on |! measured adults. Genitalia with vas. deferens/epiphallus junc tion valvular. Epiphallus (Fig. 79a) long, thick muscular, coiled: internally with longitudinal (thickenings, entering penis apically through thickened pilasters (Fig, 79b). Penis (Fig, 79b) a long, muscular tube with a thin sheath, inter- nally with a primary longitudinal pilaster, a medial, spherical thickening and secondary, smooth longitudinal thickenings. Penial retrac- tor muscle short, originating from the diaphragm and inserting on the epiphallus just prior ta its entry into the penis. Spermatophore (Fig. 79f) hook-shaped. Radula (Fig. 78e, f) with very long lanceolate mesacone on laterals, Based on three dissected adults (QMMOQ- 17015). RANGE AND HABITAT Limestone outcrops between Yessabah ard Carrai, west of Kempsey, NSW in evergreen vine thicket derived [rom the sutrounding warm temperate rainforests. Live specimens have 126 MEMOIRS OF THE QUEENSLAND MUSEUM a b c Piven FIG. 77. Rhophodon kempseyensis sp. nov. a—c, Natural Arch, Carrai S.F., NSW. QMMO17016, holotype; d-f, same data holotype. QMMO17015, AMC153720, paratypes. a—c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, base. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 127 FIG. 78. Rhophodon kempseyensis sp. nov. Natural Arch, Carrai S.F.,. NSW. QMMO17015, paratype. a, palatal barriers; b, parietal barriers; c, microdenticulation on surface of third palatal barrier; d, radula; e, central and lateral teeth; f, marginal teeth. Scale lines as marked. 128 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 79. Rhophodon kempseyensis sp. nov. Natural Arch, Carrai S.F., NSW. QMMO17015, paratype. a, genitalia; b, penis interior; c, hermaphroditic duct; d, ovotestis: e, talon and carrefour; f, spermatophore; g, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE Bronpouy > 500m fore Macquarie FIG, 80, Distribution of Rhophodon kenipseyensis sp nov, been collected from the surface of mossy lime- stolic rocks. REMARKS The lateral teeth in which the mesocone is conspicuously large, are similar to those in the sympatric Letomola contortus and Coeno- charopa yessabahensis which also have been collected live on the limestone rocks, This con- vergence may reflect similar feeding habits. R. kempseyensis is closely related to R. peregrinus and R. cousobrinus probably derived by habitat fragmentation. Rhophodon elizabethae sp, nov (Figs 81-83: Tables 15, 16) ETYMOLOGY Por Elizabeth Chaseling. COMPARISONS Small size, flammulated shell with bold radial ribs. wide umbilicus and number of apertural barriers distinguish (his species from any sym- patric charopid species. R. minudissinues 1s most similar to R. elizabethae but has fewer whorls, smaller umbilicus, and a smaller monochrome golden brown shell. In addition R, elrzabethae (Fig, 81h) has a greater number of palatal bar- riers and, unlike R. minutissimus (Pig. 34g.h), does not have the second parietal barrier shor- (ened, ta <= “TYPE MATERIAL HoLotyee: QMMO17315, Granite Creek, fvothills of Bobby Range, SEQ. 24°30.7°S, 151°30.3'E, under leaf liter, Collected by T. Carless, 7 Dec 1983. Height of shell }.1 1 mm, diameter 2.73 mm, H/D ratio 0.41. D/U ratio 2.76. whorls 5 1/44. PARATYPES: QMMO14714, | specimen, same dala as hololype; summit Mt Bonroon Booroon, SW of Miriam Vale, SEQ, MVF/Arauearia, litter, on and among rocks (17, QMMO16590. QMMO16858, 17 Sept 1985, J. Stanisic. D. Potter); base, Mt Booroon Booroon, SEQ, NVF along creek, under logs (3. OMMO16592, 17 Sepi 1985, J. Stanisic, D. Potter). DIAGNOSIS Shell minute, diameter 2.26-2.73 mm (mean 2.47 mm). with 4 3/8 to 5 1/4+ (mean 5-) tightly coiled whorls, last descending slightly. Apex and early spire (Fig. 81a) flat to very slightly elevated, SP/BWW ratio 0.05—0. 19 (mean 0.12), height 0,94-1.24 mm (mean 1.10 mm). H/D tatio 0.41-0.48 (mean 0.45), Protoconch of 1 5/8 to 1 3/4 whorls, mean diameter 520.1 um at 1 1/2 whorls. Apical sculpture (Fig. 8le) of curved radial ribs, becoming more crowded at the nuclear-post nuclear junction. Post nuclear sculpture (Fig. 81f) of bold. widely spaced, protractively sinuated radial ribs, 52-82 (mean 65.2) ribs on the body whorl. Ribs/mm 6.93-— 10,21 (mean 8.38). Microsculpture (Fig. 81f) of fine. thread-like radial riblets, 10-18 between each pair of major ribs, and fine crowded, spiral cords Which are continuous on the apices of the radial tibs. Umbilicus (Fig. §1b) wide, cup- shaped, diameter 0,92-1,13 mm (mean 1.01 mm). D/U ratio 2.07-2.76 (mean 2.46). Sutures impressed, whorls rounded below and strongly shouldered above a slightly flattened periphery (Fig. Sic), Aperture ovately lunate with three parietal, five palatal and one columellar barrier, Parietal barricrs blade-like, consisting of two lamellae and an elongate, thread-like superior trace, First barrier clongate. crescent-shaped, abruptly tapered at cach end; second with anterior taper and an expanded. downwardly deflected, posterior section. Columellar barrier low. crescent-shaped, horizontal. Five palatal barriers (Fig. 1h) consisting of four lamellac and one trace. Lamellae short. expanded. Supe- rior [race sometimes present, short, close to the palato-parietal margin. Lip simple: columella without noticeable thickening. Parietal callus developed, Colour light straw-yellow with vari- able brown radial streaks and suffusions. Based on 9 measured adults. MEMOIRS OF THE QUEENSLAND MUSEUM imm SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 131 H “Hy MG “Mc FIG. 82. Rhophodon elizabethae sp. nov. Summit, Mt Booroon Booroon, SW of Miriam Vale. SEQ. QMMO16590, paratype. a, genitalia; b, penis interior; c, hermaphroditic duct: d, ovotestis; e, pallial cavity. Scale lines as marked. FIG. 81. Rhophodon elizabethae sp. nov. a—c, Granite Creek, foothills of Bobby Range, SEQ. QMMO17315, holotype; d-h, Summit, Mt Booroon, Booroon, SW of Miriam Vale, SEQ. QMMO16590, QMMO16858, paratypes. a-—c, entire shell; d, microdenticulations on parietal barrier; e, apical sculpture; f, post nuclear sculpture; g, radula; h, palatal barriers. Scale lines as marked. 132 MEMOIRS OF THE QUEENSLAND MUSEUM Genitalia with epiphallus (Fig. 82a) reflexed before entering the penis apically through a simple pore (Fig. 82b) surrounded by a fleshy pilaster. Penis (Fig. 82b) simple with apical bulb containing epiphallic entrance and lower cham- ber containing a single large. spongy, lon- gitudinal pilaster, several shorter pilasters, and low pustules. No penial sheath. Penial retractor muscle a short tuft inserting apically on penial bulb adjacent to epiphallus, Radula (Fig, 81g) typical. Based on 2 dissected adults (QMMO16590), RANGE AND HABITAT Under logs. on rocks and amongst litter in moist notophyll vine forest of the Bobby Range, SEQ. REMARKS R. elizabethae has a very restricted range com- pared to other charopids. Its closest relative appears to be R. minutissimus. Rhophodon minutissimus sp, noy. (Figs 84-86: Tables 15, 17) ETYMOLOGY Latin minutus, little: for the small size. TYPE MATERIAL HoLoTyreE: QMMO17316, c. 1 km §$ of Raynbird Creek Rd. on Lacey's Creek Rd, Lacey's Creek, SEQ 27°14°S, 152°43°E, Collected AM/QM-ABRS, 30 Sept 1982. Height of shell 0.70 mm, diameter 1.29 mm, H/D ratio 0.54, D/U ratio 2.10, whorls 4 5/8. PARATYPES: OMMQO12094, AMC136479, 29 specimens, same locality data as. holotype; Bunya Mts N.P,, SEQ, NVF (18 QMMO6089, 3 Mar 1976, M,J. Bishop); Dandabah, Bunya Mts N.P., SEQ, behind picnic area, (26°50°S, 151°37'E) (5, OMMO12280, AMC 136604, 31 Aug 1982, AM/QOM-ABRS); sidetrack off Mt Archer Rd, Mt Mee S,F., SEQ (2709'S, 1S2°41°E) NVF/Araucaria (3, QMMO- 15078, 6 Sept 1983), J. Stanisic, D. Potter): Wratten’s Camp, Wratten’s S.F., SEQ (26°17'S, 152°20°E) NVF, litler(3, OMMO11554, 17 Jul 1980, J. Stanisic, A. Green); Little Yabba Creek, Imbil S.F., SEQ (26°28°S. 152°38°E), NVF (1, QMMOQO12023, 8 Sept 1982, AM/QM-ABRS): Fred’s Rd, Mt Mee, SEQ (27°05°S. 152°43°E) raintorest, leaf litter (1, QMMO- 17318, 14 Apr 1980, J. Stanisic, N. Hall, A. Green); Big Falls Circuit, Bunya Mts N,P., SEQ, SNVF. under logs (1, OMMO16852, 5 Nev 1985, J. Stanisic, D. Potter): Dandabah, Bunya Mis, N.P., SEQ (26°50°S, oS ” B & 24530's AX SS AS a Sy N\ ZAP35'S PANS a - Mt, Beatrice “Qs Ralt. > 400m Zhe4gis ANia “AR NAT. Bolangul 151°30'E 1S7°35'E FIG. 83. Distribution of Rhkaphodon elizabethae sp. noy. in the Bobby Range area, SEQ, 151°35°E), NVF/Araycaria (16, QMMO6069, 5 Mar 1976, M.J. Bishop). DIAGNOSIS. Shell minute, diameter 1,29-1,66 mm (mean 1.52 mm) with 4 3/8 to 4 5/8 (mean 4 1/2+) tightly coiled whorls. Last whorl descending more rapidly. Early spire and apex (Fig. 84a) flat to very slightly elevated, SP/BWW ratio 0.07— 0.12 mm (mean 0.09 mm), height 0.64-0.76 mm (mean 0.70 mm), HD ratio 0.40—0.54 mm (mean 0.46 mm). Protoconch of 1 5/8 to 1 3/4 whorls, exsert, mean diameler 391.9 um at 1 1/2 whorls. Apical sculpture (Fig. 84e) of curved radial ribs and spiral wrinkles. Post nuclear sculpture (Fig. S4f) of numerous, prominent, protractively sinuated radial ribs, 56-98 (mean 76.6 mm) ribs onthe body whorl. Ribs/mm 10.98—21.26 mm (mean 16.23 mm). Microsculpture (Fig. 84f) of fine radial riblets, 8-12 between each pair of major ribs, and very low, barely visible, spiral cords which are raised at their junction with the microradials to produce a beaded effect, Um- bilicus (Fig. 84b) wide, cup-shaped, diameter 0.60-0.72 mm (mean 0.65 mm). D/U ratio 2.09— 2.59 mm (mean 2.35 mm). Sutures impressed, whorls rounded below and shouldered above a slightly, laterally compressed periphery (Fig. 84c). Aperture ovately lunate. Barriers present, consisting of three palatal and three parietal 133 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE (ouz-#E7) PRUUFISZ (p7'2-207) PULTE (ro zs 7) AL OFLS ree (LS7-62'2) EROUF IPT ere Eta (avz-tr'2) LEWUFEST sez (gp'Z-0E2) OM UFOLT (65°2-07'2) SSO UF PRT ia irara (cb'2-O'z) SUPOFCH TS Ore OUVY VO (260-820) OU OFSR (cr1-z60) ce OF10'L 80. 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Rhophodon minutissimus sp. nov. Big Falls Circuit, Bunya Mts N.P., SEQ. QMMO16852, paratype. a, genitalia; b, penis interior; c, ovotestis; d, hermaphroditic duct; e, pallial cavity. Scale lines as marked. FIG. 84. Rhophodon minutissimus sp. nov. a—h, c. 1km S of Raynbird Creek Rd, on Lacey's Creek Rd, Lacey’s Ck, SEQ. QMMO12094, paratype. a—c, entire shell; d, microdenticulations on surface of upper parietal; e, apical sculpture; f, post nuclear sculpture: g, parietal barriers: h, palatal barriers. Scale lines as marked. 26°S 27°S 28°S FIG. 86. Distribution of Rhophodon minutissimus sp. nov. lamellae (Fig. 84g). First and third parietal lamellae large with anterior edge protruding from aperture, expanding posteriorly: second deeply recessed, short, crescent-shaped. All directed downwards. Palatal lamellac (Fig. 84h) deeply recessed, short, crescentic, Lip simple without columella expansion or reflection. Parietal callus developed. Colour golden- brown. Based on 12 measured adults. Genitalia with epiphallus (Fig. 83a) thick. coiled, muscular; longer than the penis, internal- ly with longitudinal pilasters, entering the penis apically through two fleshy thickenings (Fig. 85b). Penial retractor muscle inserted at the penis/epiphallus junction. Penis a broad mus- cular tube with sheath; internally with lon- gitudinal fleshy pilasters (Fig. 85b). Radula typical, Based on 2 dissected adults (QMMO16852). RANGE AND HABITAT Notophyll vine forests of the D’Aguilar Range, Bunya Mountains, and the Coast Range MEMOIRS OF THE QUEENSLAND MUSEUM near Gympie, SEQ. R. minutissimus has been collected in litter and from under logs. REMARKS Ahility to exist in drier open forest is probably a contributing factor to the comparatively wide distribution of this species. R. bairnsdalensis displays similar adaptability and breadth of dis- tribution. Rhophodon colmani sp. nov. (Figs 87-88; Tables 15, 17) ETYMOLOGY For Mr Phillip Colman, Australian Museum, COMPARISONS Small size, almost fat spire, widely spaced ribs, open umbilicus and apertural bartiers, ef- feclively separate R, colmani from any sym- patric charopid. Egilomen cochlidium from the McPherson Ranges ts grossly similar in shell form, but lacks apertural barriers, and has a Jarger shell with higher whorl count, higher rib count and more elevated spire. Conchologically R. colmani is most similar to R. bairnsdalensts from southern NSW and Vie- toria. However the latter species is larger, with more ribs, wider umbilicus, greater number of whorls and quite different pattern of apertural barriers. The three parietal, six palatal and single columellar barrier of R. colmani (Fig. 87f-h) contrast with the single parietal and two palatals of R. bairnsdalensis (Fig, 90a-c). TYPE MATERIAL HoLotyre: QMMO17314, Kenilworth State Forest, SEQ (26°35°S. 152°36°E) mixed notophyll vine forest. Collected by M.J. Bishop, 18 May 1976, Height of shell 0.64 mm.. diameter 1.56 mm, H/D ratio 0.41. D/U ratio 2.38, whorls 4 5/8, PARATYPES: QMMO6288, 14 specimens, same col- jection data as holotype, DIAGNOSIS Shell minute. diameter 1.34-1.56 mm (mean 1.45 mm) with4 1/4 to 4.5/8 (mean 4 1/4-) tightly coiled whorls, the last descending more rapidly. Early spire and apex (Fig, 87a) flat to very slight- ly elevated, SP/BWW ratio 0.09-0.14 mm (mean 0.11 mm), height 0.53-0.66 mm (mean FIG. 87. Rhophadon colmani sp. nov. Kenilworth S.F..SEQ. QMMO6288, paratypes, a-c, entire shell: d, apical sculpture; e, post nuclear sculpture; f, parietal barriers, 2. palatal barriers: h. columellar barrier, Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 137 138 FIG. 88. Distribution of Rhophodon colmani sp. nov. 0.61 mm). H/D ratio 0.38—0.46 mm (mean 0.42 mm). Protoconch of 1 1/2 to 1 5/8 whorls, exsert, mean diameter 357.4 wm at 1 1/2 whorls. Apical sculpture (Fig. 87d) of curved radial ribs, becom- ing more numerous at the nuclear-postnuclear junction, and fine almost obsolete spiral threads and wrinkles. Postnuclear sculpture (Fig. 87e) of bold, widely spaced, protractively sinuated radial ribs, 23-29 (mean 26.4) ribs on the body whorl. Ribs/mm 5.48-6.51 (mean 5.81). Microsculpture (Fig. 87e) of fine radial riblets, 17-25 between each air of major ribs, and low spiral cords. Umbilicus (Fig. 87b) wide, cup- shaped, diameter 0.51-0.66 mm (mean 0.58 mm). D/U ratio 2.38—2.60 mm (mean 2.50 mm). Sutures impressed. Whorls rounded below and strongly shouldered above a compressed periphery (Fig. 87e). Aperture ovately lunate. Barriers numerous consisting of three large parietal barriers (Fig. 87f) protruding from the aperture, a large columellar barrier (Fig. 87h) situated inside the aperture margin, and six pala- tal barriers (Fig. 87g) set back about 1/8 whorl. First palatal barrier small, crescent-shaped, tapering rapidly anteriorly; second similar in shape, but with longer taper anteriorly; third and fourth about twice the size of the second with long antcrior tapers; fifth almost equal in size to the second, rapidly tapering anteriorly; sixth MEMOIRS OF THE QUEENSLAND MUSEUM elongate, thread-like, about twice the length of the others. Columellar barrier (Fig. 87h) a large crescent-shaped lamella situated well inside the aperture. First parietal barrier long expanded posteriorly and directed upward; second and third, thin anteriorly, expanding posteriorly, swollen. Lip simple, without columella expan- sion or deflection. Parietal callus well developed. Colour light straw-yellow. Based on 5 measured adults. Anatomy unknown. RANGE AND HABITAT Notophyll vine forest of Kenilworth State Forest, SEQ. No data are available on micro- habitat. REMARKS Lack of soft parts and poor distributional data restrict discussion. R. colmani is most likely sympatric with R. minutissimus and it is tempting to speculate that the sharply altered shell pat- terns, in particular the increased number of pala- tal barriers, represent character displacement under conditions of sympatry. Rhophodon bairnsdalensis (Gabriel, 1930) comb. nov. (Figs 89-92; Tables 15, 17) Charopa bairnsdalensis Gabriel, 1930, p. 78, pl. 2, figs 11-12. Endodonta bairnsdalensis (Gabriel); Kershaw, 1956a, p. 137 Egilodonta bairnsdalensis (Gabriel); Iredale, 1937a, p. 329. PREVIOUS STUDIES Iredale (1937) included this species in a new genus Egilodonta based on the presence of aper- tural barriers. He did not give reasons for separating the species from the northern Rhophodon nor from the geographically more proximate Dentherona Iredale, 1933 and Bis- choffena Iredale, 1937. While R. bairnsdalensis has fewer apertural lamellae than other species of Rhophodon, shell sculpture is comparable, as is the structure of the pallial cavity and terminal male genitalia. A review of Dentherona and Bischoffena is beyond the scope of this study, but coiling patterns and microsculptural details of these genera suggest that they are not related. COMPARISONS R. bairnsdalensis can be separated from its congeners by its average size, widely spaced SYSTEMATICS OF AUSTRALIAN CHAROPIDAE at a 500m 30°S 150°E AS1°E 752°8 153°E 154°E FIG. 94. Distribution of Discocharopa aperta (Mollendorff, 1588) in southeast Queensland. Tyre MATERIAL LecToTyPe (designated Solem, 1983, p. 77); AMC- 8970, Bundaberg, Qld. ex May. Height 0.61 mm, djameter 148 mm, H/D ratio 0.45, D/U ratio 2.81, whorls 31/2, PARALECTOTYPES: AMC63489, 7 specimens, same data as lectolype. OTHER MATERIAL c. | km frum Qlsen’s Caves, MEQ (23°10°S, 150°30°S) litter, limestone outcrop (16, QMMO- 17307. 13 Jul 1980, JStanisic, A, Green, G. An- mabell. Mt Archer, Bersaker Ra,.MEQ (23°21°S, 150°35°E), remnant vine thicket (6, OMMO11922. 30 Jun 1982, J. Stanisic); c. 2 km W of Gayndah, on Gayndah-Hinesville Rd, SEQ (25°38°S, 151°36°E) bottle tree scrub/SEVT, under rocks and in litter (2, QMM011528,16 Jul 1980, J. Stanisic, A. Green); Dan Dan Scrub, SEQ, SEVT, in hitter (2, QMMO16800, 10 May 1984, J. Stanisic, D. Potter); Dawes Range, SEO (24°28'S, 151°07'E) MVF/Araucaria (28, QMMO12666, AMC136795, 4 Sept 1982, AM/QM - ABRS); c. 1,8km E of Builyan Rwy Stn, on Builyan- Gladstone Rd. SEQ (24°22°S, 151°24°E), rainforest an river bank litter (13. QMMO17308, 15 Jul 1980, J. Stanisic, A. Green); Mt Biggenden, SEQ (25°32'S, 151°S0°E), SEVT. limestone outcrop (5. QM- MOQ17309, § Jul 1980, J. Stanisic, A. Green; 1, OM- MO16834, 3 Jul 1984, J. Stanisic, D. Potter, K. Emberton; 14, QMMO11958, AMC136396, 3 Sept 1982, AM/QM - ABRS), Walla Range, E of Ban Ban Springs, SEO (25°38'S, 151°50°E) SEVT (1, OMMO- 12414, 3 Sept 1982, AM/OM - ABRS): Kalpowar S.F., SEQ MVF/Arauearia (2, OMMO17310, 15 Jul 1980. J, Stanisic. A.Green; 2, QMMO17306, 4 Sept 1982, AM/OM - ABRS): Gavial Ck, Bouldercombe, MEQ (23°34°S, 150°28°E) MVF, litter (4, QMMO- 16808, 9 Jul 1983, J, Stanisic, D. Potter); Coalstoun Lakes Rd. Coalstoun Lakes. SEQ (25°37°S, 151°53'E) SEVT (1. OMMOQ12396, 2 Sept 1982, AM/OM - ABRS); c. 54 km § of Biloela, an Biloela-Monto Rd, SEQ (24°445'S, 150°S0°E) SEVT on racky outcrop, litter (3, QMMO17311, 14 Jul 1980, J. Stanisic, A, Green); Mc Etna, MEQ (23°09°S, 150°27°E) MVF on limestone outcrop, litter (100, OMMO17312, 13 Jul 1980, J, Stanisic, A. Green, G. Annabell); Johannsen’s Caves, MEQ (23°09°S, 150°27°E) MVF on limestone outcrop, litter (19, QMMO17313, 13 Jul 1980, J. Stanisic, A. Green, G. Annabell), Cammoo Caves, ¢. 24 km N af Rockhampton, MEQ (23°10°S, 150°27°E) limestone outcrop (4. AMC152145. W. Ponder, P.H. Colman, J.B. Burch, 8 Sept, 1970); Glastonbury SF, c, 0.25 km § of forestry station (26°13°30"S, 152° 29° 45°E) (1, QMMO13305, 7 Sept 19§2.AM/OM- ABRS), S of Gympie, entrance to state forest olf Yabba Ck Road (26°28°S, 152°38°E) (3, AMC. 138110 QMMOQ13365, 7 Sept 1982, AM/QM- ABRS); Kenilworth SF. SEQ (26°37°S, 152°42'E), rainforest. leaf litter (6, QMMOS360, 22 May 1980, J. Stanisic, A-Green): sidetrack off Mt Archer Rd, Mt Mee SF, SEQ (27°04 'S, 152°41°E) (1, QMMO12173, 31 Aug 1982, AM/OQM-ABRS): Natural Bridge NP. SEQ (28°13°S, 153°14E) NVF (4, QMMO10455- AMC129266, 18 Mar 1981. AM/QM-ABRS); Fred’s Rd, Mi Mee, SEQ (27°05°S, 152°43°E) rainforest, litter (3, OMMO8357, 14 Apr 1980, J. Stanisic. N Hall, A. Green); Yarraman Forest Drive, Yarraman SF, SEQ (2657S, 151°57E) MVF (1, QMMO- 12261, AMC136586, 31 Aug 1982, AM/QM-ABRS MEMOIRS OF THE QUEENSLAND MUSEUM 146 (p6'2-65°2) OL ORLT OR'Z 097% {1Lz-90'2) CRUST 4 4 RST (ge'z-97'2) 6rururvEe'z Lez ove (te 7-42) £76'0798'2 {oLz-12'2) Sz 079P'Z (1-12) TOILE wsz trz OLLVY N/a (ov'0-seu) SHOFREO ico crt) (e040) CLV OFSED zs ero oru (Ls'0-RF'0) LZO'UFTS'O zs0 sO (1s'0-zP'0) 9Z0'0FLY'O (co0-+r'0) SHOOFHP'O (is'0-9¢'0) scororer'o aro ovo (ww) HLGIM TWOrldnwo (169¢-H9 PE) LOY OFES'SE sele cree (cee¢-60'R2) ISO EVE TE FLY sper so'se (pp'ge-R1'92) ECS PF POE PH SE (09 pe-Se'ze) SILWRLL'CE (L8'St-4L'82) LSCFIETE (ZS E67) $65 IFL6°EC SS FE SOOT WASTE (601-6) (oro-arn) Clea-ne'o) (e0-170) (Lp'0-£F'0) SRZFEYS OL OF O00 COUT TED LEUOFK'O £LO'OF9F'O ub 00 0 zo sro £6 oro oc0 £00 €r'0 (62-9) Grou) (ago-97'0) (so’o-z0'0) (zs'0-zP'0) IZ EHEL seoureed ZI OFRZO 160'OFP0'0 ZEO'OF9E'O aa) wo reo zoo Prd ol rr a 00 rr 6 Pro 670 100 sro (o11-za) (ara-aro) (ce'0-ze-0) (zp'o-se'0) SH'SF5"OH CO0FOFO LONUFEEO cou LIOOr6e0 ot or ee0 soo “ro - ir) FeO £00 1r'0 (z11-co) (EV0-90'0) (reo-ze'0) (00-200) (6v'0-1¥'0) Ch IFC OUT £Z20°0F 800 LOOUFEE'O LOU0F E00 £Z20'0FS¢'0 (101-06) (9L0-z1"0) (ge0-eL'0) (90'0-F0'0) (s5"0-zP'0) EV7FS'S6 ozo OFF 10 sz0'nr9e'0) O1G0F50°0 S90'0F6P'0 (R01-£6) (zz'0-L00) (ceo-az'0) (L0'0-Z0'0) (9p'0-se'0) 9S'TFSO0L scotzid HOOROEO z10'0=F00 s7arorzro ou zo c'0 r0'0 Pro zn ero oro cro sro (us) (wut) Sard OLLVY HLGIA NOISNYLONd OLLVa GH MANES THOHM Adon auld (QUNV¥ CNV WaS NVA) (8881 ‘SSHOONTTION) WLYIdV VdONVHIOISIC Ni NOLLVIEVA TWIT - (soI-10'1) Z1OOF EOL LOL 601 (z6'0-z8'0) ££0°0768'0) scvl sot cot (6c I-s1'1) Leoarze't ira} {ezI-sv) #20'0702'1 (6V'I-L1'1) OLOOFRIL (ez"1-£01) Pro TL ral] (us) waLaWvid S1ATaVL (6r'0-rr'0) SLOOFLPO ero LO. (cro-0e'0) ZIVOF LPO ssa oF’ sro (ora-+e'0) LOO Leo Ro'0 wa (95°0-15°0) Siores'O (e90-05°0) OLOUFLS'O (is 0-70) zenoF Reo 0 soo (uu) LHOISH (+8/6 E-B/1 ©) +H/L B/TE tT E (p/t €-8/1 £) +B/L£ B/S € we Le (p/e €-8/5 £) +B/S P/E E v/EE (a/s cor-R/¢ €) RISE (8/€ £-B/1 £) P/E (r/c evi £) REE +8/5 £ RISE STHOHM £ 89801 ONWO AaumopyrEy, t tstol ONWO adpug jenjen T ELIZL ONWO 33 WW £ 09€8 ONWO. quompuay t s9tfl ONWO 2D eqgeA ANT t 1977LOWWO uewrene A, T SOfElL ONWO aiduikp * LOELT ONWO SOAEQ S,U3S1O t 22611 ONWO soqouy IW I 00891 ONWO russ ueg eg € 99971 ONWO. odury sameq z 90£L1 ONWO “4's semodjey ’ R0£L1 ONWO ue fying I 60£41 ONWO uapuaddig 1 (mansuex ad4Qo277) ! 0L68 ONWO Suaqepung pysado vdosvyr0281¢) SNAWIDadS 40 ¥agWN HAVN SYSTEMATICS OF AUSTRALIAN CHAROPIDAE c. 15 km from Rathdowney on Boonah-Rathdowney Rd, SEQ, rocky outerop, vine thicker (2811'S, 152°46°E) (8. OMMO1086R, AMC128445, Mar 1981, AM/QM-ABRS); Ginna Burra, Lamington N.P., SEQ (28°12'S, 193°11°E) NVF/Palms (1, QM- MO 10444, 18 Mar 1981, AM/QM-ABRS); Koreelah Ck., Beaury S.F. c. 15 km W of Urbenville, NSW 24°217S, 152°20"E) 530 m, litter (2, AMC152151, 18 May 1976, PH, Colman, 1, Loch); ¢ 3 km from junction with Lever’s Platewu Rd, Tartar’s Creek, SEQ (28°28"S, 152°08"E), roadside gully (1, QMMO- }1 148.17 Mar 1981. AM/QM-ABRS): DIAGNOSIS Shell minute. diameter 0,82-1.29 mm (mean 1.19 mm) with 3 1/8 to 3 3/4 (mean 3 I/2-) loosely coiled whorls, last whorl descending in front. Early spire and apex (Fig. 93c) barely elevated, SP/BWW ratio 0.06-0,22 (mean 0.11), height of shell 0.39-0,64 mm (mean 0.49), H/D ratio 0.35-0.55 (mean 0.44). Protoconch of | 3/8 ta 1 3/8 whorls, mean diameter 693.9 jm at | 1/2. whorls. Apical sculpture (Fig, 93e.f) of widely spaced, slightly sinuous radial ribs and even finer. irregular spiral riblets. Postnuclear sculpture of crowded, protractively sinuated radial ribs, 67-134 (mean 96.70) ribs on the body whorl. Ribs/mm 24.25-40.74 (mean 31.75). Microsculpture (Fig. 93e,f) of fine radial riblets, Whose apices are crenulate. 4-10 between each pair of major ribs.. Umbilicus (Fig. 93b) saucer- shaped. diameter 0.34-0,57 mm (mean 0.44 mm). D/U ratio 2.16-3.31 (mean 2.41). Whorls rounded above and below the periphery. Aper- ture ovately lunate. Lip simple. Parietal callus developed. Colour light straw-ycllow to white. Bused on 28 measured adults, No soft parts available for dissectian, RANGE. AND HABITAT In eastern Australia from just north of Rock- hampton, south ta near the Queensland/New South Wales border. In the northern purt of its range i.e. between Biggenden and Olsen’s Caves. the species lives in dry subtropical microphyll vine forest. In the south i.e. between Gympie and Natural Bridge National Park. D. aperta inhabits warmer moist notophyll vine forests. 2. aperta lives among the ditt particles below the leaf litter. RUMARES The decision to consider all the above specimens as D. aperta is made reservedly. There are conchological differences between the northern and southern populations which indi- cate two species. However, without soft parts, recognition of two species is a tenuous proposi- lion (Solem, 1983). D. aperta is probably a complex of species which represent localised experiments in mimaturisation, Cralopa Iredale, 1941 Cralopa tredale. 94) @ p. 267: Kershaw, 1956b, p.h: Batch. 1976, p. 132. TYPE SPECIES Helix stroudersis Cox 1864 sby original desig- nation. PREVIOUS STUDIES Tredale (194 1a) introduced Cralopa for Helix stroudensis Cox, 1864 which has a small shell with weakly clevated spire, closed umbilicus, and numerous curved, bold radial ribs. ‘Cralopa’ intensa Iredale. 1941 Was added on the busts of small size und crowded radial postnuclear sculpture. Two new species are added here and ‘C. wrrensa is removed to Sinployea Solem, 1983, DIAGNOSIS Shell small, diameter 1.66-3.83 mm, with 3 3/4 to 4 1/2 tightly, to loosely coiled whorls, the last descending slowly. Farly spire and apex slightly elevated. Apical sculpture of prominent to bold, weakly curved radial ribs and low, nar- Tow spiral cords. Adult sculpture of few (carles~ st) lo many (stroudensis), widely spaced (carlessi). 10 crowded (keputarensis), bold, strongly protractively sinuated radial ribs, Microsculplure of fine radial riblets and crowded, narrow spiral cards, continuaus-on the major tibs. Umbilicus closed or only narrowly open, Sutures deeply unpressed. Whorls round to slightly shouldered ubove the periphery and rounded below. Aperture roundly lunate. Lip sinuous, strongly retracted at the sutural margin to form an apertura! sinus. Columella reflected over the umbilicus. Foot short, bluntly rounded posteriorly, colour white, ommatophores white. Mantle collar with an orange glandular section extending onto the pallial roof. Kidney weakly bilobed, with pericardial lobe longer, Ureter sigmurethrous, complete. with primary arm refleyed near origin. Terminal male genitalia with a relatively ex- panded vas deferens, very large evlindrical penis, and weakly differentiated epiphallus. In- 148 PAULEY RANCHI OP VARIATION IN CRALOMA AND ELSOHTIENA (MEAN AND RANGED TiMHIE ICAL Wwiprl SP/UWW RATIO BODY WHOL S#itbe PROTRUSION NUMBER OF SPRCIMENS, NAMIE B/U RATIO VU MM HUES WIDTH H/D RATIO HEIGHT DIAMETLR (mm) WHORLS dam) {nvm (mm) (mm) HAS be at 9 in (ha-0.26) 2AY (hit Wb Ww Cralapu (7.M6-HS2) (S247) (UP AY) {ey (Wh 201.4) (2.28- 1.43) (h45-2,22) (A 7K-4 1/40) Strdmdensis ws {het (52454) mv wad (1.66-1.75) ooo (4 HA Wis) Craleypaa Aapurareines SMR (kta 5h OLLTATY4) (0.77-1.15) 21-0.A4) {te 056 (O4R-DA DD 330 (298-474) 4a (ESR 17) aa au. (4,747.58) (aes ey MEMOIRS OF THE QUEENSLAND MUSEUM 1).2t (9871 U2) (045-14u) M2 (ha) 1,63 (1451.79) 22 (kT 0.49) (3.6 5.54) (thay 5) 8) A Side) sericaiwla Elgothera (7.48 17H YW (2) ay 144 CH2-1 58) (124-100) 0.u4 tay (0.02012) Vall (hAS=1N) iid (0), (OSOEUT) 54 443 (4.55.24) Sh a4 (2.17-2.41) 4B (4 Vited Wde) nautiliten Liserhera 724 (SRI A) wy (hte 227) WS dia Myarkera (47 62) (Uit-155) (497-664) (277-4) (44 5ky aenithecan ternally. penis with apical epiphallic pore and two large, spongy longitudinal pilasters. Penial retractor muscle inserted on penis (stroudensis, carlessi), or epiphallus near penis (kaputaren- sis). Vagina relatively short. Radula with tricuspid central and lateral teeth; central tooth reduced, about 1/2 size of the laterals, cusps lanceolate, mesocone long; lateral teeth with large lanceolate mesocone. DISTRIBUTION AND ECOLOGY The east-west distribution of Cralopa con- trasts with the more common north-south pattern shown by other genera reviewed in this study. Although the extensive rainforest distribution of C, stroudensis appears to contrast with the ‘cornered’ distributions of C. kaputarensis and C. carlessi, closer analysis reveals that C. stroudensis is also an environmentally isolated species in cool subtropical notophyll vine forests of the Great Dividing Range. PATTERNS OF VARIATION Conchological variation is conservative and probably reflects relatively recent fragmentation of populations. It involves changes in size, spire protrusion, whorl count, rib spacing and rib num- ber. C. kaputarensis, the smallest member of the genus, is a scaled down version of C. straudensis with less protruded spire (SP/BWW ratios = 0,54 and 0.99 respectively) and more crowded ribs on the body whorl. C. carlessi is larger with greater whorl count and more elevated spire. C. stroudensis has the greatest number of ribs but C. kaputarensis has most ribs/mm. C. carlessi has the widest spaced ribs. An interesting aspect of the Cralopa shell is the development of an apertural sinus. This is more commonly seen in New Zealand Charopidae. COMPARISONS Cralopa is most similar to Elsothera having shells with very similar sculpture, barely open to closed umbilicus, and relatively depressed spire. However, Elsothera is much larger with greater mean whorl count and higher rib count (Table 19), Furthermore Elsoathera has an almost unilobed kidney, tapered at the apex and the primary ureter as a Straight tube (Figs 109f, 10Se). In contrast the kidney of Cralopa is weakly bilobed with the apex of the pericardial lobe and initial part of the primary ureter weakly teflexed (Figs 96e, 99e, 102). Cralopa has a large, cylindrical penis with two large 149 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE (QL'S-RLY) Ole OHe'S IRS (LL's-17'S) BZ FHS (95°-26'9) OLVUFUZ'L (zs'8-61'R) sorarae8 ORL 06'L arg OLLVe N/G (pe0-15°0) HeU'UFIOO ica (891-990) OLO0FL9'O (spd) EL OF O (oe0-Le0) OMOOFREO fe 0 tro 10 (ut) HLGIM TOTmHnwa ors (RUS-LV'p) OCEOF INE Ob (ies-i't) Sue OF WS (169-+1'S) e207 E09 (c6'L-14"9) OLS 0769 sto SZ OE'ST (LE ZI-90'6) POL OFZ OL LV6 qTL6 (62 8-48'L) SLY'UFZER isel WHW/SEld “AONCUSISSUTYVD VdOTVYI CNV ‘AON ‘dS SISNIAVLAAVN VIOTVED ‘(r9RT *XOD) SIS ss (es-ep) EC ETE OP oP (65-95) OS TFELS (SL-Rr) THEFT (zx-L9) CLPFETL OL 69 ug (901-06) £96 OL 69 (yo1-tR) 00 1170'S6 601 SAIN (gz'-€2'0) SzOF9Z() (sou-6e'0) O900FRF'O is'0 (epa-erw) (9E0-17'0) O100FHE) (yer iza) PS FRTO Opt 6rd S10 (310-410) 60K 07910 (gr0--0) Oz0'0F91'0 ral] (sz0-17'0) vlourEcO 20 OLLVa MME/dS (20 1-r6'0) SOOT (36°0-2L'0) LEOOFGR'O OZU OFT (sv ory) 670 UF86'0 (zy 1-760) £200" UF 86'0 LW sso es) (si 1-460) ZS0°0F€0'! rou LL0 (61 1-z0'1) sg0'0Ft P60 (wus) HACIA THOHM AGOG (og0-17'0) (zs0-150) Sry UF9z th SU UFZS 0) (r9'0-060) (19'u-ss'0) PLrorero 910'0F85°0 80 aro (ep'0-¥E'0) (Ls-0-480) sprorne'y SENOFLS'O (peO- 170) (190-660) B100FL2°0 +10'0F9S'0 (pou-1z0) (ss'0-£5°0) REO'OFLEO HOOFPS'O er 950 ovo zs'0 R00 0 (410-10) (as'‘0-z0) o1o'0F90'0 S1O'OFLS'0 (LU0-£1'0) (L9-S9'0) O20 F510 QLOOF99'O 40 +90 (gsa-ss'0) 70 SLOUFLSU 17'0 £90 (wu) NOISQYLOWd OLLVH O/H aulds (ADNVU GN WAS ‘NVAIN) (6Le-Zee) SETUFISE (ps't—9'2) Pol 0FOT'E 867 (o'f-"5"€) STVUFLY'E (pS E-R6T) yOV'OF1Z'E (LE'E-H7'E) OT UFEE CE wye SL'l 991 (eve-€L7z) ZCr0F96E (osz-erz) 0900 6P'7 ara (ER E-1F°E) OIZOFZ9'E 9o% (usu) waLAWVIG (96'1-0L'D OE OFER (LV2-8S'L) ZZ1UF PRT BST (607-407) Se O-LU'Z (Ly7-zo 650° OF [R'E (ex i-se) LZOOFOR'T P07 060 060 (cg 1-8s"1) Ssy'0FL9'L (991-291) vZz0'0-49'1 stl (zz'7-L8'1) SLUOFSUT worl (ui) HOH (ail pie 2) oe Coll pit €) “H/LP BILE aie (r/l PRI £) +P (R/ p-R/L ©) +P vr +R/1 +P (H6/1 PBI) BILE (48/1 PILE) +P STHOHM MONOULS VdO'1VAI Ni NOLLYIYVA TWOOT - 0c ATAVL n nt a ot SNAWIDadS 40 HAG WON L9otP LOWY BOARD URAIQUII AL, OL7r71 OWN SOAR. MOPTNL 19671 OWV sadsnp aay 96bRT ONY nduniy Orgc OW (197 OWN (adAioyort) BESizL WY yo.9Au] pssapava ndojndy 0SL91 ONWO (adAyoyoH) £SL91]OWWO anindrey 1 siskaan)ndey pdojp2) TsRoL OWWO SHA BAUNEL PS60l ONWNO “A'S yRppaOy SH97LOWWO. “g's BuNQWODDD: UR91ZL WV SDARD YEGRSSAA (adAqoj0H) GOSE9 WY pnoag sisuapmouys vdo}D4) AWYN longtitudinal pilasters (Figs 96b, 99b, 102b) in contrast to Elsothera which has a pear-shaped penis internally divided by an apical collar (Figs 105e. 109b). The epiphallus - vas deferens junc- tion of Elsorhera is highly modified (Figs 105b, 109a) while that of Cralopa is scarcely differen- tiated (Figs 96a, 99a, 102¢). Egtlomen globosa also has prominent radial ribs and a closed um- bilicus but differs m having beaded microsculp- ture (Figs 128g, 131f). Cralopa stroudensis (Cox, 1564) (Figs 95-97; Tables 19, 20) Helix strondensis Cox, 1864, p. 20) Tryon, 1887, p.25. pl. 4.figs. 28,29. Gvracoehlea stroudensis (Cox); Hedley, 1924, p. 217, pl. 31, figs 25-27; Iredale, 1937a, p. 323. Cralepa stroudensis (Cox): Tredale, 194 1a, p, 269, COMPARISONS C. Stroudensis can be distinguished from its congeners by high rib count, relatively large size and moderate spire elevation, C. kaputarensis is smaller, has a lower rib count and almost flat spire (Pig. 98c), C. carlesst is larger with reduced rib count and strongly elevated spire (Fig. 101c). Anatomically the differences are subtle. C. stroudensis (Fig. 96b) has a large cylindrical penis with two unequal, longitudinal pilasters, and a penial retractor muscle inserted on the Penis apex; C. kuputarensis (Fig 99b) has a large penis with two large longitudinal pilasters which anastomose and divide into smaller pilasters basally. and a penial retractor muscle which is inserted on the epiphallus: C. carless’ (Fig. 102b) has the penial retractor musele inserted on the penis which has two slender equal pilasters and a low central pustular swelling internally. TYPE MATERIAL Hororyre: AMCASS00. Stroud. NSW. Coll, Rev. RL. King Height 1.62 mm, diameter 2.56 mm, H/D ratio 0.63, D/U rato $30), whorls 4. PARATYfps: AMC225, 4 specimens. sume data us holotype. OTHER MATERIAL Sherrard Falls. Dorriga, NSW. 600 m, warm temperile rainforest. in liter (1. OMMOLG778, 13 Noy 1483. AM'OM- ABRS);c. 24.5: km [rom Goom- hurra, on Goomburra S.P. road. SEO (2754S. ISP 2ZVE) G00 m CNVF/Palms (8. OMMO) 2695, OMMO1/ 1128, AMC136816, 7 Dee 1981. AM/OM - ABRS): Yessabah Limestone Cave. Shery oud Quar- MEMOIRS OF THE QUEENSLAND MUSEUM ty, Wal Kempsey NSW (31 °U6°S, 152°42'E) ground litter. in and outof cave (2. AMC121680, 9 Oct. 1979, P.H. Colman, J. Stanisic); Lower Ballanjui Falls Cir- cuit, Binna Burra, Lamington N_P., SEQ, NVF, litter (1, QMMOQ16893, J. Stanisic. D, Potter, J, Chaseling); Bunya Nts N.P. (27°53°S, 151°33°E) NVF (1, QM- MQ6072, QMMO6278. 5 Mar 1976, M.J. Bishop): initial part of Kweebani Caves walk, Binna Burra, Lamington N.P_, SEQ NVF, under log (1, QMMO- 16888, 9 Mar 1984 J, Stamsic); top of Condamine River Valley, SEQ (28°1S°S, 152°29°B) (2, QM- MOlL0410. AMC128902, Mar 1981, AM/QM - ABRS); Cherry Plain - Westcott Plain Cireuit, Bunya Mis N.P.. SEQ, NVF/MVF. under logs (43, OM- MOIGSS7. 6 Nov 1985, J. Stanisic, D. Potter); Dan- dabah, Bunya Mis N.P.. SEQ. NVF/under logs (13, QMMO 16846, 12 Mav 1984. J. Stanisic, D. Patter): Big Falls Circuit Bunya Mts N.P., SEQ, SNVF, under logs (27, OQMMOIGSS1, 5 Nov 1985, L Stanisic, D. Potter): Hails Plain, E of Emu Vale. SEQ, CNVF_ under log (1, OMMO16891, 4 May 1986, J. Stanisic, J. Chaselingi: Cunningham's Gap N.P.. SEQ, near monument. 755 m. (28°04 S$. 152°24°E) SNVF/Ferns (1, QO€MMO12685, 7 Dee |Y81, AM/QM - ABRS); c, {8 km trom Legume. on Acacia Plateau Rd, Koreelah 5.F.. SEO... SEVT. NVF. MVF (1. OMMO12718. 8 Dee 1981, AM/OM - ABRS): Bellengen River S.F., NSW (30°25°S, 152°44°E) dense rainforest (1, QM- MOIO814,. AMC128389, 12 Mar, 1981, AM/QM - ABRS): Bunya Mis S.P., SEQ (13, AMC63869_ 8 Feb 1961_ D.F. MeMichuel): Rocky Ck, Red Scrub Flora Reserve, NSW (28°38'S, 153°20°E) (1, AMC128867, 2() Mar 1981, AM/QM - ABRS); Port Macquarie (1, AMC36564. ex Hedley), Dandabah, Bunya Mis, SEQ (26°53'S. 151°36°E) (158, QMMQi2281, AMC136005, 3) Sept 1952, AM/QM - ABRS); Port Macquarie (1. AMC63855. ex Cox): Koreelah S.F., NSW. SEVT/MVF (28°21°S. 152°24'E) (11, AMCI28557, OMMOI(N9S4 15 Mar 1981, AM/QM - ABRS); Natural Arch, Carrai Caves, near Kempsey, NSW. litter (1, AMC152198. 22 Feb 1980. P.H, Cal- man), DIAGNOsts Shell diameter 2.26-3.83 mm (mean 2.89 mm) with 3 7’°S to 4 14+ (mean 44+) loosely coiled whorls. Apex and early spire (Fig. ¥5a) slightly elevated. SP, BWW ratio 0.14-0.25 (mean (1.19), height 1.45-2.22 mm (mean 1.71 mm). H/D ratio 0524167 (mean 0.60). Last whorl descending much more mupidly. Protoconch of 11/2 to [S5’8 whorls, mean diameter 593.3 jum at] 12 whorls. Apical sculpture (Fig. 95d) of numerous. high. curved racial ribs, becoming slightly more crowded toward the nuclear-post SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 151 Imm FIG. 95. Cralopa stroudensis (Cox, 1864). a—c, Stroud, NSW. AMC63500, holotype; d-f, Dandabah, Bunya Mts, SEQ. QMMO12281. a—c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, central and lateral teeth. Scale lines as marked. 152 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 94. Cralopa stroudensis (Cox, 1864). Dandabah, Bunya Mts N.P.. SEQ. QMMQ12281. a, genitalia; b. penis interior; c, hermaphroditic duct, carrefour and talon: d, ovotestis; e, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 26°S 27°S 29°S 30°S 31°S > 500m Nalt. 33°S NN 151°E 152°E 153°E FIG, 97. Distribution of Cralopa stroudensis (Cox, 1864). nuclear junction, crossed by fine spiral cords. Post nuclear sculpture (Fig. 95e) of numerous, large, strongly protractively sinuated radial ribs 69-109 (mean 91.4) ribs on the body whorl. Ribs/mm 7.84-13.57 (mean 10.08). Micros- culpture (Fig. 95e) of fine radial riblets, 3-10 between each pair of major ribs, crossed by numerous, regularly spaced spiral cords. Um- bilicus (Fig. 95b) very small to closed, diameter 0.29-0.43 mm (mean 0.36 mm), D/U ratio 7.86— 153 8.52 (mean 8.15). Sutures deep; whorls slightly shouldered above and rounded below the periphery (Fig. 95c). Aperture roundly lunate. Lip sinuous, strongly retracted at the sutural margin and forming an apertural sinus. Columel- la expanded and slightly channelled in older specimens, weakly reflected over the umbilicus. Parietal callus developed. Colour variable, white through straw-yellow to light-brown. Based on 10 measured adults. Epiphallus (Fig. 96a) not strongly differen- tiated from the vas deferens, entering the penis apically through a simple pore (Fig. 96b) sur- rounded by fleshy thickenings. Penis (Fig. 96b), internally with two large, unequal longitudinal pilasters, and a series of low, longitudinally ar- ranged pustulations. Penial retractor muscle in- serting on the penis head adjacent to the epiphallus. Free oviduct (Fig. 96a) very long, twisted and muscularised, internally with a spon- gy pad-like thickening. Vagina long with inter- nal lamellar thickenings. Radula (Fig. 95a) as for genus. Based on two dissected specimens (QMMO- 12281). RANGE AND HABITAT Cool subtropical notophyll vine forests of the Great Dividing Range between Barrington Tops, NSW and Bunya Mts, SEQ; the McPherson Ran- ges of the QLD/NSW border; and the limestone tidge which extends intermittently from Yes- sabah to Carrai, W of Kempsey, NSW. C. stroudensis lives under logs and sometimes occurs in large numbers per site. REMARKS There is significant conchological variation between populations. However insufficient material is available to investigate its extent. The Yessabah Cave shells (AMC121680) are noticeably larger, possibly due to the abundant supply of calcium for shell construction. Cralopa kaputarensis sp. nov. (Figs 98-100; Tables 19, 20) ETYMOLOGY For the type locality, Mt Kaputar. COMPARISONS Cralopa kaputarensis is separated from C. stroudensis in discussion of that species above. The barrel-shaped penis is a conspicuous char- acter which identifies C. kaputarensis. C. car- 154 MEMOIRS OF THE QUEENSLAND MUSEUM ae er ] “ey Mei 7 is “i? had . a Eis fe ; F é “ 05mm Cc FIG. 98. Cralopa kaputarensis sp. noy. ac, Summit, Mt Kaputar, NSW. QMMO16753, holotype; d-f, Dawson Springs, Mt Kaputar, NSW. QMMO16742, paratypes. a—c, entire shell: d, apical sculpture; e, post nuclear sculpture; f, central and lateral teeth. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 155 FIG, 99. Cralopa kaputarensis sp. nov. Dawson Springs, Mt. Kaputar, NSW. QMMO16742, paratype. a, genitalia; b, penis interior; c, ovotestis; d, talon, carrefour and hermaphroditic duct; e, pallial cavity. Scale lines as marked. FIG, 100. Distribution of Cralopa kapurarensts sp. nov. on the Mt Kaputar Plateau. lessi is much larger with more elevated spire and flatter whorls. Type MATERIAL HovotyPe: QMM0O16753, summit, Mt Kaputar, NSW. 1500 m, among rocks, grass and litter. Col- lected 9 Nov 1983, AM/QM - ABRS. Height 0.90 mm, diameter 1.66 mm, H/D ratio 0.54, whorls 4+. PARATYPES: Dawson Springs. Mt Kaputar, NSW. 1300 m, under logs, in fern guily (3, QMMO16742. 8 Nov 1953, AM/QM - ABRS); Juker’s Spring, ML. Kaputar, NSW, 1000 m, edges of rocky creek bed, among lichen (1, QMMO1L6750, 8 Nov 1983, AM/QM - ABRS). DIAGNos!s Shell diameter 1.66-1.75 mm (mean 1.71 mm), with 4+ to 4 1/8+ (mean 4+) loosely coiled whorls, last descending slowly. Early spire and apex (Fig. 98a) very slightly elevated, SP/BWW ratio 0.15-0.19 (mean 0.17), height 0.90 mm. Protoconch of | 1/2 whorls, mean diameter 734.0 ym at 1 1/2 whorls. Apical sculpture (Fig. 98d) of slightly curved, radial ribs and fine low spiral cords, more noticeable toward the sutures. Postnuclear sculpture (Fig. 98e) of large, crowded, strongly protractiyely sinuated radial ribs, 69-80 (mean 74.5) ribs on the body whorl. Ribs/mm 12.57-15.30 (mean 13.94), Microsculpture (Fig. 98e) of fine radial riblets, 6-10 between each pair of major ribs, crossed by crowded. regularly spaced, low spiral cords. Umbilicus (Fig. 98b) reduced to a slender crack, MEMOIRS OF THE QUEENSLAND MUSEUM more open in sub-adull specimens, Sutures im- pressed, Whorls rounded above and below the periphery (Fig. 98c). Aperture roundly lunate. Lip sinuate, retracted at the sutural margin to form an apertural sinus. Columella dilated and reflected over the umbilicus. Parietal callus well developed. Colour dark brown. Based on 2 measured adults. Vas deferens a thin tube, giving rise to a weak- ly differentiated epiphallus. Epiphallus entering penis apically through a simple pore. Penis broadly tubular, internally with two large lon- gitudinal pilasters which anastomose, and divide into numerous smaller pilasters basally. Penial retractor muscle (Fig. 99b) inserted on the epiphallus. Female reproductive system small compared with the penis, Radula (Fig. 98f) with central tooth much smaller than laterals. Based on one dissected specimen (QMMO 16742). RANGE AND HABITAT Only on the Mt Kaputar Plateau, near Narrabri, NSW. Under logs, rocks, and among lichen on the edges of springs. REMARKS C. kaputarensis is closely related to C. stroudensis and is probably a recent derivative. Cralopa carlessi sp. nov- (Figs 101-103; Tables 19,20) ETYMOLOGY For Terry Carless who collected specimens for this study, COMPARISONS The flatter more loosely coiled whorls, elevated spire and reduced number of radial ribs on the body whorl characterise C. carlessi. C, kaputarensis has an almost flat spire (Fig. 98c) and is much smaller (Table 19), Type MATERIAL HoLoTYPe: AMC 123538, Inverell, NSW. Collected by C.T. Musson, 1889. Height of shell 2.04 mm, diameter 3.62 mm, H/D ratio 0.56, whorls 4 1/2. PARATYPES: AMC153716, 2 specimens, same col- lection data as holotype: Hills of Glenrock Stn, NSW (3141S, 151°24°10"E) all. ec. 600 m, under logs and limestone slabs and litter, dry, semi-open serub (2, AMC144200, 17 Jun 1984, P.H. Colman); Wom- beyan Caves, W of Mittagong, NSW, steep scrub and grass covered limestone slope, in litter outside large SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 157 FIG. 101. Cralopa carlessi sp. nov. a—c, Inverell, NSW. AMC123538, holotype; d—e, Hills of Glenrock Stn, NSW. AMC144200, paratype; f, Inverell, NSW. AMC3640, paratype. a—c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, central and lateral teeth. Scales lines as marked. 158 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 102. Cralopa carlessi sp. nov. Hills of Glenrock Stn, NSW. QMMO144200, paratype. a, genitalia; b, penis interior; c, talon and hermaphroditic duct; d, ovotestis; e, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 29°S 30°S 31°SsS } X IN9 32°S RS NSS 33°S 34°S 3257S 36°S 148°E 149°S 150°E 151°5 1s2°s FIG. 103. Distribution of Cralopa carlessi sp. nov. cave (2, AMC142967, 28 Dec 1981, M. Shea); West side Kowmung River at Gridiron Bends, near Tuglow Caves. NSW, in litter and soil on limestone bluffs (7, AMC124279, 12 Dec 1979, W.F. Ponder, P.H. Col- man); Attunga, NSW (2, AMC28496, C. Laseron); Inverell, NSW (3, AMC123610, C.T. Musson); In- verell, NSW (11, AMC3640, W.S. Duncan); c. 1 km N of Wee Jasper, NSW, on Careys Caves Rd, in limestone scrub (2, AMC142961, 20 Oct 1980, P.H. Colman, J. Hall). OTHER MATERIAL Entrance to Devil’s Coachhouse, Jenolan Caves, NSW, litter (2. QMMO17295, 13 Dec 1979, J. Stanisic). DIAGNOSIS Shell diameter 2.98—3.79 mm (mean 3.30 mm) with 3 3/4 to 4 1/2 (mean 4 1/8) loosely coiled whorls, last descending more rapidly. Apex and 159 early spire (Fig. 101b) very slightly to moderate- ly elevated, SP/BWW ratio 0.21-0.64 (mean 0.33), height 1.58-2.17 mm (mean 1.84 mm), H/D ratio 0.48—0.61 (mean 0.56). Protoconch of 1 3/8 to 1 5/8 whorls, mean diameter 727.3 um at 1 1/2 whorls. Apical sculpture (Fig. 101d) of thin, curved, regularly spaced radial ribs, crossed by low spiral cords. Postnuclear sculpture (Fig. 101e) of numerous large, strongly protractively sinuated radial ribs, 43-82 (mean 59.1) ribs on the body whorl. Ribs/mm 4.17-7.95 (mean 5.77). Microsculpture (Fig. 101e) of fine radial riblets, 6—12 between each pair of major ribs, and fine crowded spiral cords which are higher at their intersection with the radials, producing a beaded effect. Sculpture continuous on the base. Umbilicus (Fig. 101b) narrow, D/U ratio 4.78- 7.56 (mean 5.98), diameter 0.41—0.74 mm (mean 0.56 mm), reduced to a lateral crack, or closed by the columellar reflection. Sutures impressed. Whorls rounded above and below the periphery (Fig. 101c). Aperture ovately lunate. Lip sinuous, thickened at the columellar edge. Columella dilated, reflected over the umbilicus, retracted at the suture to form a small sinus. Parietal callus developed. Colour yellowy- brown. Based on 22 measured adults. Epiphallus (Fig. 102a) weakly differentiated from vas deferens, entering penis apically through a pore surrounded by fleshy thickenings. Penis (Fig. 102b) cylindrical, without a sheath, internally with two slender equal longitudinal pilasters and an apical spongy thickening which may act as a verge, otherwise pustulose. Penial retractor muscle inserting on the penis head. Vagina tubular, about as long as the free oviduct. Atrium relatively long. Radula (Fig. 101f) as for genus. Based on one dissected (AMC144200). specimen RANGE AND HABITAT Confined to a series of isolated limestone out- crops on the western edge of the Great Dividing Range, from north of Goulburn to north of Tam- worth, NSW; under logs and rocks and among soil and litter. REMARKS Various populations of C. carlessi show some notable variation in shell characters (Table 20). However the patterns are largely mosaic and probably reflect differences in isolated popula- tions. In contrast, specimens from the Tuglow Caves (AMC124279) have a more elevated spire lan and more open umbilicus which are uncorrelated changes that may indicate speciation trends in distant populations. Clsothera Iredale, 1933 Elsothera Iredale, 1933, p. 53; Iredale, 1937a, p, 324: Iredale, 19414, p. 267; Kershaw, 19564, p. 140; Burch, 1976, p. 132. TYPE SPECIES Helix sericatula Pfeiffer, 1850; by original designation. PREVIOUS STUDIES Elsothera was introduced for Heltx sericatula Pfeiffer, [850 from the Sydney region, NSW.. Iredale (1937a, 1939) added 9 species from southern Australia, and Kershaw (1956a) another 3 from Victoria based on shell features. A number of generic groups may be represented in Elsothera@ but this matter 1s beyond the scape of the present study. Elsothera is redefined to include HM. sericatula and two species from northeast NSW and SEQ. DIAGNOSIS Diameter 3,96—-6.64 mm, with an average of 4 1/4 to 4 3/4+ normal to loosely (gerithecata) coiled whorls. Apex and early spire flat (venithecata) to slightly elevated, Apical sculp- ture of erawded, weakly curved, high radial ribs and finer, more crowded, low spiral cords which continue onto the radial ribs. Radials vague to absent on initial part of protoconch. Adult sculp- ture of numerous, very crowded, high. protrac- tively sinuated radial ribs, Microsculpture of fine, high radial riblets and equally high, narrow, more widely spaced spiral cords. Umbilicus closed to barely open (nautilodea). Sutures im- pressed to distinctly channelled (geni/hecata). Whorls shouldered above and rounded below a weakly compressed periphery, last inflated and sometimes (veutilodea) with a Weak supra- peripheral sulcus. Lip simple, columellar margin teflected over the umbilicus, Colour ight to dark-brown With darker radial streaks, Terminal male genitalia with long, mus- cularised epiphallus, sometimes partially sheathed (genithecata), entering pens through a simple pore surrounded by a collar. Epiphallus- vus deferens junction swollen, complex with vas entering laterally, Penis short, tubular with api- cal bulb, internally with a spongy or fleshy collar constricting the entrance to the main penis cham- MEMOIRS OF THE QUEENSLAND MUSEUM ber. Penial bulb with numerous short, transverse (sericatula) or longitudinal (genithecata) pilasters. Penis proper with short, longitudinal thickenings and pustulations (genithecata), or only pustulations (sericatula). Vagina short, or long with an apical caecum (genirhecata). Radula with tricuspid central and lateral teeth, ceniral smaller; laterals with very large broad lanceolate mesocone, DISTRIBUTION AND ECOLOGY Elsathera has a wide distribution from just narth of the NSW/QLD border to south of Syd- ney, NSW. It is one of the few groups of east- coast charopids which inhabits drier, open forest. E. sericatula inhabits wet and dry scletophyll forest and £. genithecata, has been found in ‘open cucalypt forest’ in the Upper Richmond Range, NSW. Ability to adapt to der environ- ments gives Elsothera the opportunity to exploit a vast amount of relatively snail depauperate habitat. The genus is quite prabably more widespread than is indicated by the coverage presented here and detailed statements about distribution are inappropriate at this time. Life- styles of the three species reviewed are diverse, probably reflecting the overal) adaptability of the genus. COMPARISONS Elsothera has a shell with average number of whorls and numerous, weakly, protractively sinuated radial ribs. In contrast the conchologi- cally similar Cralapa has fewer whorls and fewer, bolder radial ribs which are strongly protractively sinuated, Radular characteristics of the two genera are similar. The constricting penial collar. vague penial pilasters, complex vas deferens-epiphallus junction, epiphallic sheath und vaginal caecum of Elsethera are all sig- nificant differences from the Cra/opa pattern of simple penis with Jarge longitudinal pilasters, barely differentiated epiphallus, and lack of ad- ditive structures. REMARKS Elsethera is most likely a rainforest-adapted group which has managed (oO colonise drier forests. The large brown radially ribbed shell and unilobed elongate kidney are characteristics of other rainforest genera such as Gyrocachlea and Nautiliropa. Large size, combined with a complete secondary ureter were probably im- portant preadaptations which have enabled El- sothera to exploit drier habitats, DA RATIO UMBILICAL wibrit (mm) RIDS/MM 14020. (1044-315) 10.4520.945 (8.87-19,39) 13,02 RIBS (44.725.04 (195-153) 168 Oe6 18 (156-183) mn) SPUWW RATIO (), t2eD.007 (0,080, 14) (sen? (0.10-0,25) Wd (ym) 14920029 (145-153) LP e IAI AR) (166-179) BODY WITORL 154 WIDTTE (MEAN, SEM AND RANGE) SPIKE PROTRUSION (mm) OUR 20027 (0.13-0,21) 254). 047 (0.17=0.43) ty f7D RATIO OST01D (055-059) 05320,010 (0.49-0.55) ost (mm) 4.UN)24,026 513a0,117 (4.95.58) DIAMETER (4.464.005) Ao HEIGHT (nm) 2.29e01,035 (2.22234) 2.730.089 (243-298) TABLE 2} - LOCAL VARIATION IN ELSOTHERA SPRICATULA (PFEIFFER, 1451), ELSOTHERA NAUITILODEA (COX, Ith) ANDELSOTHERA GENITHECATA SP NOV ay WHORLS: aye a+ (41/44 04 412+ {4 1/44 5yHr) NUMHER OF STECIMENS, 5 NAME Elsothera teriedtula Australia AMNI 1YATIS AMC 103604 Elsathera miuitledeu Clarence River Shark Istund SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 161 as Elsothera sericatula (Pfeiffer, 1850) ae (Figs 104-105; Tables 19, 21) ~ Helix sericauida Pfeiffer, 1850, p. 127; Reeve, 1852, > pl. 132, sp, 812, 2@ Elsothera serieatula (Pfeiffer); Iredale, 1933,p. 53; =: booed # Iredale, 19374, p. 324; Iredale, 1941a, p. 269, fig, = 5. aa 5 £5 24 COMPARISONS 7.) grea A full treatment is deferred until a more com- prehensive review of the genus is completed. E. Fe sericatula resembles E. nautilodea, but differs in ae having the umbilicus closed (Fig. 104b) rather = #8 82 § €& than slightly open (Fig. 106b). ‘E.’ biretracta (Mousson, 1869) from the Illawarra region, Za NSW ts conchologically similar to E. sericatula = #8 and may prove to be conspecific. Species of = 5S Cralopa are similar in having a closed umbilicus and prominent radial sculpture but are smaller re with fewer, bolder ribs on the body whorl. The gc lectotype was selected from a lot of syntypes in 2 #2 22 8 = _ the British Muscum. ze TYPE MATERIAL x od ; + 4 Lectotype: BMNH 1987055, Australia, H. Cuming oe Collection. PARALECTOTYPES: BMNH 1987055, 3 specimens, Ee same collection dala as lectotype. zl 2 ig == 2 2 DescriPtion (of lectotype) Shell small, diameter 4.64 mm, with 4 1/2 “ normally coiled whorls, the last descending 53 more rapidly. Apex flat, spire slightly elevated, 2 88 €2 € & $SP/BWW ratio 0.11, height 2.39 mm H/D ratio. 7 Protoconch of 1 1/2 whorls, diameter 739 jum. Apical sculpture eroded with traces of radials 22 near the sutures (paralectotypes show a pattern = ae & = = & _ of high, slightly curved radials and low spirals). ae ? “Post nuclear sculpture of numerous, closely spaced, prominent, protractively sinuated radial 2 ribs, 190 on the body whorl. Ribs/mm 13.02. As Els ew 2 Microsculpture of fine radial riblets, 5-6 be- > 52 5 4 ~*~ — tween each pair of major ribs, and low crowded spiral cords. Umbilicus closed by reflection of the columellar margin. Sutures impressed. 7 a Whorls shouldered above and rounded below a slightly laterally compressed periphery. : Aperture ovately lunate, Lip simple. columellar 2g #2 = e.f 2f _} margin slightly dilated reflected over the um- ep 2h & GEC ra #2 bilicus. Parietal callus strongly developed. gi gs 2823 22 32 Colourlight-grey with darker irregularly spaced 2 ga5" 2° = radial markings. MEMOIRS OF THE QUEENSLAND MUSEUM 162 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 163 FIG. 105. Elsothera sericatula (Pfeiffer, 1850). Clive Park, Northbridge, NSW. AMC152205. a, genitalia; b. details of vas deferens - epiphallus junction; c. talon and carrefour: d, ovotestis; e, details of penis interior; f, pallial cavity. Scale lines as marked, FIG. 104, Elsothera sericatula (Pfeiffer, (850). a-c, Australia. BMNH 1987055, lectotype; c, gh, Clive Park, Northbridge, NSW, AMC152205; e-f. Shark Is, Sydney, NSW. AMC103604. a-c. entire shell: d, jaw: e, apical sculpture; f, post nuclear sculpture: g. central and lateral teeth: h. marginal teeth, Scale lines as marked, 164 OTHER MATERIAL STUDIED Shark Is, Sydney NSW (13, AMC103604, ex J. Brazier coll); Clive Park, Northbridge, NSW, in fairly dense bush (3. AMC152205, 19 Oct 1969, W.F. Ponder, D.P. Fairfax), ANATOMICAL DIAGNOSIS Terminal male genitalia with a muscular epiphallus (Fig. 105a) which is partly coiled before entering the penis apically. Vas deferens/epiphallus junction (Fig. 105b) com- plex with vas deferens entering epiphallus laterally through a large cup-shaped pilaster. Penial retractor muscle inserting at the epiphal- lus/penis junction. Penis (Fig. 105e) short, tubular with an apical bulb, internally with a simple epiphallic entrance (Fig. 105e) and numerous short transverse pilasters. A large spongy collar constricts the entrance to the lower penial chamber which has _ fine spongy pustulations arranged in vague longitudinal rows (Fig. 105e). Vagina short with fleshy, pustular interior. Radula (Fig. 104g,h) with tricuspid central and lateral teeth, central smaller. Based on 2 dissected specimens (AMC- 152205). RANGE AND HABITAT Closed and open forest from Cronulla to Gos- ford, NSW, extending westward to at least the Great Dividing Range. E. sericatula is a com- paratively adaptable species and still survives in disturbed areas where microenvironments are suitable. It lives in the litter, among leaves and accummulated debris. REMARKS E. sericatula has been reported from Victoria (Gabriel, 1930) and Tasmania (Petterd and Hed- ley, 1909). However more recent surveys of these regional faunas (Smith and Kershaw, 1979, 1985) do not list this species and I regard the earlier records as erroneous. Elsothera nautilodea (Cox, 1866) comb. nov. (Figs 106-107; Tables 19, 21) Helix nautilodea Cox, 1866a, p. 47. Helix nautiloides Cox, 1866b, p. 696; non Ferussac, 1850. Helix inusta Cox, 1868, p. 13, pl. 10, fig. 3. Helix nautilodes (sic) Iredale, 1937a, p. 324. Elsothera inusta (Cox); Iredale, 1937a, p. 324; Iredale, 194 1a, p. 269. MEMOIRS OF THE QUEENSLAND MUSEUM COMPARISONS E. nautilodea is conchologically similar to E. sericatula. However the species are allopatric and misidentification in the field is not likely. E. nautilodea has a brownish shell with very nar- row umbilicus whereas the shell of E. sericatula is light grey with darker streaks and a completely closed umbilicus. E. genithecata is larger with relatively fewer whorls, more widely spaced ribs and channelled sutures (Fig. 108a). Cralopa stroudensis is smaller with fewer and bolder ribs. PREVIOUS STUDIES Helix nautilodea appeared on 1 January, 1866. In the same year, the same article appeared in the Proceedings of the Zoological Society (April 24, 1866) with the name changed to H. nautiloides. Subsequently Cox (1868) intro- duced Helix inusta as a replacement name for H. nautiloides after realising that the latter was preoccupied. I consider nautilodea to be the correct name for this species as there was never any indication by Cox that H. nautiloides was an emendation of the earlier name. The lec- totype was chosen from topotypic material in the Cox collection. TYPE MATERIAL LectTotyPe: AMC136903, Clarence River, NSW, ex Cox. Height of shell 2.81 mm, diameter 5.24 mm, H/D ratio 0.54, whorls 4 3/4-. OTHER MATERIAL Clarence Rv (2, AMC152170, ex C.F. McLauchlan); Grafton (2, AMC152171, Jan 1904, S.W. Jackson); Clarence Rv (6, AMC152172, G. Thornley); South Grafton (3, AMC152173, ex Nat. Mus, Vict.); Har- wood Id, Clarence Rv (73, AMC152174, Feb 1961, A.A. Cameron); North Arm, Clarence Rv (9, AMC152212); on E side of tree 5 ft from ground Harwood Is, Clarence Rv, NSW (4, AMC152218, Jan 1970, A.A. Cameron), DIAGNOSIS Shell small, diameter 4.05—5.24 mm (mean 4.43 mm), of 4 1/4 to 4 3/4- (mean 4 3/8+) normally coiled whorls. Last whorl descending rapidly. Apex and spire (Fig. 106c) almost flat, SP/BWW ratio 0.02-0.12 (mean 0.09), height 2,.17-2.81 mm (mean 2.41 mm). H/D ratio 0.50— 0.57 (mean 0.54). Protoconch of 1 1/2 to 1 5/8 whorls, mean diameter 718.6 wm at 1 1/2 whorls. Apical sculpture (Fig. 106d) of curved radials and low, crowded spirals. Post nuclear sculpture (Fig. 106e) of numerous, closely spaced, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 165 2mm c FIG. 106. Elsothera nautilodea (Cox, 1866). a—c, Clarence River, NSW. AMC136903, lectotype. d-e, Harwood Island, Clarence River, NSW. AMC152174. a-c, entire shell; d, apical sculpture; e, post nuclear sculpture. Scale lines as marked. lity 151°E 152°E 153°E 154°E FIG, 107. Distnbution of Elsoriera nautiladea (Cox, 1866). prominent, protractively sinuated radial ribs, 129-190 (mean 151.9) ribs on the body whorl, Ribs/mm 10.02-11,53 (mean 10.94). Micros- culpture (Fig. 106e) of crowded spiral cords and thread-like radial riblets, 4—9 between each pair of major ribs. Umbilicus (Fig. 106b) a lateral crack to narrawly open, rarely completely closed. Sutures impressed. Whorls rounded below and strongly shouldered aboye a some- whut laterally compressed periphery with a weak subsutural sulcus formed in some specimens (Fig. 106c). Aperture roundly lunate. Lip simple, columella expanded and reflected over the umiblicus in some specimens. Parietal callus well developed. Colour brown with darker radial streaks, Based on 1] measured adults. "Animal almost black, not quite, tentacles well defined and same colour as animal, Foot lighter in colour". (Based on field notes by A.A. Cameron, AMC152174), No sofi parts available for dissection. RANGE AND FLABITAT In spite of intensive collecting in northern NSW over Six years, £. nauytifodca has not been located. Earlier collections indicate the species on Harwood Island, at the mouth of the Clarence MEMOIRS OF THE QUEENSLAND MUSEUM River. Live specimens have been taken semiar- boreally. Elsothera genithecata sp. nov. (Figs 108-110; Tables 19, 21) ETYMOLOGY Latin thecatus, encased; referring to the sheathed epiphallus. COMPARISONS E. genithecatais noteasily confused with sym- patric charopids. Large size, small number of whorls, channelled sutures, well developed aper- tural Sinus and closed umbilicus are not com- bined in other northern NSW charopids. &. naulilodea is smaller, has a less inflated body whorl and lacks sutural channelling. Gyrocochlea and Nautiliropa which have gross- ly similar adult sculpture, have a very depressed to sunken spire and wide U-shaped umbilicus. TYPE MATERIAL HoLotyer: AMC152165, Byangum, northern NSW, collected by Lower, ex Cox. Height ofshell 2.77 mm. diameter 5.63 mm, H/D satio 0.49, whorls 4 1/4, PARATYPFS: AMC153717, 2 specimens. same data as holotype: Red Scrub Flora Reserve, Whian Whian S.F., N of Lismore NSW (28°38'S. 153°19° EB) 210 m, litter (2, AMCI53718, 15 May 1976, P.H. Colman, 1. Loch); Mt Warning N.P., NSW, near base, (28°24°S, 153°16°E) NVF/Palms (2, OMMO10484, OMMO- 10502, 19 Mar 1981, AM/QM - ABRS): Richmond Runge. S.F.. NSW (28°30°S. 152°437E) 400 m, open eucalypt forest (1, QMMOo6267. 9 Apr 1976, MAI. Bishop): Big Scrub, Whian Whjan S.F.. N NSW (28°38°S, 153°19°30"E) SNVEF. under logs (1, OMMO16772. 15 Nov 19583. AM/OM - ABRS), Mt Tamborine, SEQ, in floor litter in rainforest (AMC 152164, 14 Sept 1979, PH. Colman): Upper Tweed Rv, NSW (2. AMC(S2147. Perterd! ex Cox): Emigrant Ck, Richmond Rv, NSW (4, AMCIS2168, ex Coa’): Byron Bay Scrubs, NSW (1, AMC152169, 1900, S.W, Jackson). OTHER MATERIAL Lower Ballunjui Falls Circuit, Binna Burra, Lamington W.P., SEQ. NVF, litler (1, OMMO16894, A Sept LY8S. J. Stanisic, D. Potter, J. Chaseling): FIG. 108. Elosthera genitheeuta sp. nov. a-c, Byangum, NSW. AMC1521 65. holotype: d, g-h. Richmond Range S.F.. NSW. OMMQ6267, paralype: e-l, Mt Warning N.P.. NSW. QMMOLOSO2, paratype, a-c. entire shell: d, marginal teeth: e. apical sculpture: f. post nuclear sculpture, geh, central. and lateral leeth, Seule lines us murked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 2mm : We i, We: ~ * 168 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 109. Elsothera genithecata sp. nov. Big Scrub, Whian Whian S.F., NSW. QMMO16772, paratype. a, genitalia; b, penis interior; c, talon; d, ovotestis; e, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE ILE FIG. 110. Distribution of Elsathera genithecata sp. nov. Clurence River, NSW (error!) (9, AMC103600, ex Cox). DIAGNOSIS Shell large, diameter 4,.37-6.64 mm (mean 5.67 mm), with 4 to 4 5/8- (mean 4 1/4) loosely coiled whorls, the last whorl descending more rapidly and strongly inflated, Apex and spire flat, height 2.77-3.36 mm (mean 2.94 mm). H/D ratio 0.47-0.63 (mean 0.53). Protoconch of 1 1/2 whorls, mean diameter $21.3 wm. Apical sculpture (Fig. 108e) of high, slightly curved radial ribs. and low crowded spiral cords. Postnuclear sculpture (Fig. 10&f) of numerous crowded, high, protractively sinuated radial ribs, 111-153 (mean 132) ribs on the body whorl. Ribs/mm 5.83-8.64 (mean 7.24). Microsculp- iure (Fig. 108f) of fine radial mblets, 5—10 be- tween cach pair of major ribs. and crowded spiral cords which continue onto the apices of the major tadials. Umbilicus (Pig. 105b) closed or a very small lateral crack. Sutures deeply im- pressed. Whorls shouldered above and rounded below a slightly compressed periphery (Fig, 108c). Aperture roundly to ovately lunate, Lip sinuous, simple. retracted at the suture ta form an apertural sinus (Fig. 108a-b), Columella ver- lical and thickened, reflected over the narrow umbilical opening. Parictal callus weakly developed, Colour light-brown with irregularly spaced dark radial streaks, Based on 4 measured adults. Genitalia with vas deferens-epiphallus junc- 169 lion complex; vas deferens entering almost laterally through a large spongy pilaster (Fig. 109a), Epiphallus (Fig. 109a) long, convoluted with a large muscular sheath for the last half of its length, entering penis apically through a simple pore (Fig. 109b) surrounded by a circular collar, Penis (Fig. 109b) short, tubular with an apical bulb. Apical bulb internally with short, longitudinal pilasters and a transverse spongy collar constricting entrance to lower penial chamber. Lower penial chamber with numeraus. short longitudinal thickenings (Fig. 109b), and irregular pustulations nearer the atrium. Penial retractor muscle inserted on the epiphallic sheath. Vagina (Fig. 109a) long with a caecum containing a large internal spongy thickening, and an elongate basal extension. Radula (Fig. 108d.g,h) with expanded mesocones on central and lateral teeth. Based on 2 dissected specimens (AMC- 152165, QMMO16772). RANGE AND HABITAT Warm moist subtropical notophyll yine forests of the Big Scrub area between the Rich- mond and Tweed Rivers, NSW; cooler subtropi- cal forests of the Lamington Plateau, SEQ; palm dominated forest of Mt Tamborine, SEQ; and drier scrub of the Richmond Runge, NSW. I lives under logs. REMARKS E. genithecata is one of the largest charopids in subtropical rainforests, The channelled su- tures are an unusual develapment also seen in Negairea canaliculata (Fig. 12a.c) and a number of extralimital taxa. The significance of this feature is not known. The well-developed aper- tural sinus is shared locally only with Cralopu, but is more common in New Zealand charopids, Coenocharopa gen. noy, ETYMOLOGY Latin coeno, dirt; referring to adhering dip particles on shell. TYPF SPECIES Coenacharopa sordidus sp. Nov, DIAGNOSIS Shell diameter 1.91-4.43 mm, with mean whorl count 4- to 4 3/8. Whorls normally coiled, last descending. 170 Apex and spire slightly (mudtirediata, yes- sabaliensis) to conspicuously (alata) elevated, Apical sculpture of weak (yessabahensis) to prominent, curved radial ribs which may be crowded to moderately crowded (alata), more sa al the nuclear-postnuclear junction, or regularly, widely spaced (mutliradiata). Ribs with prominent periostracal blades. Secondary apical sculpture of crowded, squiggly spiral cords which are not continuous on the radials. Posinuclear sculpture of widely spaced (a/ara) to crowded (vessabahensis) weak radial ribs with prominent periostracal blades modified into wing-like processes in some species (alata), rarely lacking (vessabahensts),, Microsculpture of thread-like radials which may have strongly developed periostracal blades (multiradiara) and low, broad (a/ata) to high, narrow (multuradiaca) spiral cords, Dirt particles adhering to shell sur- face. Umbilicus narrow to moderately wide (tacromphala). U-shaped. Sutures impressed, whorls strongly to weakly shouldered, oc- casionally with a supraperipheral sulcus (mracro- mphala). Aperture roundly (o ovately tunate. Lip simple. margins convergent. Parictal callus strongly developed, Animal with foot and tail broad, bluntly tapered posteriorly. Terminal male genitalia with slender to expanded (sordidus) vas deferens. Epiphallus sometimes partially bound io penis sheath (macromphala, vlata), entering penis apically through two fleshy thickenings. Ponts large to very large (sarcdidus), tubular, with a sheath, internally with several longitudinal pilasters which are modified in some species (vordtdus, vessabahensis), Penial retractor musele inserted on or very near to the epiphal- lus’pems junction, Rudula with mesocone of the laterals greatly expanded in C. yessahaltensis. DIstaine Hos ANP ECOLoGy Warm temperate rainforests and thickets of the Yessabah-Carrai limestone outcrop, cool sub- tropicul noatuphyll vine forests of the Great Dividing Range (from Dorrigo to the Border Ranges) and McPherson Ranges. and warmer subtrapical notophyll vine forests (with Arguecurra) in Southern Queensland. Intragieneric symputry is complex when com- pared with other charopids cxamined in this study. Symputric combinations are as follows: C, sordidusiC, macromphala, C. sardidus/C. parvicestata. C, parvicastata/C, macromipltala, Co alata’ CO. multiradiata, C. multiradiatale, par- MEMOIRS OF THE QUEENSLAND MUSEUM vicastata, C. mulliradiata/C. vessabahensis and possibly C. alaia/C, parvicostar, No details of microsympatry are available although this is likely in the ease of C. serdidus and C, macro- mphala. C. vessabaltensis has an enlarged penis und modified penial surface structures which indicate a microsympatric congener, probably C. multiradiaia. Although sympatric with both C. multiradiata and C. parvicestaia, C. alata, is probably not sympatric ut the micrahabitat level. PATTERNS OF VARIATION Shell size and shape - Whorl counts correlite positively with fnereased Size except in C, mul- Uradiate. Spire protrusion is proportionately grealest in C, vlera, Umbilical width is uniform exceptin C_ macremphalawhere itis wider (Fig. | 14e). Shell Sculpture - The prominent ealeified sculpture of most ground-dwelling charopids serves lo lessen the chance of particle adherance. Positive correlation between wider rib spacing and increased height of periosiracal ¢xtension sugecsts that in Coenocharopa the periostracal clements may be compensating for the lack of primary rib sculpture. Nonetheless most species have their shells covered by a layer of adhering dirt particles. Reduced sculpture in C, yes- sabahensis correlates with living on rock sur- faces Where adherence of dirt particles may not be a major problem. C. multivadiata (Fig, 126d- {) most closely mimics the more typical sculp- ture as in Rhoaphadan and Gyracochlea. Significantly it 1 the ‘cleanest species of Caenocharopa. Genitalia - Variavion in reproductive anatomy is confined to termina! male genitalia, In Neairea. Gyrocochlea and Rhaphadon variation in penial surface among microsym- paters involved minor changes in features of the apical chamber of the penis together with small changes. in shape and size of the longitudinal pilasters, In contrast, allerations in Coenacharapa involve major shape changes in pilasters. The basic pattern of internal penial sculpture appears to be several longitudinal pilasters as in. macromphala (Fig. }15d). C. sordidas, which 1s probably micrasympatric with this species. hus a large U-shaped pilaster and several accessory longitudinal thickenings (Fig. 112d). Insertion of the penial retractor muscle dis- plays only minor shifts from the penis/epiphallus junction onto the epiphallus adjacent to the june- lion. The epiphallus 1s typical, bul in C. macro- (MEAN AND RANGE) PAHLP 22 - KANG OP VARIATION IN COFNOCHAROMA AND EGILOMEN SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 171 2 oe Sw its = mphala and C. alata is partially bound to the $F tt 4 penial sheath (Figs. 1 15a, 118b). = 82 aa 938 35. ne ' as Radula - The enlarged mesocones of the central and lateral teeth of C. yessabahensis (Fig. a o 2S ‘4 123d,h) resemble those in the sympatric 3 2 | Letomola contortus and Rhophodon kempseyen- L' COMPARISONS = “y x i Coenocharopa is distinct among genera = bs zk et aerss reviewed in having a basically ‘smooth’ shell 3 oe . F with strongly developed periostracal sculpture. Apical sculpture resembles Elsothera and Cralopa but these genera differ in having very z oi ote at prominent radial ribs on the postnuclear whorls, = od Se 25° «OFS and closed or nearly closed umbilici. Letomola has greatly reduced sculpture but is much = = =6& & = smaller, has modified apical sculpture, wide um- i oar Int sh of bilicus and several apertural barriers (Figs 67, ae 34 933 25222 3s 68). In the Mudlo Gap area, north of Gympie, SEQ ,the conchologically similar Lenwebbia is 5. So 2 2 Se se wow sympatric with Coenocharopa. However, Len- S57 = 3 2 oF feted ae webbia is distinguished by its modified apical ge" 8A eo 85 22 6252 25 $2 sculpture which consists of irregular pits and * wrinkles (Fig. 29g), and postnuclear sculpture 2 a Me. te which includes incised spiral grooves (Fig. 29e). ae § 3 a 3 2 #8 Coenocharopa shares some genital features with SE 23 ge am Ag Sigs 33 Cralapa. These include a tubular penis, simple = ae = “as — epiphallic entrance into the penis through large fleshy thickenings, and basic penial surface 2 ceseg 5S 8 & sculpture of longitudinal pilasters (fewer in ae a oe xi oe Cralopa) and background pustulations. ' BS #93 SE se. fe ed de The postnuclear sculpture in Coenocharopa is deciduous and often lacking in older and dead # bE mem eke ye specimens. The following key is based on new- 2 = a ee adult fresh shells and may not work for gerontic ££ 83 35 56 3222 42 52 88 or litter-retrieved specimens. e FB RR FTE FS KEY TO SPECIES OF COENOCHAROPA SE <2 nd e4 ef 24 sh 22 5h a 5 ek i Taal a 1.Shell with prominent wing-like periostracal a ¢ = S. processes (Fig. 1i7e.h); spire strongly % freee § z elevated (Fig. 1170) ccccsesccssseeneseenvens alata = + os ze = 2 af Fed 2a HOE = 42° "253 "2 “S72 Fe Shell without prominent wing-like periostra- cal processes; spire weakly elevated ..........2 =4 45 rer ®©® € 3 SF = 2.Apical sculpture of regularly spaced radial ribs, z= not crowded al the nuclear-postnuclear boundary (Fig. |26d)............,...multiradiata é ig p i i BR # 2 3 Apical sculpture of curved radial ribs which z 3= 329 £3 36 F? 2f 3 fF become crowded at the nuclear-postnuclear r aS & $42 4.8 & boundary (Fig. 120f)...cc.ccecsseseecesseeeareeneeed (72 3.Postnuclear seulpture of regular, widely spaced radial ribs with periostracal blades (Figs TV. 1246, T2QE), sssscasscsrsrnssresnsersocssmsesa rete Postnuclear sculpture of low, crowded radial ridges Withoul prominent periostracal blades (Fig. | 2B} neenenerenenenteeneernere yessahahensts 4.Shell larger, mean diameter 3.92 mm: apical sculpture of weak thread-like radials (Fig. VDD Pycsccccesceceryeecrrceeettceeseaceeseneeseseeeen SOMOS Shell stnaller, mean diameter <3,00 mm; apical sculpture of low, prominent curved radial ridges (Figs 114f, 1200)... 3 5.Umbilicus narrow, V-shaped, mean D/U ratio 4.16; shell colour glossy yellow-harn Lote cau ars bron Parvieostata Umbilicus broader, V-shaped, mean D/U ratio 3.14; shell colour burnished-yellow or DEIR E a issecesssccsanins een saneaeans sort macromphala Coenocharopa sordidus sp. nov. (Figs 111-113; Tables 22, 23) ETYMOLOGY Latin sordidus, dirty: for the dirty shell, COMPARISONS C. sordidus is similar to C. macramphala but is distinguished by its larger size, narrower um- bilicus (D/U ratio 3.56 compared with 3.14 for C. macromphala) and much finer sculpture. C. sordidus (Fig. |11f,g) has apical sculpture of vague radial ridges and crowded spiral cords extending onto post nuclear whorls, C. macro- mphala (Fig. 114 f.g) has prominent curved api- cal radial ribs and more widely spaced spiral cords on apex and post nuclear whorls. Anatomically C. serdidus is distinguished trom C. macromphala by its larger terminal male penitalia which include a long epiphallus and vas deferens, and greatly expanded penis with modified pilaster pattern (Fig. 112a). C. per- vicostata Which is also sympatric with C. sor- didus, has a more rounded, smaller shell, less elevated spire, smaller umbilicus and more prominently sculptured apex. TyPE MATERIAL HOLOTYPE: QMMOQ17276, Upper Brooklield, SEQ (27°30°S. 152°55°E) NVF/Araucaria, litter. Collected hy V. Davies, R. Raven, 8 Apr 198). Height 2.60 mm, MEMOIRS OF THE QUEENSLAND MUSEUM diameter +43 mm, H/D ratio U.SY4, D/U ratio 3.17, whorls 4 12+, ParatyPes: SOUTH-EAST QUEENSLAND - QM- MO1/ 1575, 8 specimens, same data as holotype; Freds Rd. Mt Mee (27°0S°S, 152°43°E) rainforest, leaf litter (3, OQMMO9696, 14 Apr 1980, J. Stanisic. A, Green, N. Hall); Daybara - Mr Byron Rd. at crossing of North Pine River (27°15°S. 152°47°E) leaf hitter (1, OMMO- 17277, 27 Apr 1980, J. Stanisic); Mt Guyra N,P. (25°49°S. 152°35°E) NVF/Arauecarta (1, QMMO- 12073, 7 Sept 1982, AM/QM-ABRS); c | km from Raynbird Creek Rd, on Lacey's Ck Rd. Laceys Creek, open serub, litter (1, QMMO16834, J, Stanisic, G. Annabell, 2 Apr 1984): ¢ 11.3 km NE of Dandabah on Bunya Mis Rd (26°50°S, t51°33°E) (1, OMMO- }2303.. | Sept 1982, AM/OM-ABRS); Upper Brook- field, NVF/Araucaria, bitter (1, QMMO17271, 3 Mar L981. V. Davies, R, Raven; 1. QMMO17268, 11 Dec 1980, V. Davies, R. Raven), OTHER MATERIAL Gold Creek Reserve, Upper Brookfield, SEQ, closed forest, liver (1, QOMMO17278, 16 Dec 1980, V. Davies, R. Raven), Kenilworth S.F., SEQ (26°35°S, 152°42°E) NVF (2, OMMOG152, 18 May 1976, M.J. Bishop). DIAGNOSIS Shell small, often completely covered in dirt, depressed, diameter 3.41—4.43 mm (mean 3.92 mom) with 44 to 4 5/8+ (mean 4 3/8-) normally coiled whorls. Apex and spire moderately elevated, SP/BWW ratio 0,160.30 (mean 0.26), height 2.09-2.64 mm (mean 2.31 mm). H/D ratio 0.55-0.61 (mean 0.59), Protoconch of 1 1/2 to 1 5/8 whorls, mean diameter 709.9 xm at 1 1/2 whorls. Apical sculpture (Fig. 1111) of weak, curved radial ribs becoming more crowded at the nuclear - postnuclear boundary, crossed by very crowded, weakly defined, squiggly spirals. Protoconch often appearing smooth in worn shells. Postnuclear sculpture (Fig. 111g) of strongly bladed, widely spaced, protractively sinuated radial ribs. Microsculp- ture (Fig. 111g) of thread-like radials and incon- spicuous low spiral cords, which buttress both sides of the major radials. Sculpture continuous on the base. Umbilicus (Fig. 111b) narrow, U- shaped, diameter !.01-1.40 mm (mean 1.11 mm), D/U ratio 3,17—3.94 (mean 3.54), Sutures impressed. Whorls shouldered above and slight- ly flattened below the periphery (Fig. 111c). Aperture roundly lunate, margins convergent, Lip simple. Columella slightly dilated. Parietal eallus prominent, uniting lip margins. Colour 173 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE no's (po'e-99'2) O@ZOFZRZ (pS e-1V'e) OO OF LEE (ore-er'e) STOUFSTE PIT (p6'€-65'0) SLIUFLLE LYE OLLVY Vd ira go'U za (60° 1-88°0) £OV0F 660 (ze 0-040) 610 OFRLO (88°0-98'°0) HOU-Z8°0 £0! 1OT rot (ez 1-0) OOLOFEEL OF’ (uu) HALIM Tvoreawn (ceu-s10) 8U0F770 (ge0-61'0) 670 aco 910 (6z°0-S7'0) OzOFLEO iz0 OLLVY MAES “AON ‘dS VLYTV VdONVHIONTOD ONV “AON “dS ROO wL0 Seo (r6'0-S8'0) JF6R'() (p60 18'0) 810UFL8°0 (68°0-LL°0) ERO ze (es -ze1) SOLOFEr TL srl (uur) HLGIM THOHM ACOH trl 970 (oro-eV'0) Le 0F 610 (9z-0-LU0) FIZ or) REO REO (wi) NOISnNuLodd auldsS 650 850 £90 (p9°0-29'0) OLO0FLI0 (e9°0-Z5'0) SIOOFLSO (zs'0-95°0) SOOOFLS'O 190 wo 190 650 aso (o9'0-ss'0) STUOFRS 6s OLY G/H (@ONVY GNV WHS ‘NVAIN) VIVHdWONDVW VdOYVHIONTOD “AON ‘dS SNGIAGHOS (W6z-892) LOVOF6LT (€L'7-Lb2) Lro'0F6SZ (LL'2-€L72) IZ00FSL'Z ROT Ire Zot Que ese (Epr-So'r) OGL OF PTE try (ui) YALAWVIG ipl OL1 (6 1-0LD) EPUUFSL'L (QL I-87 £90096 1 (ag 1-es"1) ZZOUFSSL esl a aN a Lz wz (p9'Z-77'2) OOF ERT 09% (wu) HOH “P/E E H/LE “P/E (B/E POl+8/1 >) PLP (x/L pOe/E €) +RILE (+94) r “B/E +P (48/5 pH/1 F) “Ul +U/L STYOHM ¥dOY VHIONIOI NI NOLLVIVA TWOO7- £7 TTAVL a n a SNAWIOsddS dO aa fWnNn TYONNO Rung, RULE 1961 ONWO. (ad&ioyej4) 6L7L1 ONWO “aS yRpAOy pjojo ndoswwyrouan) Ip7z1 ONWO RTRYL OWWO Pe ng ypel a 9LS11 ONWO (adAjojoH) £9741 ONWO pp i ppoydwossow dosnyr0us0> €L0Z1 OWWO cukn WW £071 ONWO yeqrpurg JO aN L891 ONWO 4Q shaoey] LLZLLOWWO o10gheq SLS11 OWWO (addiopoH) 9LZL1 ONWO. PPY AON snpipsos vdospy20“a0) AWYN 174 Imm ak i Wy Jy: tb ee MEMOIRS OF THE QUEENSLAND MUSEUM Ae ne cere : Sve Qs went Sibeabiatlt De at "tal v eters & SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 175 FIG, 112. Coenacharopa sordidus sp. nov. Upper Brookfield, SEQ. QMMO17268, paratype. a. genitalia and details of penis interior: b, hermaphroditic duct; c. talon and carrefour; d, ovotestis; e, pallial cavity. FIG. 111. Coenocharopa sordidus sp. nov, a—c, Upper Brookfield, SEQ. QMMO17276, holotype; e-g, Fred’s Rd, Mt Mee, SEQ. QMMO9696, paratype: d, h, Upper Brookfield. SEQ, QMMO17271, paratype. a—c, entire shell; d, central and lateral teeth: e, entire shell; f, apical sculpture: g, post nuclear sculpture: h, lateromarginal teeth. Scale lines as marked. 176 25°S 26°S 27°sS aw 151°E 152°E 153°E 154°E FIG. 113. Distribution of Coenocharopa sordidus sp. nov. dirty yellow-horn when clean. Based on 7 measured adults. Vas deferens (Fig. 112a) large, convoluted, giving rise to a more expanded, muscular epiphallus which is as long as the penis. Epiphallus (Fig. 112a) entering penis apically through a spongy pilaster. Penial retractor muscle inserted at the penis-epiphallus junction. Penis (Fig. 112a) with an enlarged upper cham- ber receiving the epiphallus and enveloped in a thin sheath; internally (Fig. 112a) with a very large spongy horseshoe-shaped pilaster opposite the epiphallic pore, and a series of short lon- gitudinal thickenings below. Lower chamber with irregular spongy pustules arranged in vague longitudinal rows. Radula (Fig. 11 1d,h) without unusual features. Based on 3 dissected specimens (QMMO- 17268, QMMO17278, QMMO17271). RANGE AND HABITAT Drier araucarian vine forests between Brook- field, Mt Guyra, and the Bunya Mountains, SEQ. MEMOIRS OF THE QUEENSLAND MUSEUM It has been caught in pitfall traps set for ground dwelling spiders and probably lives among fri- able soil and litter. C. sordidus has been col- lected in degraded forest at Laceys Creek and along the Dayboro-Mt Byron Road, SEQ. These are areas with little rainforest that have been largely cleared for farming. REMARKS C. sordidus is identified by its size and almost completely dirt-encrusted shell (Fig. 111a). Itis inconspicuous in the field. Coenocharopa macromphala sp. nov. (Figs. 114-116; Tables 22, 23) ETYMOLOGY Latin macro-, large; Greek omphalos, um- bilicus; referring to large umbilicus. COMPARISONS C. macromphala may be confused with C. sordidus but the wider umbilicus, smaller shell, more prominent apical sculpture and more wide- ly spaced spiral cords on the postnuclear whorls, distinguish it from that species. C. parvicostata (Fig. 120a—c) has more rounded shell whorls, flatter spire and much smaller umbilicus. Len- webbia protoscrobiculata is sympatric in the Mt Mudlo area, north of Gympie, SEQ. It has apical sculpture of irregular pits and wrinkles, and postnuclear sculpture that includes incised spiral grooves (Fig. 29e,g). Type MATERIAL HOLOTYPE: QMMO17263, Upper Brookfield, SEQ (27°30°S, 152°55°E), litter, NVF/Araucaria. Col- lected by V. Davies, R. Raven, 8 Apr 1981. Height of shell 1.83 mm, diameter 2.98 mm, H/D ratio 0.61, D/U ratio 2.64, whorls 4 3/8-. PARATYPES: SOUTH-EAST QUEENSLAND - QM- MO11576, same data as holotype; Upper Brookfield, CNVF/Araucaria (2, QMMO17262, 3 Mar 1981, V. Davies, R. Raven); beside Bruce Hwy, Blackbutt Range, under logs, NVF/Araucaria (8, QMMO16828, 12 May 1984, J. Stanisic, D. Potter); c. 7.4 km S of Benarkin, Blackbutt Range (26°53°S, 152°11°E) (12, QMMO 12241, AMC136567, 31 Sept 1982, AM/QM- ABRS); Upper Brookfield, NVF/Araucaria, litter (2, QMMO17265, 28 Nov. 1980, V. Davies, R. Raven); Mt Mudlo, Kilkivan S.F., (26°01°S, 152°13"E), FIG. 114. Coenocharopa macromphala sp. nov. a—c, Upper Brookfield, SEQ. QMMO17263, holotype. d—h, same data as holotype. QMMO17265,QMMO11576, paratypes. a—c, entire shell; d, central and lateral teeth; e, entire shell; f, apical sculpture; g, post nuclear sculpture; h, radula. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 177 178 MEMOIRS OF THE QUEENSLAND MUSEUM (? i? 4 ‘yh So CO, 4 es Sf wy xf ‘ eS we & nm a gre oO wenn nee KH Hor Des ¢ [aes a FIG. 115. Coenocharopa macromphala sp. nov. Beside Bruce Hwy, Blackbutt Range, SEQ. QMMO16828, paratype. a, genitalia; b, talon and carrefour; c, ovotestis; d, details of penis interior; e, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 152°E 153°E 154°E FIG. 116. Distribution af Coenocharapa macro- mphala sp. nov, NVFiAraucaria, leaf litter (4, QMMQ15974, 17 Jul 1980, J, Stanisic, A. Green), OTHER MATERIAL Coolabunta Pine Scrubs, Kingaroy, SEQ (1, AMC32999, Jun 1908, S.W, Jackson). DIAGNOSIS Shell small, depressed, diameter 2.47-2.98 mm (mean 2.69 mm) with 3 3/4- to 4 3/8- (mean 4+) loosely coiled whorls, last descending more tapidly. Apex and spire (Fig. | 14c) moderately clevated. SP/BWW ratio 0.15-0.33 (mean 0.25), height 1.28—-1.83 mm (mean 1.57 mm). H/D ratio 0.52-0.64 (mean 0.58). Protaconch of 1 3/8 to | 1/2- whorls. mean diameter 647.8 p.m at 1 1/2 whorls. Apical sculpture (Fig. !11f) of crowded, protractively sinuated radial ribs which become more crowded at the nuclear- postnuclear boundary, crossed by fine spiral cords. Postnuclear (Fig. 1142) sculpture of ir- regularly Spaced. protractively sinuated radial tibs with conspicuous periostracal blades that are often worn, 20-30 ribs on the bedy whorl. Microsculpture (Fig. 114g) of numerous low thread-like curved radial riblets and low crowded spiral cords. Umbilicus (Fig. 114b) 179 broadly V-shaped, diameter 0.70-1.13 mm (mean 0.87 mm), D/U tatio 2.64-3.54 (mean 3.14). Sutures strongly impressed. Whaorls (Fig. 114c) rounded above and below the periphery, with a supra-peripheral sulcus. Aperture round- ly to ovately lunate. Lip simple, columella di- lated and slightly twisted toward the umbilicus. Parietal callus strongly developed and connect- ing lp margins. Colour burnished yellowy- beige, Based on 11 measured adults. Genitalia with large, muscularised epiphallus (Fig. 115a); internally with longitudinal thicken- ings and entering the penis. apically through a simple pore. Epiphallus partially bound to the penis sheath by short fibres. Penis (Fig. 115d) tubular, internally with low pustules in the apical portion giving rise to low, spongy, longitudinal thickenings in the penis proper, Penis sheath present. Penial retractor muscle long, inserting at the epiphallus/penis junction. Vagina mus- cular with low pustules internally, Radula (Fig. 114d.h) similar to that of C. sor- didus. Based on 3 dissected specimens (QMMO- 16878, QMMO17265), RANGE AND HABITAT Prefers drier araucarian notophyll vine forests between Brookfield and Mt Mudlo, SEQ. Like C. sordidus, C. macromphatla has been collected in pitfall traps and probably lives in friable earth and litter. Distribution is poorly known, Coenocharopa alata sp. nov. (Figs 117-119: Tables 22, 23) ETYMOLOGY Latin alatus, a wing: relers to wing-like periostracal processes. COMPARISONS Live specimens are recognised by their large Wing—like periostracal extensions (Fig. 117e,h) Which are deciduous and often lost in dead specimens. Worn shells can be distinguished from C. macromplala by their comparatively higher spire and smaller umbilicus; from C. par- vieostata by their strangly elevated spire and flatter periphery: and from C, multiradiata by their larger size and less regular sculpture, C. alata has a more immediate resemblance to some species of the punctid genus Paralaoma Iredale, 1913, That genus is much smaller and hus apical sculpture of low. rounded spiral cords. 180 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 181 FIG. 118. Coenocharopa alata sp. nov. a,c. 12.2km from Acacia Plateau-Killarney Rd, Koreelah S.F., NSW. AMC128563, paratype. b-e, Coombadjha Walk, Washpool N.P., Gibralter Range, NSW. QMMO17270, paratype. a, genitalia; b, details of penis interior; c, hermaphroditic duct; d, ovotestis; e, pallial cavity. Scale lines as marked. FIG. 117. Coenocharopa alata sp. nov. a—c, c, 12.2km from Acacia Plateau-Killarney Rd, Koreelah S.F., NSW. QMMO17279, holotype: d, same data as holotype. QMMO10961. paratype: e-g, Wiangarie S.F., c. 24km N of Kyogle, NSW. AMC150094, paratype; h, Gibralter Range, NSW. QMMO17270, paratype. a—c, entire shell; d, lateromarginal teeth; e, entire shell; f, apical sculpture; g, post nuclear sculpture; h, details of periostracal process. Scale lines as marked. 182 28°S 29°S 30°S SJeit, > 500m J Sh i51°E 154°E FIG, 119. Distribution of Coenecharopa alata sp. nov, TYPE MATERIAL HOLOTYPE: QMMO17279_ ©, 12.2 km trom Acacia Plateau - Killarney Rd, Koreelah $.F.. NENSW, SU m, SEVT/Araucaria, Collected 15 Mar 1981, AM/OM- ABRS. Heightol shell t.70mm. diameter 2.73 mm, H/D rauia 0.63, D/U ratio 3.32. whorls 4 1/4-. Paratyres: OMMOI0961, AMCI28563, 21 speci- mens, same dala as holotwpe; Binna Burra, Lamington N.P., SEQ. 850m, ONVF (28°13°S. 153° 12°B) (1, OM- MO6210. Apr 1976, M.J, Bishop); Wiangarie S.F..c. 25 km NE of Kyogle. NENSW, 1000 m (28°23°S. 153 Ui" E) litter (6. AMCTSOU94, 17 May 1976, P.H. Colman, |. Loch); Moonpar S.F. NENSW,0.6 km along Mills Rd trom Mounpar Rd intersection. NSW (30°13°5. [52°39"E). under moss on log (1, QMMO17264, 7 Mar 1987. J, Stunisic. D, Potter); Coumbadyha Walk. Wash- pool NLP. Gibralter Range, NSW (29°28°S, 152°16°E) (1. OMMO17270. 8 Mar 1987. J. Stanisic. D. Potter): Gibralter Range, N.P.. NENSW, rainforest (1. OMMO17289,G. Monteith, [0 Nov 1980). Beaury S.F,, e. 15 km W ot Urbenville, NNSW (28°27'S. 152°24E). Sim (7, AMC 154730, 1S May 1976. PH. Colman, | Loch); Red Serub FR Whiag Whinn Soko, NNSW (2N°38°S. ISS |WE) 210m (2. AMCISL791, 15 May 1976, PH. Colman. !. Lowh), OTHER MATERIAL Wesiern end. Gibralter Runge. NENSW (20°32°S,. MEMOIRS OF THE QUEENSLAND MUSEUM 152°17°E) NVF. under moss on rock (1, QMMO17272, 8 Mar 1987, J. Stanisic, D. Potter); top of Condamine River valley, alang Croftby Rd (28°1S°S, 152°29°B) (2, QMMO10541, Mar 1981. AM/OM-ABRS); Gibralter Range. NENSW, rain- forest, litter (1, QMMQO1727S, 10 Nov 1980, R, Raven). DIAGNOSIS Shell minute, glossy, depressed, diameter 2.30-2.73 mm (mean 2.49 mm) with 3 3/4- to 4 1/4- (mean 4-) loosely coiled whorls. Last whorl {in adult specimens) descending much move rapidly. Apex and spire (Fig. 117c) strongly elevated, SP/BWW ratio 0,29-0.45 (mean 0.37), height |.36-1.70 mm (mean 1.49 mm), H/D ratio 0.580.603 (mean 0.60). Protaconch of 1 5/8 to 1 3/4 whorls, mean diameter 629.7 pm at 1 1/2 whorls. Apical sculpture (Fig. 117f) of curved radial ribs, becoming more crowded toward the nuclear-postnuclear boundary, and low spiral cords, Postnuclear sculpture (Figs. 117e,h) con- sisting of widely spaced, prominent, protractive- ly sinuated radial ribs which have large wing-like periostracal processes at the periphery; processes deciduous and not present in larger, worn specimens. Microsculpture (Fig. 117g) of numerous, fine, thread-like radials, crassed by low, crowded spiral cords. Sculpture continuous on the base. Umbilicus (Fig. 11 7b) narrow. U-shaped, diameter 0.41-0.82 mm (mean 0.60 mm), D/U ratio 3.32-5.60 (mean 4.38). Sutures impressed, whorls strongly shouldered above and rounded below a flattened periphery (Fig. 117c). Aperture roundly Junate. Lip simple. columella slightly dilated. Parietal callus strongly developed, connecung lip mar- gins. Colour yellow-horn, Often covered in dirt particles. Based or 3 measured specimens, Genitalia with a short, expanded epiphallus bound to the penis sheath for its entire length (Fig. LtSb). Epiphallus tubular, entering penis apically through a fleshy pilaster, internally with Jongitudinal thickenings. Pental retractor muscle inserted on epiphallus prior to the epiphullus-penis junction. usually originating from the diaphragm but in figured specimen (OMMOl0691). arising from tentacular branch of columellar muscle. Penis (Fig. (1&b) tubular. swollen apically, with a sheath in the expanded upper portion: internally with apical pustulations and basal spongy longitudinal pilasters. Radula (Fig. 117d) typical. Based on four dissected Specimens (QMMO- 183 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE (Z0'p-hL't) 1SOUF88'C S6'£ (or'p—o't) YOUOFRO'P fot tor (RSP 18) Ar Ordre ram (ez"p-LR't) TUOFLO Rot Ob 79'p-LH'E) REUSE (FS pret) ZHOFSTY (cp'b-S0'b) £0 0F6Tr (ogee) ROOFRROE rr OLLVY VG (cs0-69') O10 eeu) (estFir) S10 1S 1s) (rot 150) TLOUFLS ara (og1 ST OFS O oro seu (Ly'0-s¢'u) TROOFES | (r9°0-S5 0) Sur ng Dall (zR'0-9L'0) BOOUF OB (eo 1-8L'0) TEOOFLN'0 RHO (ui) TLLGIN Tvolldnn (coL-96"9) S8VOFSTL (ZS O-RL SI SID UFRED TAY WW/SE1d tr (zp) OO 10 0P Le sar (s1o-1ro) ANOFTI'O +10 (9r'U-x0'0) s1o0szE0 peu (170-910) CO0F6VO sul (970-9 1-0) EVOFITU cra) RIO (ez0-91'0) P0770 (sz'0-91'0) COOFIZO reo (ez'0- EVO) 1007610 (yz0-S-0) Lo-0FHE0 peu OLLVa MAAS (zL0-99°0) (ov'0-B0°0) SLO'OFUL'0 50007600 FL) ovo (rL0-ZL-0) (z1'0-90'0) SOU OFFLO E1060 reo avo 1L0 avo (LV0-e ru) gO ScONFST'O 89'0 20 (izit-ev'a) 18) Scu'0FL EU 180 er L£6°0 oLvo (tR0-ZL'0) (41'0-£1'0) EpOFLLO [z00=S1'0 (ax'U-LL'0) (izo-€r0) ZZ0'0 P80 91U0FRE'O re ocu Gorin (9z0-£1'0) ZLUFS VI zeoarze') (pe I-1Y) (ag0-L1-0) SIOUFSE'T SLOOF EZ 71 oro (uuu) (uw) TLLGIAN NOISNY.LOUd THOHM AGOU aulds (ADNV¥ CNY Was ‘NVaW) (290-0) BLOOFRSO ssu (Ls tr es'0) OLOOFSS'O aU) 190 (LoUre st) 20'0FSs'0 ns ist “0 ooo (6s0-s8'0) zorLs 0 (o90-€5'0) Lov0FLS'O ust (19°0-05'0) 10'0F9S'0 (95°0-8F'0) 1007 "S'0 6s'U OLLVa C/H (107-16) 1ZU'0F 961 Wz (prz-se'l) OPN OFINE LIZ (gp't-972) QROOF PET Elz (OE 7-217) Cpu'UFOE'Z (6€°2-60'2) OPLOFHTZ (is z-e¢'2) 9100F RET Be (sp'g-al'e) PZ UF EEE (ER€-47'£) 180°0F 9F'E SLE (wu) WaLAWVIC (61 I-Lt SeO'UFEL Lut (61 1-60'1) TRU FEL aL ayt (cere) SCUFRZ"A Re’ bbl (e147 EPO UF RET (6p I-22 1) TeoOF LP’ gs't (961-991) PZU'0F SRL (po'z-0L 1) LEOOFLE'L we (witu) L1o1gH (a/t t-H/L *$ WILE (+8/1 pOr-B/L ED + WEF “RLY CH/L port) +h “BILE (g/t b-A/L £) +h PEE +H/EP “BILE 8/1 PO-R/L £) + (3/S pOrR/L P) RIED (+8/€ t-8/1 ») +H/1 rier STHOUM “NON AS VILVIOVHLLIOW.: VdOUVHOINIOD UNV “AON “AS VIVLSODIANVd VelOUVHIONTO.D “AON “dS SISNTHVAVSSIA VdOUVHIONTOO NI NOLLVINVA TVOO1- Pz TEV r £6RU1L OWWO “A'S auadeyid VW. ! RELEST OWYV ha RS6TRIOWV SOABD IEE (adAqojoH) I £2681 ONWO HEA, 8, UOS| AL, piniposnjna ndosmys0uae) t Z8L01 OWWO “PN OFLOG +7801 ONWO ‘a's Addeyorg a 1 9r691 OWAG auadryid ‘TWA 9f601 OWND WOO{O0 J, 1 L809 ONWO, yeqepurq I L9TLLOWWO quomi vay t ssezt ONWO s #S¢8 OWWO OPW IW (adMojoy) 1 99741 ONWO. 23) 1A eimsoanaed pdosnysqueay rai OR9IZLOWY L £1021 ONWO (add10j0H) i 19741 ONWO SOAR) HEQESSIA sisuaynqussak pdosnysour02 SNAWIDadS AO YaNWON FINN 184 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 185 FIG. 121. Coenocharopa parvicostata sp. nov. Kenilworth §.F., SEQ. QMMO17264, paratype. a, genitalia; b, details of penis interior; c, talon; d, ovotestis; e, hermaphroditic duct; f, anterior edge of pallial cavity. Scale lines as marked. FIG, 120. Coenocharopa parvicostata sp. nov. a—c, Freds Rd, Mt Mee, SEQ. QMMO17266, holotype; d-f, Mt Mudlo, Kilkivan $.F., SEQ, QMMO8358, paratype: g, Diehappy S.F., NSW. AMC128388, paratype; h, c. 8.3km from Woodenbong-Legume Rd, Tooloom Scrub, Beaury S.F., NSW. AMC128532, paratype. a-c, entire shell; d, post nuclear sculpture; e, spire; {-h, apical sculpture. Scale lines as marked. INO 10270, OMMOLO269, QMMO17289, AMC- 128563). RANGE AND HABITAT Cooler subtropical notophyll vine forests of the Great Dividing Range between Darrigo, NSW and the QLD/NSW border, and the notophyll vine forests (and possibly microphyll fern forests) of the McPherson Ranges, SEQ. Live specimens have been collected from among moss on logs and rocks. REMARKS Probably Setomedea’ aculeata (Hedley 1899) comes closest in shell form, However, the periostracal extensions of \S". aculeata are more slender, more numerous and regularly spaced along the major radial ribs, Coenocharopa parvicostata sp. nov. (Figs 120-122; Tables 22. 24) EFyMOLOGY Latin parvus, small; and cos/atus rib; refers to the postnuclear sculpture. COMPARISONS C, parvicostatais sympatric with C, sordidus, C. macromphala and C. multiradiata, and dead shells of these species are apt to be confused. However C. parvicostata can be distinguished from C. sordtdus by its smaller size, more rounded whorls, less elevated spire and more prominently sculptured apex: from C. macro- miphala by the lack of a supraperipheral suleus, Jess elevaied spire and smaller umbilicus; from C. multiradiata by the more numerous apical radial ribs which are crowded at the nuclear- postnuclear boundary; and from C. alala, by having a less clevated spire and more rounded whorls. Tyre MATERIAL HoLoTyPE: OMMO17266, Fred's Road, Mt Mee, SEQ, 2705'S. (52°43'E. rainforest. liner. Collected hy J. Stanisic, N. Hall, A. Green, 14 Apr L980, Height of shell 1.58 mm. diameter 2.81 mm, H/D ratio 0.56. D/U ratio 3,34, whorls 4 3/8. PARATYPES: Mt Mudlo, Kilkivan S.F.. SEQ (2h 01'S, (52°13 E), NVE/Aradearia, leaf litter (9, OMMOS358, 17 Jul 1980, J. Stanisic, A. Green): Kenilworth 8.F,, SEQ (26°37°S. 152°42"E) rainforest. Teal Vitter (1, OMMO1 7267. 22 May 1980. J. Stamste. A. Green): ¢. 8.3 km trom Woodenbong - Legume Road, Togloom Serub., Beaury S.F.. NENSW MEMOIRS OF THE QUEENSLAND MUSEUM (28°29°S, 132°24'E) CNVF (7, AMCI28532, OMMO10936, [5 Mar 1981, AM/OM-ABRS); Dan- dabah, Bunya Mts N.P., SEQ (26°53'S, 151°35°E) (2, QMMO6087, 5 Mar 1976, M_J. Bishop): c. 6,5 km E of tickgate. Mt Pikapene S.F.. NENSW, (2901'S, 152°45°E) rainforest (1, QMMO16946, Mar 1981, AM/QM-ABRS): creek culling, Diehappy S.F.. NENSW (30°28°S, 152°38°E), broad leaf scrub (7, OMMO10824. AMCII8388, Mar L981 AM/QM- ABRS); Newell Falls, Dorrigo N.P,, NENSW (30°24°S, 152°45°E) NVE (6. OMMO1L0752, AMC- 128333, Mar 198), AM/QOM-ABRS); Mudlo Gap S.F.. SEO (26°01'S, IS214°E) MVF/Araucaria (5, OMMO012355, AMC136656-_2 Sept 1982, AM/QM- ABRS): ¢.0.5 km S of ‘Bellbird’ parking area, Kenil- worth S.F,. SEQ, rainforest litter (1, OMMO17264, 28 Feb J9S80, J. Stanisic, A. Green). OTHER MATERIAL Little Yabba Creek. Imbil S.F., SEQ (26°287S, (52°38'E) NVF (5, OMMO12026, AMC136440, § Sept 1982, AM/QM-ABRS): Bellinger River, NENSW (30°27'S, 152°37°E) rainforest gully (3, OMMO10832. AMC128399, Mar 1981, AM/QM- ABRS): Mt Warning N.P.. NENSW_ near base (28°24'S, 153° 16°E) NVF/Palms (4. OMMO10480, AMC 129285, Mar (981, AM/QM-ABRS); side track aff Mt Archer Rd. Mt Mee S.F., SEQ (27°047S, IS241°E), NVFideauceria (1, QMMQ15S079, J. Stanisic, D. Potter); Manorina N.P., Mt Nebo, SEQ (2729'S, 15247 E) wet scleraphyll (1, OMMO0258, Aug 1976, M.J, Bishop); Wralten’s Camp, Wratlens S.F., SEO (26°17°S. 152°20°E) NVF, litter (3, OM- MOQO11563. 17 Jul 19S80_J. Stanisic, A. Green); Cherty Tree North S.F., ¢, 40 km W of Casino, NSW 28°54°S, [52°45°E), 400 m (100 +, AMC154733, 19 May 1976. PW, Colman, J. Lach); Ree Serub F.R., NENSW (28°38°S, 153°20°E), rainforest (1. AMC|28867_ 20 Mar 1981, AM/QM-ABRS): Mt Lindesay, Upper Richmond Ry, NSW (3, AMC 154734. 27 Apr 1948. L. Price); Koreelah Ck, Beuury S.F.. c 1S km W of Urbenville, NSW (2873108, 152°20°E) (52. AMC 154736, 18 May 1976, PH. Colman. |, Loeb), DIAGNOSIS Shell small, depressed. diameter 2.09-2.81 mm (mean 2.36 mi) with 3 3/4 to 4 1/4 (mean 4+) whorls, last whorl descending, Spire (Fig. 120c) moderately elevated to depressed, SP) BWW ratto 0.164032 (mean 0.21). Height 1.07-1.58 mm (mean |.24 mm), H/D ratio 0.50- 0.61 (mean 0.57). Protoeonch of 1 1/4 ta | S/8 whorls. mean diameter 364.5 jm at 1 1/2 whorls. Apical sculpture (Fig. 120f-h) of fine, protrac- SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 26°S 27°S 28°S 29°S 30°S 31°S 151°E 152°E 153°E 154°E FIG, 122, Distribution of Coenocharopa parvicostata sp. nov. tively sinuated radial ribs, more crowded near the nuclear-postnuclear boundary, crossed by irregular spiral cords. Postnuclear sculpture (Fig. 120d) of regularly spaced (often obsolete) blade-like, protractively sinuated radial ribs, continuous on the base. Microsculpture (Fig. 120d) of numerous, thread-like radial riblets and low, broad crowded spiral cords. Umbilicus wide, V-shaped, diameter 0.45-0.84 mm (mean 0.57 mm). D/U ratio 3.344.62 (mean 4.16). Sutures impressed; whorls rounded below and slightly shouldered above the periphery (Fig. 120c). Aperture roundly to ovately lunate. Lip simple, columella dilated and twisted toward the umbilicus. Parietal callus present. Colour light- yellow horn with a pink tinge in some specimens. Genitalia with epiphallus (Fig. 121a) tubular, 187 internally with longitudinaly thickenings, enter- ing the penis apically through a simple pore surrounded by a muscular collar (Fig. 121b). Penial retractor muscle inserting on the epiphal- lus prior to its entry into the penis chamber. Penis (Fig. 121b) expanded, tubular, internally with crowded pustulations and weak lon- gitudinal pilasters. Vagina short, expanded, in- ternally with spongy pustulations. No data available on radula. Based on one dissected specimen (QMMO- 17264). RANGE AND HABITAT From the Dorrigo area, northern NSW to Mt Mudlo, north of Gympie, SEQ. It has the largest distribution of any Coenocharopa species and is found in both warm moist notophyll vine forest and drier araucarian notophyll vine forest. Al- though there is some range overlap with C. alata, C. parvicostata has only been found sympatri- cally with that species in the Beaury State Forest, northern NSW. No data available on micro- habitat. REMARKS In spite of the comparatively large number of dead shells available for study only one sub- adult specimen was suitable for dissection and more material is needed to confirm structures of the terminal male genitalia. As may be expected in a wide ranging species there is a degree of interpopulational variation in shell characters. Southern specimens are larger and have bolder sculpture on the protoconch. Coenocharopa yessabahensis sp. nov. (Figs 123-125; Tables 22, 24) ETYMOLOGY For the type locality, Yessabah. COMPARISONS Its relatively large size and postnuclear sculp- ture of numerous low, crowded, radial ribs dis- tinguish C. yessabahensis from congeners. C. sordidus is most similar but has widely spaced radial ribs with conspicuous periostracal blades. Anatomically the penial pilaster pattern of apical stimulatory pad and large basal U-shaped thick- ening contrasts with the typical pattern of simple longitudinal thickenings. C. multiradiata which is possibly sympatric with C. yessabahensis, has regular widely spaced radial ribs, more con- spicuous microsculpture and is much smaller. MEMOIRS OF THE QUEENSLAND MUSEUM 188 mm 1 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 189 FIG. 124, Coenocharepa yessabahensis sp. nov. Yessabah Caves, via Kempsey, NSW. QMMO17013, paratype. a, genitalia; b, details of penis interior; c, ovotestis; d, hermaphroditic duct and talon; e, pallial cavity. Scale lines as marked. FIG. 123. Coenocharopa yessabahensis sp. nov., a—c, Yessabah Caves, via Kempsey, NSW. QMMO17261, holotype; d—h, same data as holotype. QMMO17013, AMC121680, paratypes. a—c, entire shell; d, marginal teeth; e, spire; f, apical sculpture: g, post nuclear sculpture; h, central and lateral teeth. Scale lines as marked. 190 30°30'S 31°S 31°30'S 153°E FIG. 125. Distribution of Coenocharopa yessabahen- sis Sp. nov. TYPE MATERIAL HOLOTYPE: QMMO17261, Yessabah Caves, via Kempsey, NENSW (31°05°30"S, 152°51°E), vine thicket on limestone rocks. Collected by J. Stanisic, D. Potter, P.H. Colman, 3 March 1987. Height of shell 2.22 mm, diameter 3.75 mm, H/D ratio 0,59, D/U ratio 4.24, whorls 4 3/4. PARATYPES: 130 live, and 20 dead adults and sub- adults, QMMO17013, same data as holotype; Yes- sabah limestone cave, Sherwood Quarry, W of Kempsey, NSW (31°06’S, 152°42’E), ground litter in and out of cave (63, AMC121680, 9 Oct 1979, P.H. Colman, J. Stanisic), DIAGNOSIS Shell small, depressed, diameter 3.19-3.83 mm (mean 3.40 mm), with 4 1/8 to 4 3/4 (mean 4 3/8) normally coiled whorls. Last whorl des- cending slightly in front. Apex and spire (Fig. 123c) only slightly elevated, SP/BWW ratio 0.11-0.26 (mean 0.20). Height 1.66-2.22 mm (mean 1.88 mm). H/D ratio 0.48-0.61 (mean 0.55). Protoconch shiny with 1 1/4 to 1 5/8 whorls, mean diameter 644.5 wmat | 1/2 whorls. Apical sculpture (Fig. 123f) of weak, squiggly, irregularly spaced spiral cords and vague, low radial ribs. Postnuclear sculpture (Fig. 123g) of low, broad, crowded spiral cords, crossed by weak thread-like radial and microradial ribs, continuous on the base. Umbilicus (Fig. 123b) narrow, U-shaped, diameter 0.76-1.03 mm (mean 0.83 mm). D/U ratio 3.73-4.43 (mean MEMOIRS OF THE QUEENSLAND MUSEUM 4.12). Sutures strongly impressed; whorls shouldered above and rounded below the periphery (Fig. 123c). Aperture roundly lunate. Lip simple, slightly thickened at the columellar margin. Columella dilated and slightly reflected over the umbilical opening. Parietal callus developed. Colour yellowy-brown. Based on 20 measured adults. Body colour dark grey, with ommatophores and pedal glands darker. Terminal male genitalia with epiphallus (Fig. 124a) reflexed, entering the penis apically through two spongy thickenings. Penis (Fig. 124b) thick, muscular with a thin sheath for its entire length; internally with a spongy horseshoe-shaped pilaster (Fig. 124b) in lower part of the penial chamber, and a small subcircular thickening in the apical half, other- wise with pustules. Penial retractor muscle in- serted on the epiphallus prior to its junction with the penis. Free oviduct short, muscular, internal- ly with rather vague, transverse, spongy thicken- ings. Vagina short, thin with internal longitudinal pilasters. Radula (Fig. 123d,h) with mesocone of the central and inner laterals greatly enlarged. Based on four dissected adults (QMMO- 17013). RANGE AND HABITAT Only on the limestone outcrop at Yessabah, near Kempsey, NSW, which supports a low vine thicket. Crawling on limestone rocks. REMARKS C. yessabahensis, Letomola contortus and Rhophodon kempseyensis are restricted to the limestone outcrops between Yessabah and Car- rai, near Kempsey, NSW suggesting that these outcrops are important refugia for terrestrial snails. Coenocharopa multiradiata sp. nov. (Figs 126-127; Tables 22, 24) ETYMOLOGY Latin multi -, many; and radiata radial; refer- ting to the numerous radial ribs. COMPARISONS Sculpturally, C. multiradiata is the most dis- tinctive species of Coenocharopa described herein. The sculpture is regular and in many ways simulates the more typical reticulate pat- tern seen in Gyrocochlea and Rhophodon. Com- pared with its congeners, C. multiradiata has SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 191 Imm FIG. 126. Coenocharopa multiradiata sp. nov. a-c, Top of Condamine River valley on W slopes of Wilson’s Peak along Croftby Rd, NSW. QMMO18973, holotype; d-f, Natural Arch, Carrai Caves, NSW, AMC 142958, paratype. a-d, entire shell; e, apical sculpture; f, post nuclear sculpture. Scale lines as marked. \) 20°S. 41°5 FIG. 127. Distribution of Coenecharopa mulliradiata Sp. nov. more numerous, Iess protractively sinuated radial ribs and microradials which have con- spicuous periosiracal blades. In addition, the microspirals are less crowded, more slender and raised to produce a conspicuous reticulate pat- tern. C, alata is larger, has fewer radial ribs with extremely expanded periostracal processes and has amore reduced microsculpture. C. parvicas- tata Which is most apt to be contused with C. multiradiata, is larger with more elevated spire und has fewer, less regularly spaced, more protractively sinuated radial ribs. In subadult specimens, these two species can be differen- tiated by their apical sculpture which consists of few. widely spaced, well defined, radial ribs in C. multiradiata, and crowded, low ridges, be- coming more crowded al the nuclear- postnuclear boundary, in C. parvicastata. C. vessahahensis is larger with very crowded, weak, radial ribs and reduced microsculplure. TYPE MATERIAL HOLOTYPE; QMMO18973, top of Condamine River MEMOIRS OF THE QUEENSLAND MUSEUM valley on W slopes of Wilson's Peak, along Crofiby Road, SEQ (28°146'S, 132°28'E). rainforest/hoop pine. Collected AM/QM-ABRS, 16 Mar 1981. Height of shell 117 mm, diameter 2.16mm, H/D ratio 0.54, D/U ratio 3,93, whorls 4 1/8. PARATYPES: AMC1 28634. same collection dita as holotype: Natural Arch, Carrai Caves, near Kempsey, NSW, leaf lilter (9, AMC 142958, 22 Feb 1980, P.H. Calman): ¢. 6.5 km E of Tickgate, Mr Pikapene S.P.. NSW (2801'S. 13248 EB). riverine rainforest (11. QMMOLO893, 14 Mar ]98), AM/OM-ABRS), OTHER MATFRIAT Limestone caves al Natural Arch, W of Kempsey, NSW (30°59°S, [52°21°E). under slubs of limestone (7, AMC143728, 10 Mar 1981. W, Ponder, OL Gril- fiths): Koreelah S.B.. NSW. c. 12.2 km from Acacia Platedu-Killarnevy Rad. (28°20°S, |52°24°E) SEVT/MVF (1, AMC 128553. 15 Mar 1951, AM/QM- ABRS). DIAGNOSIS Shell diameter |.91-2.16 mm (mean 2,03 mm), with 3 7/8- to 4 1/S+ (mear 4-) normally coiled whorls. last whorl descending slightly, Apex and spire (Fig. 126c), very lightly elevated, SP/BWW ratio 0.08-0.16 (mean 0.13), height of shell 1.07-1.19 mm (mean 1.14 mm), H/D ratio 0,43-0.62 (mean 0.54). Protoconch of 1 1/2 to 1 5/8 whorls, mean diameter 511.1 jum at | 1/2 whorls. Apical sculpture (Fig, 126e) consisting of widely spaced, conspicuous, slight- ly curved radial ribs with raised periostracal blades and numerous, narrow, low spiral cords. Post nuclear sculpture (Fig. 126f) of numerous, regularly spaced, weakly protractively sinuated tadial ribs which have prominent periostracal blades that are often worn, 38-47 (mean 42,5) ribs on the body whorl. Ribs/mm 5.78-6.96 (mean 6.63). Microsculpture (Fig. 126f) of fine thread-like radial riblets, 6-13 between each pair of major ribs, and fine, slender conspicuous spiral cords which are not raised at their junction with the microradials. Sculpture continuous on the base. Umbilicus (Fig. 126b) narrow, U- shaped, diameter 0.47-0.55 mm (mean 0.51 mm). D/U ratio 3.79-4.36 (mean 3.98). Sutures impressed, whorls shouldered above and rounded below a slightly flattened periphery (Fig. 126c). Aperture roundly lunate. Lip simple, columella not expanded, Panetal callus developed, Colour light-horn. Based on 10 measured adults. Anatomy unknown, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE RANGE AND HABITAT From the top of the Richmond Range, NSW and from the Carrai limestone outcrop near Kempsey, NSW; in dry cool subtropical vine forest and warm temperate rainforest. It is prob- able that C, multiradiata extends further south in warm temperate forest. No data are available on microhabitat. REMARKS C. mulriradiata differs from average Coenocharapa patterns in a number of features. Nonetheless these altered character states are teadily derived from typical Coenocharopa. In the absence of anatomical detail, placement in this genus. is preferable to creating another nomenclatural entity based on shell characters. Egilomen Iredale. 1937 Egilomen Iredale, 1937a, p. 326; Iredale, 19414. pp. 267, 269; Kershaw, 1955, p. 29; Burch, 1976, p. 132, ‘TYPE SPECIES Heltx cachlidium Cox 1868; by original desig- nation. PREVIOUS STUDIES Fgilamen was introduced for five species with bold radial ribs and wide umbilicus. Egilomen is herein redefined to include species that haye apical sculpture of prominent radial ribs with spiral elements absent or reduced, postnuclear sculpture of bold radial ribs. and a microsculp- lure of low rounded spiral cords, ‘Helix’ cochlidium, and a new species from northern NSW and SEQ are assigned. Other species pre- viously allocated to Egilomen probably belong elsewhere. DIAGNOSIS Shell diameter 1.62-3.07 mm, with 4 1/4 to 5 1/8 tightly coiled whorls, last descending rapid- ly. Apex and spire weakly to strongly (g/obosa) elevated. Apical sculpture of broad, bald, wide- ly spaced. weakly curved radial ribs, and weak spiral cords, Postnuclear sculpture of broad. bold, moderately to widely spaced (cochlidium), weakly protractively sinuated to vertical radial ribs. Microsculpture of low, rounded, crowded spiral cords crossed by fine, thread-like radial riblets, Umbilicus wide, U-shaped (cochlidium) to closed (g/ohosa). Sutures strongly impressed. Whorls shouldered above and rounded below a weakly compressed periphery. Lip thickened, rarely with a distinct basal knob (some popula- tions of elobosa), retracted at the suture to form an apertural sinus. Animal with foot shart and broad; colour white without any accessory pigmentation. Palljal cavity with complete secondary ureter and strongly bilobed kidney. Talon with very short stalk, subcircular. Terminal male genitalia with well-differentiated epiphallus and simple epiphallic pore. Penial retractor muscle insert- ing on epiphallus. Penis subcylindrical, inter- nally with a single, large longitudinal pilaster and an accessory circular pad, Female genitalia with reJatively long vagina and free oviduct. Radula with central tooth much smaller than laterals, otherwise without unusual features, COMPARISONS Conchologically Egtlomen resembles Cralopa and Rhophodon, Rhephodon differs most markedly in having apertural barriers, Micros- culpturally Egilomen (Figs 128g. 131f) has prominent, rounded, continuous spiral cords crassed by fine, thread-like radials in contrast to Rhophoden, which has low, barely visible spiral cords and prominent, high microradial mbs with elongate beads at their intersection (Figs Slt, 87e). Anatomically the strongly bilobed kidney of Egt/lomen (Figs 129d, 132f) contrasts with the weakly bilobed. to almost unilobed, condition of Rhephodon (Figs 72f, 75h). Terminal male genitalia of Egi/omen have the penial retractor muscle inserted on the epiphallus (Figs 129b. 132b) whereas in Rhophadon it is always in- serted al the penis-epiphallus junction (Figs 72a, 75e), Cralopa differs in having more prominent api- cal spiral cords, more raised microsculpture and less differentiated epiphallus. Egilomen cochlidium (Cox, 1868) (Figs 125-130; Tables 22, 25) Helix cachlidiim Cox. (R68, p. 13. pl. 8, fig. 1. Egilomen cocllidiwn (Cox). Wedale, 19374, p. 328, Tredale. 1941a. p. 269. COMPARISONS E. cachlidtum differs from E, globosa in its flattened shell and wide umbilicus. It is most similar to species of Rhaphodon but lacks aper- tural barriers and has less crowded apical sculp- ture. Discocharopa aperta (Fig. 93) has a wide umbilicus aad prominent radial sculpture but is 194 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 195 FIG. 129. Egilomen cochlidium (Cox, 1868). a, c,d, Upper Pine Ck, Canungra, SEQ. QMMO16905. b, Mills Rd, Moonpar S.F., NSW. QMMO1I7290. a, genitalia: b, details of penis interior: c. talon and carrefour; d, pallial cavity. Scale lines as marked. FIG, 128. Egilomen cochlidium (Cox, 1868). a-c, Clarence River, NSW, AMC63505. lectotype: d, h. Mills Rd, Moonpar S.F., NSW. QMMO17290; e-g, The Island, off Grafton, NSW. AMCS57247. a—c, entire shell; d, marginal teeth; e, entire shell; f, apical sculpture: g, post nuclear sculpture; h, central and lateral teeth, Scale lines as marked, 196 MEMOIRS OF THE QUEENSLAND MUSEUM much smaller, lacks regular spiral sculpture and TAULE 25- LOCAL VARIATION IN AGILOMEN COC HLM (COX, lak) ANB RGILOMEN GL UHASA SE NOV = es has finer, more crowded radial ribs. = SS, ee oF = ! A — Tyre MATERIAL Lecrotyre:; AMC63505, Clarence River, NSW, 23 Height of shell 1.25 mm, diameter 2,26 mm. H/D ratio wee ekd Eo = 0.55. D/U ratio 3.14, whorls 4 1/2. oT See! So PARALECTOTYPES: AMC153714, 2 specimens, same collection data as holotype. ae £2 z wen 22 = ¢ gee 2 9 OTHER MATERIAL 2 pte ME 3 6 ero of Dorrigo, NSW (1, AMC63790); The Island, off Graf- ton,NSW (12, AMC57247); O’Reillys, Lamington A r N.P., SEQ (28°14'S, 153°15°E), MFF (1, fn te ie AMC129243, 17 Mar 1981, AM/QM - ABRS); W = 2 45 ag a & = 32 2 ¢& — side Richmond Range, NSW, tributary of Duck Ck (28°33’S, 152°40’E) (2, AMC128524, 14.Mar 1981, on . Heme “es AM/QM - ABRS): top of Condamine River Valley, zs 22 ; zoe W slope of Wilson’s Peak, NSW (28°16’S, 152°28’E) 3 = 282 22 € & 2 22 28 8 (1, AMC128637, 16 Mar 1981, AM/OM - ABRS); Upper Pine Creek, Canungra, SEQ, NVF/Araucaria, 2 i. om he under rock (1. QMMO16905, 21 Jan 1987, J. Stanisic, Bs ‘ == 32 25 D, Potter); Mills Rd. Moonpar §.F., NSW (30°13’S, raf o£ £2 = ¢ 228 #2 $ 152°39°E) rainforest (1,QMMO17290, 7 Mar 1987, = a 7 J. Stanisic, D. Potter); Carr's Creek, Clarence River, = NSW (1, AMC8737, presented C. Hedley, ex Brazier, 3 54 26 «485 Rossiler, Cox: 1, AMC152253, ex Brazier); Cherry 3 ses $2 x = 3 #E 2S = Tree North S.F, NSW (28°54’S, 152°45’E) litter (5, 3 J FS F_*= = FS SS = AMC153714, AMC154732. 19 May 1976, P.H. Col- 7 man, I, Loch); Goomburra S,F., SEQ (27°59’S, s . as ee. as 152°21"E) (1, AMC128618, 16 Mar 1981 AM/QM - ~ & 33 . =3 37 ABRS); N of Dorrigo, Clouds Ck S.F., NSW a 428 82 3 Efe G5 = (30°07°S. 152°41°E) dense rainforest (1, AMC- fe 128319, 11 Mar 1981, AM/QM - ABRS); ¢. 5.5 km N . re “=, 2. of Dorrigo NSW (30°23'S, 152°44"E) (1, QMMO- a =F Ee 52 10794, Mar 1981, AM/QM - ABRS); Cunninghams zE $22 42 § $ 8 32 §2 = Gap, SEQ, leaf litter (1,QMMO17291. 18 Jun 1979, ? 7 sal pal G. Annabell); Koreelah Ck, Beauty S.F., NSW (28"- 21°S, 152°20°E) (3, AMC154735, 18 May 1976, P.H. =e 5a 33 ag Colman, I. Loch). 2- 925 Fe 8 = Bes 98 8 = Vm bee de | owe re are?” BMGHOSE Shell diameter 1.91—3.07 mm (mean 2.35 mm) 4 2 z= 2 with 4- to 5 1/8- (mean 4 1/2+) lightly coiled & axe AD 2 | = ad ¢2 « whorls, the last descending rapidly in front. 2 S55 35 % ~ 553 32 5 Apex and spire (Fig. 128c) flat to slightly 4 elevated, SP/BWW ratio 0.09-0,27 (mean 0.18), 32 height 0,95-1.83 mm (mean 1.30 mm), H/D #0 ToT patio 0.49-0.61 (mean 0.55), Protoconch of 1 ze 1/2 to 1 5/8 whorls, mean diameter 491.8 jxm at 5 a 1 1/2 whorls. Apical sculpture (Fig. 128f) of Zeogt, » if 38 3 RE? widely spaced, slightly curved, broad, prominent . gegehe & 22 €& 2525 25 radial ribs, and low, inconspicuous spiral cords. 2 #82ise 22 FS EE 2322 22 2 Postnuclear sculpture (Fig. 128g) of broad, * ae eee SS se FF widely spaced, protractively sinuated radial ribs, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 28°S 29°S 30°S FIG. 130. Distribution of Fgilomen cachlidium (Cox, 1868), 30-47 (mean 36.8) ribs on the body whorl. Ribs/mm 4.22-5,99 (mean 5.07), Microsculp- ture (Fig. 128g) of fine radial riblets, 8-12 be- tween cach pair of major ribs, and fine crowded spirals. Sculpture continuous on the base. Um- bilicus (Fig. 12sb) wide to very wide, U-shaped, diameter 0.60-1.03 mm(mean 0.82 mm), D/U ralio 2.59-3.47 (mean 2.93). Sutures impressed. Whorls flattened below and rounded above a laterally compressed periphery (Fig. 128c). Aper- lure ovately lunate. Lip simple, thick, margins convergent. Columella only slightly dilated. Parietal callus well developed. Colour white to light straw-yellow, often covered in dirt particles. Based on 10 measured adults. Vas deferens a thin tube. Epiphallus (Fig. 129b), cylindrical, greatly expanded, reflexing prior to entering penis apically through a simple pore surrounded by two fleshy lips. Penis (Fig. 129b) short, internally with a large, apical, lon- gitudinal pilaster, and a basal, circular pilaster. Penial retractor muscle (Fig. 129b) inserted on the epiphallus prior to its junction with the penis, Free oviduct short, muscular, internally with lon- gitudinal pilasters. Radula (Fig. |28d,h) with central tooth greatly reduced, Based on two dissected specimens (QMMOQ- 16905, QMMO17290), RANGE AND HABITAT Mainly confined ta cooler subtropical notophyll vine forests on highlands of the Great Dividing Range (between Dorrigo, NSW and the Mistake Mountains, SEQ), and the McPherson Ranges on the NSW/OLD border. The early Grafton/Clarence River records need to be con- firmed. The only data available on microhabitat are from a specimen collected from soil under j1 rock at Canungra, SEQ, REMARKS The apical Jongitudinal penial pilaster and basal penial stimulator contrast with the apical stimulator and basal longitudinal pilaster of E globasa. This difference in penial surface sculp- ture is probably a species recognition change brought about by sympatric interaction. &. cochlidium ts poorly represented in collections. Egilomen globosa sp. nav. (Figs 131-133; Tables 22, 25) ETYMOLOGY Latin globosa, rounded; referring to shell shape. COMPARISONS F. glabosa differs from F. coelhlidium in having a more globose shell with strongly elevated spire.more crowded radial ribs, more prominent spiral cords and closed umbilicus. £, globosa is most apt to be confused with the sympatric Cralopa stroudensis (Fig. 95) which has similar shell shape, prominent radial ribbing and closed umbilicus, However, C, stroudensis has fewer whorls, 4 more conspicuous apertural sinus, less elevated spire, thinner lip. and a greater number of ribs on the body whorl. El- sothera genithecata (Fig. 108) and £. nautilodea (Fig. 106), from northern NSW, have closed umbilici and prominent radial ribs, but are much larger, with flatter spires, and comparatively less crowded radial ribs.on the body whorl. TYPE MATERIAL HOLOTYPE: QMMOQ17292. sidetrack off Mt Archer Rd. Mt Mee §.F., SEO 27°04'S, 152°41°E. Collected 31 Sept 1982, AM/QM-ABRS. Height of shell 1,87 mm, diameter 2.47 mm. H/D ratio 0.76, wherls.4 3/4. PARATYPFS: QMMOQO12175, AMC136521, 20 specimens, same collection data as holotype; Mt Cura N.P., SEQ (25°49°S, 152°35°E) MVF/NVF/ Araucaria (9, OMMQO12068 AMC136463, 7 Sept 1982. 4M/QM - ABRS): near base, Mt Warning N.P.. NSW (28°24’S. 153°16'E) NVF/Palms (2, OMMO- 10483, Mar 1981, AM/QM - ABRS); Mt Warning. N.P., NSW (28°24°S. 153°17°E) (3, AMC129305, 19 Mar 1981, AM/QM - ABRS); Natural Bridge N.P.. SEQ (28°13'S, 153°14E) NVF (8, QMMO10459, AMC129270, 18 Mar 1981. AM/QM - ABRS); Terania Creek, Nightcap Range N.P., NSW (28°34'S, 153°19°E), subtropical rainforest, under bark on log (15. OMMOI7O12, 11 Mar 1987, J. Stanisic, D. Pot- ter: 13. OMMOITLA8. 11 Apr 1987. J. Stanisic. J, Chaseling). OTHER MATERIAL Toonumbar Forest Read, Toonumbar 8.F.. NSW (28°33°S, 15245’ FE) CNVPF(),QMMO10911, 14 Mar 1981, AM/QM - ABRS); Rainforest Park, near Maleny, on Maleny-Montville Rd, SEQ, NVF (1, OMMO11882, 5 Dee 1982, J. Stanisic): sidetrack off Mt Archer Rd, Mt Mee S.F. SEQ (27°(4'S. 152°41°E) (2. QMMO15075. OMMO15076, 6 Sept 1983, J. Sianisic, D, Potter; 2. QMMQ12177, AMC136520, 30 Sept 1982, AM/QM - ABRS); Fred's Rd. Mr Mee, SEO (27°0S°S, 152°43°E) NVF, litter (1, OMMO- 15983, 28 Oct 1984. J. Stanisic, D, Powter): c. 1.6 km N af Numinbah Valley tirnoff on Burleigh- Springbrook Rd, SEQ, NVF/Palms, litter (1. OMMO- 17293, 17 Dee 198U, J, Stanisic); Kenilworth S.F., SEQ (26°37'S, 152°42°E) raintorest, ltier (1, QM- MQ17294, 22 Muy 1980.5. Sianisic, A. Green); Byan- gum, NSW (1. AMC 152199, Lower); Murwillumbah, NSW (1. AMC28495, ex Tech, Mus.); Northern Rivers. NSW (1, AMC63823, Petlerd, ex Cox): lop of Condamine River Valley, NSW (28"15°S, 152°29°E) (1, AMC128902. 16 Mar 1981, AM/OM - ABRS). DIAGNOSIS Shell diameter t.62—2.47 mm (mean 2.14 mm) of 43/8 to 5 (mean 4 5/8+) tightly coiled whorls, last whorl descending more rapidly. Apes und spire (Fig. 131¢) moderately to strongly elevated, SP/BWW ratio 0,25-0.55 (mean 0.43), heizht of shell 1,19-2,04 mm (mean 1.64 mm). HD ratio 0.71.96 (mean 0.77). Protoconch of 1 WSta |) 3/8 whorls, mean diameter 497.5 pum at | l/2 whorls. Apical sculpture (Fig. 131¢) of slightly curved, evenly spaced, broud radial ribs. FG. ISI MEMOIRS OF THE QUEENSLAND MUSEUM Postnuclear sculpture (Fig. 131f) of strong, slightly protractively sinuated radial ribs, 53-80 (mean 64.1) ribs on the body whorl, Ribs/mm 7.75-12.19 (mean 9.04). Rib interstices about 4-5 times their width. Microsculpture (Fig. 131f) of thread-like radial riblets crossed by low, crowded spiral cords. Umbilicus closed or teduced to a lateral crack. Sutures impressed; whorls slightly shouldered above and rounded below a laterally compressed periphery. Aper- ture roundly Tunate. Lip expanded at the basal and columellar margins. occasionally with a basal denticle, Columella strongly dilated with a tidge-like thickening, reflected over the um- bilicus. Lip sinuous. retracted at the suture to form an apertural sinus. Parietal callus strongly developed. Colour white to light straw-yellow. Based on 14 measured adults. Genitalia with a strongly expanded, muscular epiphallus (Fig. 132b). Epiphallus with ascend- ing arm parallel to the penis and reflexed before entering the penis apically. Penial retractor muscle (Fig. 132b) inserted on the epiphallus prior to its junction with the penis. Penis (Fig, 132b) internally with a apical longitudinal spon- gy pilaster and a basal pad-like thickening. Radula (Fig. 131d.h) similar to F. cochiidium, Based on 6 dissected specimens (QMMOQ- 17012. OMMO17148). RANGE AND HABITAT Warm, moist notophyll vine forests of the Big Scrub area, NSW and the D’Aguilar Range. SRQ, [tis sympatric with E. cochlidtum in the McPherson Ranges. An outlier population at Mt Guyra, north of Gympie, SEQ, is the northern limit of the species, FE, globosa lives under logs, REMARKS Specimens of F. glabasa (QMMO1206%, AM- €136463) trom Mt Guyra are smaller than average and have an altered lip with a small baso-columellar thickening, Omphaloropa gen. nov. Er Maton From the Greek omphalos, umbilicus: refer- ring to the prominent umbilicus. Fgilomen globasa sp, noy, a-c. Sidetrack off Mi Archer Rd, Mt Mee $.F.. SEQ. QMMQ17292. trolly pes sh, he Terania Ck, NSW. OMMOL7I48, paratype. cet, Mt Mee S$... SEO. QMMOQ12178, paratype: Mi Guyra N,P.. SEQ, OMMO/ 2008, paratype, a-e, cotire shell: d. marginal teeth; &, apical sculpture: 1 pose nucledr sculpture: g, entire shell sowing deruicle on lips he central and lateral teeth. Seale lines as marked, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 199 {mm vi te. é i TRS. 200 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 132. Egilomen globosa sp. noy, Terania Ck, Nightcap Range, N.P., NSW. QMMO17012, paratype. a, genitalia: b, penis interior; c, talon and carrefour; d, hermaphroditic duct; e, ovotestis; f. pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 27°S 2B°S = 152°E 153°E 154°E FIG, 133. Distribution of Egilomen globosa sp, nov. Tyre SPECTES Omphaloropa varicosa sp. nov. DIAGNOSIS Shell small, mean diameter 2.23 mm, with lightly coiled whorls (mean 4 1/2-), last descend- ing rapidly. Apex and spire moderately elevated, Protoconch with low spiral cords. Post nuclear sculpture of very widely spaced, prominent, protractively sinuated radial nbs and microsculpture of low radia! riblets and spiral cords which have raised knobs at their intersec- tion. Umbilicus wide, cup-shaped. Sutures strongly impressed. Aperture roundly lunate. lip simple. Columella only slightly dilated. Terminal male genitalia with epiphallus poor- ly differentiated from Vas deferens, entering penis apically through a papillate verge. Penis with a sheath and internal longitudinal pilasters, Female genitalia without unusual features, Radula with very small tricuspid central tooth, otherwise typical. COMPARISONS Omphaloropa is conchologically distin- guished by small size, apical spiral cords, few widely spaced radials on the adult whorls and wide cup shaped umbilicus. Rhophodon is also Zui small, with wide umbilicus and many to few radial ribs on the adult whorls, but in contrast has curved. spaced radials. on the protoconch, and few to many lamellate barriers in the aperture. Sinplovea intensa has a spirally lirate protocanch but more crowded postnuclear radial ribs. Omphaloropa varicosa sp. nov. (Figs 134-136: Tables 26. 27) ETYMOLOGY Latin varix, dilated vein; referring to the prominent postnuclear radial ribs. COMPARISONS Within its tange, O. varicosa is most apt to be confused with Sinployea intensa and species. of Rhophodon, §. intensa has a brown shell with Spiral cords on the protoconch but has very crowded radial ribs on the body whorl and much smaller umbilicus (Fig. 137). Sympatric Rhophodon also have monochrome or flammu- lated shells with wide umbilicus and few to many radial ribs on the adult whorls. However, in contrast to O. varicosa, they have a radially lirate protoconchs with weak spiral elements, and few to numerous apertural lamellae. Tyre MATERIAL HoLotTyPE: QMMOQ17281. Gavial Creek, Boulder- combe, SEQ (23°34’S, 150?28°E) MVF, in litter, collected by J, Stanisic, D. Potter. 9 Jul 1983. Height of shell 1.42 mm. diameter 2.36, H/D ratio 0.60, D/U ratio 2.21, whorls 4 5/8. PARATYPES: SOUTH-EAST QUEENSLAND - QM- MO16805. 88 specimens, same data as holotype; sidetrack off Mt Archer Rd. M1 Mee S.F., (27°04'S, 152°41°E) NVF/Araucaria (3, QOMMO15077, 6 Sept 1983, J. Stanisic, D. Potter); Dawes. Range (24°28'S, 15S1°O7°E) MVF/Araucaria (600+, QMMO12671, AMC136800; AMC136780 QMMQ12651_ 4 Sept 1982. AM/QM-ABRS): Bouldercombe Falls, SEVT (2, OMMO17279, 29 Mar 1983, G, Annabell); Dan Dan Scrub, SEVT, in litter (5, OQMMO16803, 10 May 1984, J. Stanisic, D, Poster); Freds Rd, Mt Mee (27°05°S, 152°43'E), rainforest, leaf liter (7, OMMO- 8363, 14 Apt 1980, J. Stanisic, N. Hall, A Green); Kenilworth S.F. (26°40'S, 182°36°E) Araucaria, leaf litter (15. OMMO17280, 22 May 1980, J. Stanisic, A. Green). OTHER MATERIAL SOUTH-EAST QUEENSLAND - Entrance to slate forest off Yabba Creek Rd. Yabba Creek S.F. (26°28°S, 152°38°E) (1, OMMO13370, 7 Sept 1982, 202 MEMOIRS OF THE QUEENSLAND MUSEUM lmm Imm SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 203 FIG. 135. Omphaloropa varicosa sp. nov. Gavial Ck, Bouldercombe, SEQ. QMMO16805, paratype. a, genitalia: b, penis interior; c, ovotestis; d, talon; e, hermaphroditic duct; f. pallial cavity. Scale lines as marked. FIG. 134. Omphaloropa varicosa sp. nov. a—c, Gavial Creek, Bouldercombe, SEQ. QMMO17281. holotype; d—h, same data as holotype, QMMO16805, paratypes. a—c, entire shell: d. radula; e, apical sculpture; f, post nuclear sculpture; g—h, central and lateral teeth. Scale lines as marked. 204 22°S 24°S 25°S 27°S 2875 30°S 153°E 154°E FIG. 136, Distribution of Omphaloropa varicosa sp. nov. AM/QM-ABRS); Fred’s Rd, Mt Mee (27°05’S, 152°43°E) NVF, litter (1, QMMO15981, 28 Oct 1984, J. Stanisic, G. Annabell); Cunningham's Gap, near monument (28°04’S, 152°24°E) 755m, SNVF/ferns (8, AMC136808, QMMO12679, 7 Dec 1981, AM/QM-ABRS); Little Yabba Ck, Imbil S.F. (26°28°S, 152°38°E) NVF (6, QMMO12020, AMC136437, 8 Sept 1982, AM/QM-ABRS); Montville Rd, near Kondalilla Falls N.P. (26°40°S, 152°52°E) vine thicket on side of road (1, QMMO- 11979, 8 Sept 1982, AM/QM-ABRS). DIAGNOSIS Shell diameter 2.03—2.34 mm (mean 2.23 mm) MEMOIRS OF THE QUEENSLAND MUSEUM with 3 3/4 to 4 3/4 (mean 4 1/2-) tightly coiled whorls last descending rapidly. Apex and spire (Fig. 134c) moderately elevated, SP/BWW ratio 0.18-0.44 (mean 0.29), height 1.03-1.42 mm (mean 1.26 mm). H/D ratio 0.50-0.64 (mean 0.7). Last whorl descending more rapidly. Protoconch white, glossy, exsert, 1 1/2 to 1 3/4 whorls, mean diameter 505.1 pm at | 1/2 whorls. Apical sculpture (Fig. 134e) of 12-15 low spiral cords. Postnuclear sculpture (Fig. 134a) of widely spaced, prominent, protractively sinuated radial ribs, 18-35 (mean 26.8) ribs on body whorl. Ribs/mm 2.79-4.93 (mean 3.83), Microsculpture (Fig. 134f) of fine thread-like radial riblets, 18-35 between each pair of major ribs, crossed by numerous low spiral cords with distinct raised knobs produced at their intersec- tion. Umbilicus (Fig. 134b) wide, cup-shaped, diameter 0.70-1.07 mm (mean 0.91 mm), D/U ratio 2.16-2.92 (mean 2.48). Sutures strongly impressed. Whorls rounded below and shouldered above a rounded periphery (Fig. 134c). Aperture roundly lunate. Lip simple, thickened slightly. Columella only slightly di- lated. Parietal callus strongly developed. Colour brown with alternate light and dark brown radial streaks. Based on 29 measured adults. Genitalia with a poorly differentiated epiphal- lus which enters the penis apically through a large papilla (Fig. 135a,b). Penial retractor muscle inserting on the penis adjacent to the penis-epiphallus junction. Epiphallus internally with longitudinal pilasters. Penis (Fig. 135b) pear-shaped with a distinct bulb and sheath, in- ternally with apical papilla and low, spongy lon- gitudinal pilasters. Based on 3 dissected specimens (QMMO- 15077, QMMO16805). RANGE AND HABITAT Humid subtropical notophyll vine forests in the southern part of its range; drier microphyll vine forests (with Araucaria emergents) and semi-evergreen vine thickets in the north. O. varicosa has been found living among litter and in soil, but not under logs. REMARKS Conchologically O. varicosa could be linked with Sinployea but the simple internal penial structures relate O. varicosa to other charopids reviewed in this study. O. varicosa may be abundant in certain localities e.g, 600+ specimens were collected in SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 205 the Dawes Range, SEQ (QMMO12671, QM- MO12651, AMC136800, AMC136780) in Sep- tember, 1982. s Es Tite (nS Rs4) 449 (UK-527) 2H (2, lheQuay 4a CTA a) Sinployea Solem, 1983 (W.23-0,27) (036-051) (550,78) 35 = Sinployea Solem, 1983, p.81. 7) TYPE SPECIES Sinployea peasei Solem, 1983; by original designation. (27a a = 2 T a = = (5.604743) in HLLUS/MM 6.02 PREVIOUS STUDIES Solem (1983) introduced Sinployea for 49 Pacific Island charopids that show a uniform pattern of anatomical variation, The penis inte- rior is characterised by an apical verge, muscular collar and stimulatory pads in the lower part of the chamber. Conchologically Sinployea is con- servative with fine apical spiral cords, rounded whorls with radial ribs, and microsculpture of radial riblets and fine, low spiral cords in all species. Sinployea includes ‘Cralopa’ intensa Iredale, 1941. whose characters are close to the average Pacific Island species. = = HIBS u5 4 (ST 4ay We GESI) ut Aly Sreuww RATIO wR HLOYEA AND ROTACTIAROMA shy 2-1) ibaa (072-1095) (pt) wr (1th) (hh 2) DISTRIBUTION AND ECOLOGY In the Marianas in Micronesia; parts of Melanesia including Kiribati (= Solomon Is) and Vanuatu (= New Hebrides); and into Polynesia as far east as the Society Islands. Locally S. intensa is confined to subtropical notophyll vine forests of northern NSW and SEQ. Its northern limit is the Rundle Range near Rockhampton. S. intensa often occurs in strand-line rainforests within this region e.g. al Byron Bay, NSW; Cooloola, SEQ; Eurimbula, near Bundaberg, SEQ; and Burleigh Heads, SEQ. 1b fid) ILSay < rh ms (hae8 wit (he TALIM) TART dh RANGIC OE VARLATION IN OMI IVD RATIO as DIAMETER (mm) 200 (77-24) 2.15 C5240) Lat (1, 7H-2, }Ay 10 (2419-2.1) {hyS 2.55) —|,52) WEIOWeL (mm) 1.2K (tL 13-1 4) (1h 1S<1.en) (hua 42) (OMT bh D (3h s Las (I COMPARISONS Sinplayea is not easily confused with other Charopidae. The small, reddish-brown shell, slightly elevated spire, apical spiral cords and postnuclear crowded radial ribs are distinctive. Setomedea is larger, has apical spiral cords, and more complex adult sculpture with periostracal a setae on the major ribs. Cralopa has prominent radial ribs, narrow to closed umbilicus, apertural sinus, and apical sculpture of low, broad, radial ribs and weak, low spiral cords. Omphaloropa (Fig. 134), has apical spiral cords but few major radials on the body whorl, a wide cup-shaped umbilicus, and low microsculpture. fe (4 TRS SIRE) (Fe Ta (aaa) z p = & 5 =x = 2 =< we = t (5 Wht WR) 41/2 A HVR+ + x ve svt uy in i} ™ | dundee NAML Kirtacharipe. MEMOIRS OF THE QUEENSLAND MUSEUM Sinployea intensa (Iredale, 1941) (Figs 137-139; Tables 26, 28) : a 2.21 2.39 2.8020.061 (2.70-2.91) Cralopa intensa Iredale, 1941a, p. 269, fig. 5. COMPARISONS S. intensa is distinguished from sympatric charopids by its small brown shell which has few whorls, small umbilicus, apical spiral cords, and postnuclear sculpture of crowded, protractively sinuated radial ribs. Setomedea nudicostata (Fig. 35) has a flammulated shell and squiggly apical spiral cords. Omphaloropa varicosa (Fig. 134) has a brown to flammulated shell with few prominent radial ribs. Discocharopa aperta (Fig. 93) has crowded radial ribs on adult whorls but is smaller, white to straw-yellow in colour, with wide umbilicus, radially lirate apex, and complex postnuclear spiral sculpture. 0.8420.020 107 0.90 (0.70-00.76) 2.950.090 (2.79-3.10) 3.91 RIBS/MM x = $e 27 tat ol Sa (18-20) ra = G 32 (0.18-0.29) (02-044) O31 0.31 TYPE MATERIAL HOLotyPE: AMC63496, Bryon Bay, NSW. Height 1.15 mm, diameter 2.30 mm, H/D ratio 0.50. D/U ratio 4.00, whorls 3 5/8+. 0.6920.007 (0.66-0.72) 0.640.031 (0.58-0.68) BS Se 3 = ee (mm) 0.72 0.66 WIL BODY WHORL (0.64-0.70) OTHER MATERIAL AMC31650, AMC63769, 4 specimens, Cape Byron, NSW. Collected Lower, ex J.C. Cox; Kenilworth S.F., SEQ (26°37°S, 152°42’E) rainforest, leaf litter (1, QMMO17284, 22 May 1980, J. Stanisic, A. Green); Mt Mudlo, Kilkivan S.F., SEQ (26°01’S, 152°13’E) Araucaria/NVF, litter (13, QMMO17283, 17 Jul 1980, J. Stanisic, A. Green); Kalpowar S.F., SEQ (24°43°S, 151°21°E) MVF/Araucaria, litter (12, QM- MO17287, 15 Jul 1980, J. Stanisic, A. Green); rain- forest walk, Kalpowar S.F., SEQ (24°41°S, 151°21’E) MVF/Araucaria (1, QMMO12576, 4 Sept 1982, AM/QM-ABRS); Below summit, Mt Warning, NSW (28°24’S, 153°16°E) (1, QMMO10511, Mar 1981, AM/QM-ABRS); Mt. Fort William, Kalpowar S.F., SEQ, 833 m, CNVF (1, QMMO16824, 3 Jul 1984, J. Stanisic, D. Potter, K. Emberton); Whian Whian S.F. N. of Lismore, NSW, rainforest (1, QMM06294, 12 Sept 1976, M.J. Bishop); Mt Guyra N.P., SEQ (25°49’S, 152°35’E) MVF/NVF/Araucaria (2, QM- MO12075, 7 Sept 1982, AM/QM-ABRS); Burleigh Heads N.P., SEQ, vine thicket, under logs (2, QMMO- 16917, 21 Jan 1987, J. Stanisic, D. Potter); Freds -- « ¢ ” 7s Road, Mt Mee SEQ (27°05’S, 152°43’E) NVF, litter (1, QMMO15980, 28 Oct 1984, J. Stanisic, G. An- nabell); Cooloola N.P., SEQ (25°55’, 153°08’E) 50m, CNVF on sand (1, QMMO06350, 25 Oct 1976, M.J. 0.21 (MEAN AND RAN 0.5520.027 (0.50-0,59) 0.60 0.54 H/D RATIO 2.2120.025 an f¢ aoe So za aS nici (2.12-2.36) 2.0420).007 DIAMETER (mm) 36 2.16 TABLE 27- LOCAL VARIATION IN OMPHALOROPA VARICOSA SP. NOV. 2,28 mam — HEIGHT (mm) 1.2620.030 (1.21-1.34) w a 1.42 iy 1.23 ( a. (ut (4 3/R+to4d 3/4) K = 2 45/8 4 3/8+ 45/84 45/8- (33/4-4+) 48+ 45/8+ (4 1/8458) Fs 5 ae ge Bishop); sidetrack off Mt Archer Rd, Mt Mee S.F., 2 32 2 32 332 SEQ (27°04’S, 152°41°E) (1, QMMO12180, 31 Sept z go 5 35 365 1982, AM/QM-ABRS; Byangum, Tweed River dis- UIMUILEC AL (MULAN, SEM AND RANGL) TALL 24 - LOCAL VARIATION IM SANELOYEAINTEASA (IREDALE, 1941) SINR NAMI SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 207 trict, Nth NSW, serub, (2, AMC63779); Cape Byron, SVE Ww HOODY WHORL = ss ES Nib NSW, (2, AMC140493, 1 May 1958, L.. Price); 5S 268i #5 8 SS 554 § Byron Bay, NSW (2, AMC140483, C.F. Me- = . a Lauchlan); South Grafton, NSW (2, AMC.63785); Mudlo Gap. Mudlo $.F., SEQ. MYF/Araucaria, = 2 32 Ze (26°01°S. 152°14°B). (55, AMC136654. QMMO- Sf 23: #2 ¢ « 3 He zs 12352, 2 Oct 1982, AM/QM-ABRS); Yarraman 2 fS~ Sa AB FEF © Forest Drive, Yarraman S.F., SEQ, MVF, (26°50’S, 2 _ -181°57°E). (3, AMC136587, OMMOQ12262, 31 Sept RA xi 1982, AM/QM-ABRS); Cooloola Nat, Park, SEQ, 5 eit, -¢ £ & ze SNVF on sand (25°57’S, 153°06°E) (2. AMC 136489, i == QMMO12112, 6 Sept 1982, AM/QM-ABRS); Bobby Range S,F. SEQ, NVF, (24°37°S, 151°32°E) (10, _ 2 = AMC136505, OMMO12141, 6 Sept 1982, AM/QM- z de, z= 0 oz ABRS): Blackbutt Range, c. 7.4 km § of Benarkin, ES os: os » £ 8 See 8 SEQ, MVF/Araucaria (26°53°S, 152°11'E), (5, AMC136565, QMMOQ12239, 31 Sept 1982, J. bes wie Stanisic, D. Potter); Mt Fort William, Kalpowar SF, = zs a= SEQ, NVF (24°39°S, 151°20°E) (2, AMC136759 S = s2 2 3 3 ES QMMO12690, 4 Sept 1982, AM/OM-ABRS); NE of ~~ Monto, Kalpowar SF., SEQ, vine thicket/serub litter (24°42°S, 151°21'E)(2, AMC137851,QMMO13153, a g zz = 4 Sept 1982, AM/QM - ABRS); NW of Miriam Vale. Sst 2 So* tee ae Eo oF Colosseum Ck., SEQ, rainforest along creek (24°23'S, ae SS ye eee 151°27°E) (7, AMC137894, OMMO13264, 6 Sept ' 1982, AM/QM-ABRS); Bulburin SF, S of Gladstone, 2 zs «2 z = SEQ, 540 m, CNVF-AI forest type (24°31'S, ®¢ <2 234 22 » 2 « 2 Z —-151°29°E) (1, AMC152213, 8 May 1975, W.P. a= 587 eS) 3 Fe = Ponder, J.B, Burch, PH. Colman); Eurimbula S.F.. S of Gladstone, SEQ, 10 m (24°11°S, 151°50°E) (4, = ew, - es, AMC152214, AMC152215,7 May 1975, J.B. Burch, z - 28 ef =o W.F. Ponder. P.H. Colman); Rundle Range, SE of eg ¥ ee 2 2 2 282 &2 Rockhampton, SEQ, 30 m (23°39°S, 150°59°E) i “ft A , (many, AMC 152216, AMC152217 6 May 1975, J.B, i Ps Burch, W.F. Ponder. P.H. Colman), ZF #. 2£ 5 » z DIAGNOSIS Shell diameter 1.95-2.55 mm (mean 2.31) with 3 1/2+ to 3 7/8- (mean 3 5/8+) loosely coiled a5 whorls, last whorl descending more rapidly. ‘ Hi 4 js « Apex and spire (Fig, 137c) flat to slightly ; TS (Sees elevated, SP/BWW ratio 0.020. 19 (mean ().12), height of shell 0.97-1.3! mm (men 1.13 mm). a = H/D ratio 0.45-0,54 (mean 0.50), Protoconch of = 2ee £ ye Fae ae 1 3/8 to 1 1/2 whorls, mean diameter 574.8 pm 2 a a “5 at | 1/2 whorls, sometimes exsert but generally flattened. Apical sculpture (Fig. 137d) of c. 12 z glossy. evenly spaced, fine spiral cords. = cere te = & = er WF Postnuclear sculpture (Fig, 137c) of complex, > protractively sinuated, crowded radial ribs, §1— NUMMER OL 116 (mean 92.1) ribs on the body whorl. _- @ = Ribs/mm 10,65-16.79 (mean 13.11). Micros- Sa = culpture (Fig. 137e) of fine radial riblets, S11 gz #2 23 between cach pair of major ribs, and low spiral <3? 8 =° cords which form raised knobs at their intersec- 208 MEMOIRS OF THE QUEENSLAND MUSEUM Cc FIG. 137. Sinployea intensa (Iredale, 1941). a—c, Bryon Bay, NSW. AMC63496, holotype; d-e, Mudlo Gap, Mudlo S.F., SEQ. QMMO12352; f-g, Cherry Tree North S.F., via Casino, NSW. AMC154769. a-—c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, central and lateral teeth; g, marginal teeth. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 209 Gtr be pera ee: FIG. 138. Sinployea intensa (Iredale, 1941). Burleigh Heads N.P.. SEQ. QMMO16917. a, genitalia; b, d, details of penis; c, talon and carrefour: e, ovotestis: f, pallial cavity. Scale lines as marked. malt) 2A°s 259s 26°S 27°S 26°S a ISS Rjalt. > 500 WR 30°s 150°E 151°E FIG. 139. Distribution of Sinployea intensa (Iredale, 1941). tion with the microradials. Sculpture continuous on the base. Umbilicus (Fig. 137b) wide, V- shaped, last whorl decoiling more rapidly, diameter 0.55-0.78 mm (mean 0.66 mm). D/U ratio 3.17—4.00 (mean 3.78). Sutures strongly impressed, Whorls rounded above and below the periphery (Fig. 137c). Aperture roundly lunate. Lip simple, slightly thickened. Columel- la weakly twisted toward umbilicus. Parietal cal- lus well developed, margins convergent. Colour brown. Based on 17 measured adults. Epiphallus (Fig. 138d) short, muscular, reflex- ing before entering the penis apically, internally with longitudinal pilasters. Penis (Fig. 138a) short; externally differentiated into an upper and MEMOIRS OF THE QUEENSLAND MUSEUM a lower chamber; without sheath. Details ot upper chamber and epiphallus entry unavailable. Lower chamber with a semicircular (horseshoe- shaped?) pilaster (Fig. 138b). Penial retractor muscle (Fig. 138) inserting on the penis ad- jacent to the epiphallus-penis junction. Radula (Fig. 137f.g) typical for genus. Based on | dissected adult (QMMO16917). RANGE AND HABITAT Humid sub-tropical notophyll vine forest in northern NSW and SEQ; drier microphyll vine forest between Gympie and Rockhampton, SEQ: and strand line vine forests at Bryon Bay. NSW. Burleigh Heads, Cooloola and Eurimbula, SEQ. The latter habitat preference is unusual in the context of Australian species but as a remnant of a much wider Pacific Island distribution it is not unexpected. S. intensa lives under the bark of fallen trees and has the typical grey markings also present in semi-arboreal Pacific Island Sinployea. REMARKS S, intensa is similar to. 8. euryomphala (Solem, 1959) from Vanuatu in external appearance of penis, insertion of penial retractor muscle, shape of epiphallus, unusual muscular collar on the free oviduct, and pallial configuration (Solem, 1983). However S. intensa has more postnuclear ribs, is smaller jn size, and has a less elevated spire. Subfamily ROTADISCINAE Baker, 1927 This taxon was introduced (Baker, 1927) for endodontoid snails with incomplete secondary ureter. and kidney witha rectal lobe much Jonger than pericardial lobe. Solem (1983, p. 70) in- cluded the subfamily in the Charopidae for Species with or without a complete secondary ureter and weakly to strongly bilobed kidney. He considered that the pecultar apical sculpture and unusual variations in genital morphology were (he main distinguishing features and in- cluded the Northern Hemisphere Rotadiscus Pilsbry, 1926, Radiadiscus Pilsbry and Ferriss, 1906, Radioconus H.B. Baker, 1927 and Radiodomus H.B. Baker, 1930, together with Microcharopa Solem, 1983 from the Pacific region. Rotacharopa gen. nov. ETYMOLOGY A combination of syllables from Rotadiscinae and Charopidae. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE TYPE SPECIES Rotacharopa annabelli sp. nov. DIAGNOSIS Very small to minute Charopidae, diameter range 1.77—2.46 mm, with 4 5/8+ to 6 1/8 very tightly coiled whorls, last descending more rapidly. Apex and spire weakly to moderately elevated. Apical sculpture of short, slightly twisted segments, organised into spiral rows, and weakly curved, low radial ridges. Postnuclear sculpture of very crowded (densilamellata) to moderately spaced (annabelli), protractively sinuated radial ribs. Microsculpture of very fine radials and low, broad spiral cords which are raised at their intersection with the microradials to form an elongate bead. Umbilicus very nar- row, U-shaped to almost completely closed by reflection of the columella. Sutures impressed; whorls shouldered above and rounded below a laterally compressed periphery. Aperture roundly lunate. Parietal callus weakly developed. Animal with slender foot and tail. Kidney with well developed, vaguely triangular, elongate pericardial lobe and a reduced rectal lobe. Ovotestis two teardrop-shaped lobes lying in the apical whorls of the digestive gland. Terminal male genitalia with epiphallus differentiated into ascending and descending arms. Descending arm of epiphallus with transverse thickenings internally. Epiphallus entering penis through a simple pore surrounded by a large fleshy pilaster. Penial retractor muscle inserted on the epiphal- lus, or on the epiphallus - vas deferens junction (annabelli). Penis large, barrel-shaped, with (kessneri), or without a verge, normally with longitudinal pilasters which may be modified (annabelli). Penis with short (annabelli), long (kessneri) to extremely long (densilamellata) preputial tubular extension. Vagina long, rarely short (annabelli), internally with longitudinal thickenings. Free oviduct short. Atrium without unusual features. Spermatheca typical. Radula with tricuspid central and lateral teeth (central slightly smaller) which have a long slender lan- ceolate mesocone and short, pointed ectocones. DISTRIBUTION AND ECOLOGY Rotacharopa is confined to moist notophyll vine forests and drier microphyll vine forests and vine thickets of south to mid-east Queensland. Undescribed rotadiscines, with flatter shells and more open umbilici, also occur in this region (Stanisic, unpublished). Rotacharopa kessneri 211 and R. annabelli have been collected in close proximity to each other near Gladstone, SEQ, and further fieldwork may demonstate sympatry. Both species have modified penial surfaces which indicate possible microsympatry but this sympatry may also involve undescribed taxa. The two southern species, R. densilamellata and R. kessneri live under logs, whilst R. an- nabelli is a litter dweller. COMPARISONS Other charopids reviewed in this study which have similar-sized shells lack the peculiar apical sculpture of Rotacharopa. Rhophodon has a flattened shell with wide umbilicus, apical sculp- ture of prominent radial ribs, and apertural bar- tiers; Discocharopa has apical sculpture of prominent radial ribs and weak spiral wrinkles, and a saucer-shaped umbilicus; Omphaloropa has a very wide cup-shaped umbilicus and con- tinuous spiral cords on the protoconch; and Sinployea intensa has continuous apical spiral cords, fewer whorls and a more open umbilicus. Microcharopa Solem, 1983 from Fiji is readily distinguished from Rotacharopa by its lower whorl count, smaller size, flatter shell and very wide umbilicus. KEY TO SPECIES OF ROTACHAROPA 1.Postnuclear sculpture of widely separate radial ribs (mean ribs/mm 6.62) (Fig. 140a) Load dea Mie upeshiedstet cl cangeanedsn dean ivddyaae annabelli Postnuclear sculpture of more crowded radial Tibs (Figs 143a, 1464) oo. eeeeeeeseeeeeeees 2 2.Postnuclear radial ribs very crowded; umbilicus more open (Fig. 146b), mean D/U ratio 4.93 AAS TORY AVES FAP TIRLANY.5 POOP PO densilamellata Postnuclear radial ribs less crowded; um- bilicus more closed (Fig, 143b), mean D/U TATOO. TD ee eccccssecesessseseesenteeeeeenes kessneri Rotacharopa annabelli sp. noy. (Figs 140-142; Tables 26, 29) ETYMOLOGY For Graeme Annabell who provided assistance with collecting. 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VdOUVHOVLON ONY “AON AS ITEIIVNNY VdOUVHOVLOY NI NOLLWRIVA WOOT (66'1-S6'1) 9100FL6'L (9b 2-972) 190'0FRE'T 661 (207-76) TI OFEOT rand wee 6g (91z-S8' 1) SSUFIO'Z Lratg (OZ'2-81'2) OLOOF ONE fal (97-912) MOL 0F9T'Z (9z'2-91'2) 9ZNOFIZZ (seI-Le 0) HOUT T 681 (uw) waLaWVIG (ez 1-171) a1odFr2't (ose) RSOOFOST tv (uz t-ee) eoruryz' | cel fee svi (ez I-or) 072007 12'1 ort (os t-Rb 1) GLU OF HPT rel 6T0UFLP (eri) HMOFSUL tet (wu) LHOIH 60 PAV, (B/E SO}+H/L #) RIS (48/5 So-z/I S$) “B/S S (py sHIL Y) +s +H/ES tus “RLY (s-R/L P) +H/L +P RIL S WSS (R/L SO1+H/6 S) +e/ES (-8/1 9-R/S S) *RILS SIE S) +S a STHOHM £ £97£1 OWWO y2aat) WINISSO|O.) t zOR91 ONWO qrung ung UEC 1 P0LL1 ONWO MRWOUTYOOL IAL £ Sorel ONY 9CR9OT OWWO uopuad tig iW. ! sac! ONWO mur I LZI£| ONWO yaar auoysauiyy £0€41 ONWO (adAiojoH) i Z0¢L1 ONWO “ys anmadey, uaaussay pdospysmioy nn t c0zt1 ONWO MoO, IW. 1 LILEL OWWO uraoW IW t L787 OAWO + LOR9L OWWO aquioaapyNog o RPER OWWO (ado) t 00¢21 OWWO sayy 1 yjaqouut pdoanyomoy SNAWIOAdS AO WAIN AINYN SYSTEMATICS OF AUSTRALIAN CHAROPIDAL large whorl count, narrow umbilicus, details of internal penial sculpture and position of penial retractor muscle insertion separate R. annabelli from its congeners, TYPE MATERIAL HoLotyee: QMMQ17300, Mt Archer, Rock- hampton, MEQ, leaf litter, Collected by G. Annabell, 23 Dec 1978. Height of shell 1.23 mm, diameter 1.89 mm, H/D ratio 0.65, D/U ratio 8.36, whorls 6 1/8. PARATYPES: QMMO8348, 26 specimens. same col- lection data as holotype: Mt Archer, Rockhampton, MEQ, about one third of the way up (23°21'S, 150°35*E), remnant vine thicket in gully (62, OMMO- 11659, OMMQ11919, 30 Jun 1982, J. Stanisic); Gavial Creek, Bouldercombe MEQ (23°34’S, 150°28°E), MVF, litter (24, QMMO16807, 9 July 1983, J. Stunisic, D. Potter); Bouldercombe. W of Rockhampton, MEO, rainforest litter (10, QMMO- 12827, 3 Feb 1980. V, Kessner); ¢. 7 km E of Mt Morgan, MEQ (23°387S, 150°28"E) MVF/NVF, litter (1, OMMO17301, 11 Jul L980, J. Stanisic, A, Green, 5. OMMO13117, 29 Jun 1982. J, Stanisic; 4, QMMO- 11633. 29 Jun 1982: J. Stanisic); W side of Mt Moore. SW of Gladstone, MEQ, in patch of vine thicket, liner (24°02°S, 151°05°E) (5, AMC137875, OMMO13203. 5 Sept 1982, AM/QM-ABRS). QITHER MATERIAL. Calliungal, MEQ (1, AMC264. CT, Musson); W of Gladstone, MEQ. underleaves.on groundin bottle tree serub (1, AMC32996, Aug 1908, SW. lackson), DIiAGwosis Shell diameter 1.77—2.36 mm (mean 2,00 mm) with 5 3/8 to 6 1/8 (mean 5 3/4-) tightly coiled whorls, the last descending more rapidly. Apex and early spire (Fig. 140c) flattened to depressed, SP/BWW ratio 0.11-0,26 (mean 0.16), height 1.13-1.54 mm (mean 1.28 mm). H/D ratio 0.52-0,69 (mean 0,63), Protoconch of 15/8 to 1 3/4 whorls, mean diameter 343.5 jm at | 1/2 whorls. Apical sculpture (Fig. 140e) of short segments arranged in 14-18 spiral rows crossing a series of curved, broad radial ribs and ridges, Postnuclear sculpture (Fig. 140f,g) of line, widely spaced, weakly protractively sinuated radial ribs, 34-51 (mean 41.3) on the body whorl. Ribs/mm 5.60-7.45 (mean 6,62). Microsculpture (Fig. 140f) of very fine radial riblets, 9-16 between each pair of major ribs, crossed by low spiral cards which are raised at their junction with the microradials, producing a beaded effect. Umbilicus (Fig, 140b,h) very nar- raw, U-shaped to completely closed by reflec- lion of the columellar margin, diameter 0.23- 0,27 mm (mean 0.25 mm). D/U ratio 6.85-8,54 (mean 7.66). Sutures impressed. Whaorls shouldered above and rounded below a laterally compressed periphery (Fig. 140c), Aperture roundly lunate. Lip simple, columella slightly to strongly reflected over the umbilicus. Parietal callus well developed. Colour golden-yellow horn, Based on 19 measured adults. Epiphallus (Fig. 141a) muscularised with as- cending and descending arms, internally with well developed transverse thickenings (Fig. 141b), entering penis through a simple pore sur- rounded by a large circular pilaster. Penis (Fig. 141b) elongate, with a short basal extension (Fig. 141a), internally with strong pustules and ir- regularly shaped pad-like thickenings (Fig. 141b). Penial retractor muscle (Fig. 1414) in- serted at the epiphallus - vas deferens junction. Radula (Fig. 140d) with central tooth slightly smaller than laterals. Based on 3 dissected specimens (QMMO- 11633). RANGE AND HABITAT In leaf litter in dry microphyll vine forests and vine thickets between Gladstone and Rock- hampton. SEQ. REMARKS The shift in insertion of the penial retractor muscle to the epiphallus - vas deferens junction is most likely related 1o whorl count increase. Rotacharopa kessneri sp. nov. (Figs 143-145; Tubles 26, 29) ETYMOLOGY For Vince Kessner, in recognition of his can- tribution to Australian malacology. COMPARISONS R. kessnert differs from R, densilamellata in having coarser sculpture, a less elevated spire and narrower umbilicus. Anatomically it lacks the very long penial extension of R. densilame!- lata. TYPE MATERIAL HovoTyPe: QMM()17302, ¢. 8 km from Kalpowar on Fireclay Rd, Kalpowar S.F.. SEQ (24°43'S, 151°21°R), MVFi/Araucaria, litter. Collected by J. Stanisic, A, Green, 15 Jul 1980. Height of shell 140 mm, diameter 2.28 mm, H/D ratio 0.61, D/U ratia 11,12, whorls 5 1/44. MEMOIRS OF THE QUEENSLAND MUSEUM 214 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE FIG, 141, Rotacharopa annabelli sp. nov. c. 7km E of Mt Morgan, SEQ. QMMOQ11633. a, genitalia; b, details of penis interior; c, ovotestis; d, hermaphroditic duct; e, pallial cavity. Scale line as marked. FIG. 140. Rotacharepa annabelli sp. nov. a—c, Mt Archer, Rockhampton, MEQ. QMMO17300, holotype: d, c. 7km E of Mt Morgan, MEQ. QMMO1 1633, paratype; e—h, same data as holotype,, QMMO8348, paratype. a—c, entire shell; d, lateral teeth; e, apical sculpture: f, post nuclear sculpture; g, details of microsculpture; h, base. Scale lines as marked. Bal) 22°95) FIG, 142. Distribution of Rofacharapa aniabelli sp. nov. PARATYPES: QMMOQ17303, 6 specimens. same data as holotype: Mt Goonaneman, via Childers, SEQ, rainforest (3, OMMO1 7304, 7-8 Feb 1981, R, Raven, G. Monteith); high up on river bank beside road. Cania Gorge. SEQ (24°4 1S. 150°S8"E), rainforest, litter (2, OMMO17305. 14 Jul 1981, 7. Stanisic, A. Green); Dan Dan Serub, SEQ. SEVT. litter (4, OMMO16802. 10 May 1984, J. Stanisic, D. Potter); Mt Biggenden., SEQ, SEVT on limestone outcrop, Jitter (1, QM- MO16836, 3 Jul L984, J. Stanisic. D. Potter, K. Em- berton): SW of Miriam Vale, Old. Colosseum Ck. rainforest along creek (24°23°S. [5127°F) (8, AMC137895, QMMO13265, 6 Sept 1982, AM/QM- ABRS): Mi Fort William, Kalpowar $.F.. SEQ. NVF (24°39°S_ 151720" E) (3, AMC 136755, QMMQ12599, 4 Sept 1982. AM/OM-ABRS): W of Childers, Lime- stone Ck §,F. (25°15°S, IS1°SS°E) (5, AMC137830. OMMO13127, 3 Sept 1982. AM/OM-ABRS); Ruin- forest walk. Kalpowar S.P.. MVE. SEQ (24°41°S, MEMOIRS OF THE QUEENSLAND MUSEUM 151°2VE) (3, AMC136743, QMMO12575, 4 Sept 1982, AM/QM-ABRS). OTHER MATERIAL Bulburin S.F..'S of Gladstone, SEQ, 580 m (24°34'S, 151°29°E) (1, AMC1{52210, 8 May 1975, J.B. Burch, W.F. Ponder, P.H. Colman); Bulburin S.F., S of Gladsione, SEQ, 540 m, CNVF-A1 forest type (24°31°S, 151°29°E) (11. AMC152211, 8 May 1975, W.F. Ponder, J.B. Burch, P.H, Colman); Bobby Range S.F., SEQ, NVF (24°37’S, 151°32'E) (12, AMC- 136504, QMMO12138, 6 Sept 1982, AM/QM- ABRS); Clifton Range §.F.,SW of Fairlics KnobN,P., SEQ, Vine thicket/Araucaria (25°34°S, 152°16'E) (2, AMC136672, QMMO12376, 2 Oct 1982, AM/QM- ABRS); c. 12.1 km NW Granite Creek Crossing, on Bobby Range - Mt Perry Rd, SEQ, NVF/palms, under logs (2, QMMO16844, 16 Sept 1955, J. Stanisic, D. Potter): summit, Mt Booroon Booraon, SW of Miriam Vale, SEQ, MVFi/Araucaria, among rocks (5, QM- MO16849. 17 Sept 1985, J. Stanisic, D. Porter). DIAGNOSIS Shell diameter 1.85—2.46 mm (mean 2.15 mm) with 4 7/8- to 5 5/8+ (mean 5+) tightly coiled whorls, last whorl descending more rapidly. Apex flat (Fig. 143c), spire slightly to moderate- ly elevated, SP/BWW ratio 0,10-0.24 (mean 0.15), height 1.15-1.60 mm (mean 1.3] mm), H/D ratio 0.57—-0.65 (mean 0.61). Protoconch of 1 5/8 whorls, mean diameter 385.7 wm at 1 1/2 whorls. Apical sculpture (Fig. 143c) of short Stgments arranged in spiral rows and curved, broad radial ridges and ribs. Postnuclear sculp- ture (Fig. 143f.g) of fine. moderately spaced to crowded, weakly protractively sinuated radial ribs, 57-146 (mean 95.8) ribs on the body whorl. Ribsimm §.83-19.32 (mean 14,19). Micros- culpture (Fig. 143f) of fine radial riblets, 5—L0 between each pair of major ribs, and low spiral cords which are raised at their junction with the microradials to form a bead. Umbilicus (Fig, {43b,h) narrow, U-shuped, diameter 0.16-0.29 mm (mean 0.23 mm). D/U ratio 7.23-11.90 (mean 4.71). Sutures impressed. whorls shouldered above and rounded below a broadly rounded periphery (Fig. 143c). Aperture broad, roundly lunate. Lip simple. columella margin slightly twisted toward umbilicus. Parietal cal- FIG. 143. Rotacharapa kessnert sp. nav. ac. c. 8km from Kalpowar on Fireclay Rd. Kulpowar $,F., SEQ. QMMO! 7302. holotype: c. 12.)km N of Granite Ck crossing Bobby Range. SEQ. OMMO16844: c-h. Colosseum Ch. SW of Miriam Vale. SEQ, QMMO1326S. paratype. aec. entire shell; d. central und Jaleral teeth: ©. apical sculpture: f post nuclear sculpture: 2 details of micrasculprure: h. hase, Seale lines as marked, 217 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE —— (ee 218 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 144. Rotacharopa kessneri sp. nov. Summit, Mt Booroon Booroon, SW of Miriam Vale, SEQ. QMMO16849, a, genitalia: b. details of penis interior: c, hermaphroditic duct: d, talon and carrefour: e. ovolestis: f, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 25°S 26°S 150°8 151°E 152°E 153°5 FIG. 145. Distribution of Rotacharapa kessneri sp. nov. lus developed. Colour beige to light-golden horn. Based on 18 measured adults. Epiphallus (Fig. 144a) with short, narrow as- cending arm and much wider descending arm that has thick, transverse, fleshy pilasters inter- nally (Fig. 144b). Epiphallic pore (Fig. 144b) simple, surrounded by a weak fleshy collar. Penis (Fig. 144a) with a wide upper chamber, a lower, cylindrical section, and a basal extension (Fig. 144a), Internally, penis with strong pus- tulations arranged into longitudinal thickenings (Fig. 144b) and an apical conical verge. Penial retractor muscle (Fig. 144a) inserting on the epiphallus at the point of reflexion. Radula (Fig. 143d) typical. Based on 4 dissected specimens (QMMO- 16849. OMMO16544). RANGE AND HABITAT Under logs in dry microphyll vine forests and vine thickets between Biggenden and Calliope, SEQ. It has also been found in the wettet warm subtropical notophyl! vine forests of the Bobby Range and Mt Fort William. The species appears (o be able to tolerate drier conditions and occurs in ‘fringe’ rainforest in the Cania Gorge, near Monto, SEQ. 219 REMARKS R. kessneri may be sympatric with RX, annabelli because the two species have been collected within only a few kilometres of each other west of Gladstone, SEQ. Rotacharopa densilamellata sp. nov. (Figs 146-148; Tables 26, 30) ETYMOLOGY Latin densus, crowded; and lamella, plate; referring to the postnuclear sculpture of crowded ribs. COMPARISONS The small, globose shell with many tightly coiled whorls. elevated spire, crowded, fine radial ribs. and small umbilicus, identify R. den- silamellata in SEQ. Anatomically the peculiar preputial extension of the penis (Fig. 147a) is distinctive. The only other species apt to be confused with R. densilamellata is the sympatric Egilamen globasa, but it has adult sculpture of fewer, broad radial ribs, a completely closed umbilicus, and a sinuate lip with apertural sinus. TYPE MATERIAL HoLoryre: QMMO17296, Kenilworth S.F.. SEQ (2637'S, 152°42’E) rainloresl, leaf litler, Collected by J. Stanisic. A. Green. Height of shell 1.38 mm. diameter 1.95 mm, H/D ratio 0.71, D/U ratio 5.27, whorls 4 3/4-. PARATYPES; QMMO17297, 4 specimens, same col- lection dala us holotype; sidetrack off Mt Archer Rd, Mt Mee S8.F., SEQ (27°04°S, 152°41°B) NVF/Araucaria (20, QMMO1508), 6 Sept 1983, J. Stanisic, D, Potter); Kenilworth $.F., SEQ (26°35'5, 152°3G°E) mixed NVF (12, QMMQ6285, 18 May 1976, M.J. Bishop); Fred’s Rd, Mt Mee, SEO (27°085°S. 152°43'E) rainforest. leaf litter (11, OMMO8347, 14 Apr 1980, J. Stanisic, N. Hall, A. Green; (6, OMMO15S9S4, 2K Oct 1984, J, Stanisic, G, Annabell); Litthe Yabba Creek, Imbil 5.F,, SEO (26°28°S. 182°38'E) NVF (4. OMMO12018, AMC- 136435, 8 Sept 1982, AM/QM-ABRS); Kenilworth S.F., SEO (26°40'S, 152°36 E) Araucaria, leat litter (12, OMMO8346_ 22 May 1980, J. Stanisic, A, Green); c. 25.5 km and 28.5 km trom Goomburra on Goomburra 8.F. Road, SEQ NVF/Palms/Araucaria (8, OMMOQO12694. AMC136815_ 7 Dec 1981, AM/QM-ABRS; 1, QMMO11125. 16 Mar 1981, AM/QM-ABRS); “Rainforest Park’, near Maleny, on Maleny-Montville Rd, SEQ. NVF, under log (1, QM- MO1LI&S3_5 Dec 1982, J. Stanisic): Kenilworth S.F,, SEQ (26°35°S, 152°42°E)c. 80 m, rainforest/eucalypt 220 PATIO LOCAL VARIATION IN HOPI OHARODA DUN STEAMELEA TASTE ISI [MEAN SEM AND RANTES UMBILICAL SAW Ww RATIO HOY WHE Su MUIMMINK Cb NAME Di RATIO Whit RIUS/MM MS WIDE (mm) VOU STON HE TGHEE DIAMETER WO RAL (rm) tere) (mm) WHONKTS, SECIMENS, {men 52 =n Ss =5 vist 4.7 > 2 = = 5 = = <= 5 = ” r a > x a 2 > o x 6 G < = = - = by 2 = = 2 : < 3 ca z= = =r 3 33 2= 22 2= <= == 22 2- 3 > = MEMOIRS OF THE QUEENSLAND MUSEUM SAS a7 (IM ALE TEAK) Thee (20 527 Sb (iy) Hh thsiLNas (0 aany (76-178) = Aa mt AP HEEAIKIS (On TU) 1 AMS LUNT Et) (ashe Dee MI Mee = 3 a 21,72 = RO — aire we (ead ay 5 (MMOD 1 S0KT (1.04 44) ah }s2 that = al (0.49) O47 2M lna OW 36 AK Tey (ha iu rs = = = = = a a7 ia 4 al they AS 47% Lak 12H W399 ad Ty at > iso emergents (22. QMMO6284, 3 May 1976, M.J. Bishop); Cunninghams Gap N.P., SEQ. near monu- ment, 755 m. SNVF/ferns (28°04'S, 152°24°E) (104, AMC136805, QMMO12676, 7 Dee 1981, AM/QM- ABRS); Montville Rd near Kondalilla Falls N.P., SEQ, vine scrub on roadside (26°40°S, 152°52’E) (2. AMC136411, QMMO11980, 8 Sept 1982, AM/QM- ABRS); Cunninghams Gap N.P., SEQ, near monu- ment, MVF/Araucaria (28°04'°S, 152°24°E) (10, AMC128609, OMMO10993, 16 Mar 1981, AM/QM- ABRS); sidetrack off Mt Archer Rd, Mt Mee S.F., SEQ NVF/Araucaria (27°04°S, 152°41°E) (6, QMM0O12176, AMC136522. QMMO12179, 31 Sept 1982, AM/OM-ABRS). OTHER MATERIAL. Little Yabba S.F,, SEQ (26°34°S, 152°39'E) c. 160 m, eucalypt and vine forest (1, QMMO6301, 3 May 1976, M.J. Bishop); Maiala N.P., Mt Glorious, SEQ, NVF, leal litter (1. QMMOII853, 20 Jun 1982, MSA party): Boombana N.P., Mt Nebo, SEQ, wet sclerophyll (1. QMMO6298, Aug 1976, M.J. Bishop): Campbell’s Camp, Kenilworth S.F,, SEQ (26°42°S, 152°37°E) CNVF, leaf litter (1, QMMO17298, 1 Jul 1980, J. Stanisie, A. Green): Cunningham's Gap, SEQ, leaf litter (1, QMMO17299, 18 Jun 1979, G. Annabell): ¢.0,3 km N of Litthke Yabba Ck, on Maleny- Kenilworth Rd, Kenilworth S,F.. SEQ (26°37'S, 152°42°E) litter (2. QMMO13434. 19 May 1983, K. Collins. J. Stanisic). DIAGNOSIS Shell diameter !.79-2.16 mm (mean 1.94 mm) with 4 5/8+ to 5 7/8- (mean 5 1/8+) tightly coiled whorls, last whorl descending slowly. Spire elevated (Fig. 146c), apex tending to be flat- tened, SP/BWW ratio 0.21-0,53 (mean 0,34). height 1.23-1.52 mm (mean 1.35 mm), H/D ratio 0.67-0.78 (mean 0.71). Protoconch of 1 1/2 to 1 5/8 whorls, mean diameter 428.3 «tm at | 1/2 whorls, Apical sculpture (Fig. 146d.e) of short segments arranged in 19-25 prominent crowded spiral rows overlying broad, curved radial ribs and ridges. Postnuclear sculpture (Fig. 146f) of narrow. crowded. weakly protrac- lively sinuated radial ribs, 93=206 (mean 139) on the body whorl, whose intertices are +—5 times their width. Ribs/mm 16.18-36.68 (mean FIG, 146. Ratacharapa densilamellata sp, nov. a-c, Kenilworth S.F.. SEQ. QMMO1 72496, holotype: d-f. same duta as holotype. QMMO8345, paratype: g-h. Cunningham's Gap, SEQ. QMMO 12676. paratype. a-c.entire shell: d. apical sculpture: e. details of upical Spiral cords; f, post nuclear sculpture: g. central and literal teeth: A, marginal teeth, Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 221 222 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 147. Rotacharopa densilamellata sp. noy. Cunningham's Gap N.P., SEQ, QMMO10993, QMMO12676, paratypes. a, genitalia; b, details of penis interior; c, hermaphroditic duct and carrefour; d, talon; e, ovotestis: f, pallial cavity. Scale lines as marked. SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 223 25°S 26°S 27°S 28°S 29°S I54°E 153°E 151°E 152°E PIG. 148. Distribution of Rolacharepa densilamel- lata sp, hov. 22.10). Microsculpture (Fig. 146) of very fine radial riblets, 6-10 between each pair of major ribs, and equally fine spiral cords, which are raised at their intersection with the micraradials producing a beaded effect. Umbilicus (Fig. 146d) narrow, U-shaped, diameter 0.35-(,51 mm (mean 0.41 mm). D/U ratio 4.08-5,27 (mean 4.93), Sutures impressed. Whorls shouldered above and rounded below the periphery (Fig. 146c). Aperture roundly lunate. Lip simple, columella slightly deflected toward umbilicus. Parietal callus strongly developed. Colour light-brown. Based on 14 measured adults. Epiphallus internally with transverse pilasters, entering penis apically through a simple pore (Fig, 147b). Penial retractor muscle (Fig. 147a) inserted on epiphallus at the point of reflexion. Penis (Fig. 147b) thin walled, with a very long preputial extension (Fig. 147a). internally with a simple, apical epiphallic entrance, and strongly pustulose walls; pustules arranged in vague lon- vitudinal rows (Fig, 147b). Radula (Fig. 146g,h) typical. Based on 4 dissected specimens (QMMO.- 10993, QMMO12676). RANGE AND HABITAT Disjunct distribution in cool subtropical notophyll vine forests of the Great Dividing Range in southern Queensland, and the warmer subtropical notophyll vine forests of the D’- Aguilar, Conondale and Blackall Ranges, SEQ. It lives under logs. REMARKS R. densilamellaia has a more elevated spire than either of its congeners and the longer prepu- tial tube may be asecondary modification related to this height increase, There appears to be considerable interpopula- tional variation in the shells of R. denstlamellata but much of the available material is subadult and not suitable for statistical analysis. EXTRALIMITAL RELATIONSHIPS Attempts to relate the Charopidae reviewed in this study to extralimital charopids remain tenta- tive because of the lack of knowledge about some of the major regional charopid faunas. Solem (J983) reviewed the Pacific Basin species but only briefly mentioned the large radiations of New Caledonia, Lord Howe Island and Norfolk Island, Clima (1969, 1970, 197], 1978, 1980, 1981, 1983) reviewed a large num- ber of New Zealand species, but omitted impor- tant comparative data on shell sculpture and anatomy. Additionally, a large number of New Zealand charopids are still undescribed (Solem, Climo and Roscoe, 1981, p, 454). Locally. the Australian charopid fauna has thus far received scant atlention with only Solem (1984), Smith and Kershaw (1985) and more recently Stanisic (1987) presenting significant detail on a handful of species, Nevertheless broad comparative ob- servations are possible, SHELI. PATTERNS The extent of conchological variation in Australian species is similar to that in New Zealand charopids but contrasts with the relative stability Shown by Pacific Basin laxa, With the exception of Micracharepu momnula Solem, 1983 and the widespread Discacharopa aperta (MGllendorff, 1888), apical sculpture in Pacific Island species consists of spiral cords. Patterns of apical sculpture in New Zealand species may be spiral, radial, reticulate, smooth ot otherwise modified (Climo, 1969b, 1970, 1981, 1983), Among species examined in {his study, a combination of curved radial ribs and ape | weouker spiral vords is Common. Spiral aptesl sculpture, dominant among Pacific Island species, is present only in Sctemedea, Onr phaleropa varicasa and the extralimitally re- lated Siyploved ptensa. The pitted apical sculprure of Bionplialopa, Nautilirapa omicran and Gyracachlea curtisi- ama (Pigs 60e, 63d, 34e, S7c) has not been reported for Pacilie Island or New Zealand Charopidae but is seen in the primarily Papua New Guinean Pilshrvchurepa, Solem (1984) recorded this sculptural feature for Pilsbrycharopa timidus (Odhner, 1917) from the Northern Territory, Australia, ‘These taxa may be distantly related bul considering that this sculpture is derived from the more common reticulate pattern (see discussion for Gyro- eorlilea eurtisiana), it is likely ta have developed independently in the New Guinean genous, The pitted upical sculpture seen in Nvatrea murphy, Letomola contortus, Lenwebbia protoscrobicul- ata and Hedleyoconche (Figs 9f, 23c, 26d, 29g, 67d) is grossly Similarto that ofthe New Zealand Paracharopa Climo, 1983. In these cases how- ever, the macroscopic similarity has quite dif- ferent underlying microstructure which correlates with major anatomical differences. Apical sculpture of Raracharopa is identical to that in the Pacific Basin Microchuropa mimula Solem, 1983 and North American Radiodiscus millecastarus Pilsbry and Ferriss, 1906 although whorl numbers and coiling pattern indicate im- portant regional differences in these rotadiscines. The postnuclear sculptures of Australian, New Zealand and Pacific [sland species share similarities which are probably convergent and related to sharing a common niche. However. in contrast with the conservatism of Pacific Basin species, the Australian charopids display the same rich range of variation seen in the New Zealand fauna. Sculptural reduction, common among New Zealand and Australian species, is rare in Pacific Island charopids. Comparison of rib counts of Australian (Table 1) and Pacific Island species (Solem, 1983. Table |) shows that the median rib count in the latter is greater. [tis premature to say whether this will prove to be a basic difference between these hwo charopid faunis, The most unusual postnuclear sculptural ele- ments amung Australian subtropical Charopidac are the incised spiral lines of Ngairea. Leaweh- Ata imd ir lesserextent Mussonnula, This feature, which is more typical of helicarionids, is con- fined to the group of genera with incomplete MEALOTRS OF THE QUEENSLAND MUSEUM seeuridary ureters. No equivuledt word iter iis been reported in anv extralimital charopids. Shell shape of Australian and New Zealand species ts mure diverse (han thal of Pacific Lsland species which, with a single exception. are a monophyletic assemblage spread over anumber of islands (Solem, 1983). It appears that rhe Australian and New Zealand charopids have had a More complex evolutionary history involving many radiations und extinctions under condi- tions of high intra-site diversity where competi- tion for food and space has led to considerable variation in shell form. Consequently shell elevation and associated umbilical closure js a more frequent occurrence in Australian and New Zealand charopids than in Pacitic Island species. Accordingly, Solem and Climo (1985) shawed that development of a keel, periostracal proces- ses and shell colour in New Zealand species are sile-preference cortelated and not based on monophyletic assemblages. These authors ulso found that inthe development of large sympatric snail communities evolution appears to favour an increase in diversity of shell shape. In Such situations convergences will occur and are high- lighted jn the Australian-New Zealand species pairs of Hedlevoconeha delia - Serplia kev (Gray, 1853) and Se/omedea seticostata - Suteria ide (Gray, 1850), Barrier formation is extensive among Pacific Island and New Zealand Charopidae but occurs to a lesser extent in Australian subtropical taxa, Solem (1973; 1983. p. 15) discussed the polyphyletic origin of barriers in Pacifie Basin species und Climo (1978) provided evidence for olyphyletic origin in New Zealand species, Che form, number and microsculpture (Figs 78e, Std) of the barriers in Rhophadon make them comparable to the barriers in Pacific [sland taxa such as Semperdon Solem, 1983, and New Zealand Prychadon Ancey. 1888. However the reduced structures of Lefomola contartus (Fig. HSa.b) wre less easily related to Pacific taxa, being grossly similar to the barriers seen in New Zealand heetola tredale, 1915 (Clima, 1978), ANATOMICAL PATTERNS Pullial configurations of species reviewed are most similar to those of New Zealand Charopidae. All endemic Pacific Island species possess a Kidney in which the lobes.are subequal, equal or the rectal lobe is much longer than the pericardial lobe (Solem, 1983). In contrast New Zealand species for which data is available (Climo, 1980, 1983: Solem, 1983). have pericar SYSTEMATICS OF AUSTRALIAN CHAROPIDAE dial longer than rectal lobe. Letomola contortus, Egilomen cochlidium, E. globosa and Sinployea intensa have equal to subequal kidney lobes (Figs 69g, 129d, 132f, 138f) while the remainder have the pericardial lobe much longer than the rectal lobe. An incomplete secondary ureter was pre- viously recorded in the Charopinae only in Oreokera cumulus (Odhner, 1917) and O. nim- bus Stanisic, 1987 from north Queensland, and Amphidoxa marmorella (Pfeiffer, 1850) from Juan Fernandez. This is a primitive condition in the Charopidae (Solem, 1983; Stanisic 1987). Its presence in genera such as Hedleyoconcha, Setomedea and Ngairea, which all show typical- ly disjunct relict distributions, is not surprising, and a similar condition in southern Australian and New Zealand species would not be unex- pected. Shortened secondary ureters have been reported for some extralimital Rotadiscinae (Baker, 1927) but were not present in rotadiscine species reviewed here. Intrusive mantle-gland tissue was noted in the majority of species examined. This structure is rare in Pacific Island species and present only in Graeffdon graeffei (Mousson, 1869) and Semperdon xyleborus (Solem, 1983). Extent of its occurrence in New Zealand species is not known but it has been recorded for Charopa coma (Gray, 1843) and Phenacohelix pilula (Reeve, 1852). In local species the ovotestis usually consists of two clumps of palmately clavate to finger-like lobes of alveoli oriented parallel to the plane of coiling and embedded in the apical whorls of the digestive gland. Occasionally lobes assume a right angle orientation and may vary in size and number. All Pacific Island species have ovotestis lobes parallel to the plane of coiling. Informa- tion on New Zealand species is scarce although both parallel and right angle orientation have been reported. Stanisic (1987) considered that a tight angle orientation of the ovotestis was a primitive character. However, from this study it appears that ovotestis orientation is related to shell height. Hedleyoconcha, Ngairea and Len- webbia have elevated shells and right-angle orientation of the ovotestis. In contrast, Setomedea, which possesses a primitive pallial configuration, has a less elevated shell and the typical parallel orientation. Gyrocochlea vinitincta has a biconcave shell, with secondarily enlarged body whorls, and has the ovotestis oriented at right angles. Number and size of clumps are variable and probably related to variability in whorl count and shell size. Ngairea murphyi and Hedleyoconcha delta have high whorl counts and multiple clumps of ovotestis (Figs 10d, 24e) while the small Sinployea intensa and Letomola contortus have single bilobed clumps (Figs 69c, 138e). Among Pacific Island and New Zealand species the ‘two clump’ condition is typical. Some Pacific Island Semperdon Solem, 1983 and Sinployea Solem, 1983 show reduction to a single clump but there are no examples of the increase in clumps recorded in some Australian and New Zealand species. The penis of Pacific Island species is pear- shaped with the penial retractor muscle inserting on the penis head, or epiphallus, very near to the epiphallus-penis junction. The penis lacks a sheath, and internally, has a basic pattern of apical verge, circular ridges and a pocket stimulator, Among Australian subtropical taxa verges are rarely present, penial collars are con- fined to a handful of genera and a true pocket stimulator is seen only in Nautiliropa omicron (Fig. 65c). Instead there is a simple, apical (oc- casionally subapical) entrance of the epiphallus and few to many longitudinal pilasters which may be modified under conditions of sympatry. The penes of Hedleyoconcha and Ngairea have an apical muscular collar (Figs 4b, 10b, 13b, 16b, 24c, 27b) which is comparable with that of the New Zealand Phenacohelix pilula (Reeve, 1852) (Solem, 1983, p. 28). Addition- ally the large fleshy penial pilasters of Cralopa are similar to those of the New Zealand Cavellia sterkiana (Suter, 1891) and C. colensoi (Suter, 1891) (Climo, 1969b). According to Climo (1980, 1983) Pulch- ridomus, Aeschrodomus, Charopa and Phenaco- charopa have a constricting ‘sphincter’ dividing the penis into distinctly sculptured upper and lower chambers, A similar pattern is seen in Elsothera sericatula and E. genithecata (Figs 105e, 109b). Most Australian, New Zealand and Pacific Island species have a well defined epiphallus. However, some New Zealand species lack an externally differentiated epiphallus. A com- parable Australian species is Setomedea janae in which the epiphallus is partially incorporated into the penial complex (Fig. 39c). RADULA The typical charopid radula has tricuspid central and lateral teeth (usually the central tooth is slightly smaller) with long lanceolate an mesocone and short pointed ectocones. The marginal teeth are multicuspid and in many in- stances 4 transitional zone of lateromarginal teeth is present. This pattem is the rule in Pacific {sland and. most Australian subtropical species. Notable local exceptions are Egi/omen globosa, E, cochlidium (Figs 128h, 131h), Ompha/aropa varicosa and Letomola coniertus (Figs 134h. 68e) which have reduced central teeth, This latter condition is common in New Zealand species (Climo, 1969b, 1970). The broadly triangular mesocone and bicuspid laterals of Neairea (Figs 3f, Gd, 12d, [Sd) are similar to those of the New Zealand Phenacohielix (Cumber, 1961). Significantly both are arboreal to semi-arboreal. However, presence of anterior flares on teeth in groups such as Ngairea and Hedlevecancha has no known correlative in either New Zealand or Pacific Island taxa. Nautlliropa amicron has slender hook-like teeth and V-shaped row arrangement (Figs 63c,f). This is 2 major departure from other subtropical species and also has no equivalent among Pacific Island or New Zealand Charopidae. NV, omicron is sympatne with the conchologically convergent Gyrocachlea and the unusual radula may be a local feature related (o intergeneric sympatry. In summary, Australian subtropical Charopidae display a number of conchological and anatomical features which suggest relation- ships with New Zealand charopids but also indi- cate evolution in isolation for some considerable period, At the same time development under similar environmental conditions has led to many convergences in shell features among specics of these two regional charopid faunas. Analysis of southern Australian:species will be critical in Understanding the full extent of pos- sible relationships. PHYLOGENY “lhe fifty species of Charopidae reviewed dis- play a Wide variety of anatomical and con- chological pallerns, yel hey represent only a small subset of an extensive Australian charopid fauna, The prolific Tasmanian and. Victorian faunas, extensive radiations in mideast and northeast Queensland, and a further 60-70 poor- ly known or undescribed subtropical species are still to be reviewed, Hence the following discus- sion fs preliminary, Negairea, Hedleyoconcha, Mussonula, Len- MEMOIRS OF THE QUEENSLAND MUSEUM webbia and Setomedea ave incomplete secon- dary urelers and possess a number of con- chological specialisations (incised spiral lines, broad spiral grooves, periostracal setae, punctate apical sculpture) Which isolate them from the genera with the more typical charopid pallial configuration in which the secondary ureter is complete. They probably had a contmon ances- tor, Their penes have a tVpical internal pattern of apical collar and longitudinal pilasters. This is well illustrated in Ngairea and Hed- leyeconcha. Together with Mussonula and Len- webhia these genera are further characterised by clevated shells with very narrow umbilici. An exception 1s N, murphyi which has secondarily narrowed whorls and a slightly more open um- bilicus. Lenwebbia can be derived from Ngairea through modification of shell sculpture, penis and kidney, and shares the unusual feature of incised spiral lines with Ngairea and Mussonula. Setomedea appears to represent an attempt at habitat specialisation among the taxa with in- complete secondary ureters, The relatively small size. peculiar coiling of |he primary tireter, and interspecific variability In whorl numbers and shell coiling pattern, are inconsistent with other genera in this group, Unlike Ngairea and Hed- levaconeha fand probably Mussonula), Setamedea is nol arboreal or semi-arboreal, It usually lives in moist rotting logs and the periostracal sctae-are probably an adaptation for preventing dirt accumulation on the shell in this comparatively wet microhabital, Among the more generalised radially-ribbed groups. Cralopa and Elsethera stand apart in having altered penial and epiphallic complexes, closed umbilici and conspicuous apertural sinuses. This latter feature is relatively rare in Australian charopids and suggests & possible relationship with New Zealand particularly through Cavellia Iredale, 1915. Cralopa and Elsothera are probably more closely related to each other than to other groups reviewed here. In spite of size differences, Biomphalopa and Gyroceciiea show structural consistencies in shell and anatomy which suggest common an- cestry, Gyrececilea is an evolutionary experi- ment in whorl size increase which reaches its extreme in G. vinisinetie. Nauriliropa with pitted apical sculpture, depressed spire, and strong radial ribbing may be related to the Gyrocochlea—Biomphalopa com- plex even though radular morphology, penial structure, shell coiling pattern and microsculp- jure are against close relationship, SYSTEMATICS OF AUSTRALIAN CHAROPIDAE The highly modified Letomola contortus is possibly a derivative of the Gyrocochlea-level of specialisation which has undergone size and shell sculpture reduction associated with a shift in microhabitat to life on rock surfaces. En- larged, bulbous protoconch and elongate genitalia indicate descent from a large ancestor whose organ systems are consistent with the elongate Gyrocochlea pattern. Barrier forma- tion and sulcus development in L. contortus can be correlated with change to small size and al- tered lifestyle. The bilobed kidney (with acutely angled ureteric arms) is probably a secondary modification directly attributable to reduction in whorl count. Rhophodon also has apertural lamellae but these are independently derived from those of Letomola. The coiling pattern, whorl profile and strong radial sculpture suggest affinities with Gyrocochlea lineage. However the apical sculp- ture does not. Coenocharopa is diverse in subtropical rain- forests. The various species lead relatively specialised lifestyles among the dirt particles of the litter, on rock surfaces, or among moss on logs. Their shells have undergone reduction in primary calcified sculpture and developed fine deciduous periostracal ribs and extensions. Radial apical sculpture, enlarged penial com- plexes and expanded epiphallic pore suggest dis- tant relationship with the Cralopa/Elsothera complex. Problems associated with Discocharopa have been discussed in the systematic section. In essence the subtropical Discocharopa may be a a locally derived group. Apical sculpture indi- cates possible relationships to the Cralopa-El- sothera group. Egilomen is represented by the depressed, widely umbilicate and strongly ribbed E. cochlidium and the globose, almost imperforate, more heavily ribbed E. globosa. The bilobed kidney (without evidence of whorl number reduction) suggests a quite separate derivation to other radially-ribbed genera reviewed herein. Omphaloropa varicosa with spirally lirate apex, wide umbilicus and large tubular penial verge does not appear to have any close equivalent among other subtropical species. Another species with spirally lirate protoconch, Sinployea intensa, represents a remnant of a more widespread Pacific group. Epiphallic and penial modifications distin- guish Rotacharopa while the peculiar apical sculpture relates the genus to the Pacific Island Microcharopa and North American Radiodiscus. Knowledge of Australian Charopidae is still at a rudimentary level and reflected by the mosaic nature of morphological trends outlined above. Production of a formal generic phylogeny would be a futile exercise at this stage. Rather I have proposed a scheme of hypothesised relationships (Fig.149) which summarises the preceding dis- cussion and information given elsewhere in this study. It identifies the probable multiple origins of the fauna and outlines major trends discussed herein. PAUP A preliminary computer-based cladistic analysis using PAUP was carried out. Seventy- five characters (42 anatomical, 33 conchologi- cal) (Appendix) were chosen and ordered. The resultant cladogram (Fig. 150) shows that taxa with highly differentiated shells such as Coenocharopa and Omphaloropa are distin- guished from the more generalised groups such as Gyrocochlea and Rhophodon. However, there are also some unusual associations where a high degree of conchological convergence is involved, viz. Ngairea levicostata and Mus- sonula verax ; N. murphyi and Hedleyoconcha delta. Egilomen globosa is associated with the Rotadiscinae even though the latter has marked- ly different sculptural and anatomical features. Elsothera and Cralopa are associated with taxa lacking a complete secondary ureter and the conchologically specialised Setomedea is separated from other species lacking a complete secondary ureter. Gyrocochlea, Biomphalopa and Nautiliropa were grouped together, and the highly modified Letomola contortus was placed near these genera. BIOGEOGRAPHY The Australian charopid fauna possibly con- sists of 250-300 species but prior to the present study, which introduces 27 species, only 110 species level taxa had been named. Hence, with probably less than half the available species described, and an even smaller number subjected to detailed analysis, it is premature to be confi- dent about understanding their biogeography. The treatment below provides a synoptic review of climatic, geologic and vegetational changes in the east coast region during the Tertiary; brief discussion of charopid distribution patterns; and PP MEMOIRS OF THE QUBENSLAND MUSEUM synthesis placing the taxa of this study into broader biogeographic context. BACKGROUND In subtropical and tropical eastern Australia distribution of land snails and rainforest arc linked, In the Charopidae, which is acknow- edged to be a Gondwanan family (Van Bruggen, 1980), this relationship probably had its origins in (he Late Cretaceous when mesic forests were more widely spread than at present. Since the Cretaceous few Charopidae have made the tran- silion lo drier sclerophyll forest, in spite of potent enVironmental change which has fayoured the development of xerophilic communities. This indicates a strong ecological bond between these tiny snails and closed forests which is probably due to convergent habitat preference, The ecological factors which favour development and. persistence of rainforests ¢.¢. moisture and eutraphic soils, are also those which favour the sucecss of Lerrestrial molluses, For sniils, rain- forest also provides shelter and food. These conclusions are not very different from those of Solem, Climo and Roscoe (1941) who found that ministurc atid spatial quality were significant fac- lors in the outta of snail species in the Manukau Peninsula area of New Zealand, and Boyeott (1934), who considered that shelter, lime and moisture were key factors in determin- ing local occurrences of British land molluscs. Australia separated [rom Gondwanaland inthe carly Tertiary reaching its present position at the edge of the Asian plate in the mid-Miocene (Kemp, 1981; Kemp and Galloway, 1981). In the early Tertiary, most of Australia would have resided within a weak band of wind circulation and the surrounding warm seas would have resulled im Widespread precipitation. But with opening Of the southern ocean and drift northward into lower latitudes, significant chan- ges in climute occurred, Circum-polar circulia- tion significantly reduced heat transport between the eyuator and pole resulting in latitudial stratification of water temperatures. Conse- quently around the middle of the Miocene, the sea surface cooled and precipitation on land declined. From that lime aridity has been a domsnant factor in Australia’s climate with wet- ler phases occurring only briefly, Rainforest communities, Widespread at the beginning of the Tertiary, retreated to refugia and were replaced by u xeric adapted biota. However, in eastern Australia, two geological events contributed to the maintenance of mesic communities in the wake of the arid phases which elsewhere resulted in the removal of the widespread rainforest flora (Martin, 1978), Firstly, episodic uplift of the eastern highlands began in the Paleocene with major elevation occurring in the late Miocene and Phocene. Secondly there was widespread igneous activity in eastern Australia along and adjacent to the eastern highlands (Wellman and MacDougall, 1974), In northern NSW vyul- canism continued inta the mid-Miocene while in SEQ peak activity occurred in the Oligocene and Plio-Pleistocene, The eastern highlands ensured orogr apnic rainfall and enabled species to radiate alliludinally, while indirectly, vulcanism provided favourable soil-water conditions in a basically dry infertile continent. The fact that most land snails in eastern Australia now occur in ‘rainforest’ indicates that the persistence of moist refugia was critical to (heir survival in this region, The tmajor refuges during these dry potions were in the moist uplands, and in owlands along major drainage lines— areas which were not large, but wel. Climatic changes in the late Quaternary were similar in amplitude to those experienced in the Miocene and Pliocene bul occurred in more tapid succession, Galloway and Kemp (1981) concluded that these changes must have pluced considerable stress on montane and coustal en- vironments and that modern communitics. in these situations are recen! phenomena consisting of biota Which have survived in isolated refuges, Recent studies by Kershaw (1980, 1981) on pollen samples from crater lakes on the Atherton Tableland, confirmed thut the climatic fluetus- tions of the late Tertiary had dramatic effeets on the Australian vegetation. Kershaw’s results showed that between 83,500 and 79,000 BP, complex rainforest covered the Atherton Tableland under rainfall much higher than today, These conditions persisted until about 38.000 BP when a major change occurred and wet-adapted araucurian forests of the Atherton Tableland were replaced by drier communities charac- terised by Evecalyptus and Casuarina, These dominated the area until B-10,000 BP when rain- forest again emerged from retreats. Kershaw (1980, 1981) was further able to demonstrate that these local, environmental changes could be ap- plied to other areas along the east coast. The palynological studies of Kershaw (1981) FIG, 149. Hypothesised relationships of charopid genera revieweel in Uns study 229 SYSTEMATICS OF AUSTRALIAN CHAROPIDAE atqetaea ‘ajduwis ‘aunzdinos Leorde aye[Noiqau aunqdinos uea_onuzsod daqaun Ayepuodsas azaldwoo 4e_ pod Leotde yqim ALLensn stuad xa[duiod aungdpnos aeaponuzsod daqyaun Axepuosas ayaldwoout * 9 ¢ aunzdinos ayenuazze fpazeaa aunydtnos [eoide LeLpeu * s_eatds pue st B4NzyXlui aunja Lenbaqns saqo, Aauply * LeLbizsaa AL ysow aqo, Aauply LepIe aeqyas Leoeuysolued * aunzdinos Leoide jeuids + W4AdSWOLAS [epowig tazeLNoILqze4 aunjdpnss ava_onuzsod aanjdtnas Leoide paqzid * proyso3 Lays * YHONOQO Ad 103 lpautsap Lam sn__eydida sql4 [eLper pLoq NAWO 1194 aunzdp nos Leoide azelnoizau * quauLwoud aunydinos Lerpea SLeLpeu pape_q ao steutds pastour YTLM aunqdpnos seal anuzsod suoLsuayxa [eoeuqsoluad * paonpau aunzdpnos * YdOUVHOONSOD sLeLpea yuauLwoud joe Lawe, Leanquade * LLews Avan LLays * NOGOHdOHU paljsipow uo s_peitds yeam YzLM sLeLpea go aunqdinos jeorde proysouqqns [Lays | VAUTVON Lenbaqns sago, Aauply aellawe, peunquade ajzeuauabap aunqdynas * LLews Awan [Lays * WIOWOLS1 abueuo yyhtag AaupLy aunydtnos leotde pazqid * asogobqns LLaus * YWIGdSMN3 1 S[PLpea aye, pue s[eards Ajaea Jo aungdinos eorde - PLoysoaz | Leys * WIANOSSNW SnoLLEquin apLm AuoA aynuLuw [Lays YdOUVHIOISTIG pidsnorg yjaaz Leurbaew uaajjzed uaqzse_id xapduoo HuLpuassap you [4oyM zseL YWdOUTTILAVN Sql LeLpe4a papmoss pazeLjuasassip Leordeqns uorzount Ai yeam snyleydida * sn_Leydida/suauajap sea sqla LeLpea plog * papLAtp stuad abuel ALPALZeL aU WdO Wud YWadHLOS 13 peorde jeatds ° abuar ueln sqla LeLped paseds-A|apLm * YdOUO IWHdWO Sqtu [eLpes papmoud * abuel AL@aAtqzelau * pidsnoizinw yzaa} [euLbuew 4 dayseid Letuad ayGurs * WdOIWHdWOTd xa_dwoo petuad/oL,leydida parytpow xa[dwod aunqdinos Leorde aye_ndiqau aunqdpnds aeayonuzsod * 4aqyaun Asepuodas azaldwos - Lenbagns saqo, Aauply * suaysetid xapdwoo sqlu [eLpea papmosd WAA0 IdNIS qny ° Aut} snoepLqun * Lieus Auaa Sasoqol6 Lays * WdOuWHOVLOY BAROUOILG * Bulpuadsap [4oym yse, 7 suayse_id peruad atdurs Auew * YA THOOIONAD 2) and Murtin (1981) demonstrated the drastic ef. feets of mid to late Tertiary climate on mesic communities and exposed some important prin- ciples pertinent to east coast biogeography. Firstly, areas and their faunas have been isolated on a number of occasions thus promoting al- lopatric speciation, and subsequently unified to allow for dispersal and the evolution of sym- patric congeneric pairs. Secondly, cerlain ele- ments of the biota found in the “modern” northern Australia have been present in Australia for a lang time and many of (hose taxa presently confined (u (he north were present in south-east- ern Australia when it Was adjacent to Antarctica. Thirdly, some species in northern Australia have become extinct in fairly recent times. These principles have particular application toa low vagility Gondwanan group of organisms such as the charopid fand snails. Webb (1959, 1968) expressed the resulis of past environmental changes through a present day classification of structurally based rainforest types. Ina biogeographic synthesis, Webb and Tracey (1981) demonstrated that the popular concept of tropical and subtropical rainforest encompasses a complex mosaic of community types cach reflecting a past scrics of climatic change. This division of the "ramforest massif" into regions and provinces associated with present climatic regimes, which at the same lime reflected quite different histories of climatic change. provides a practical {ramework for inter- preting the impact of these changes inthe context of the Fragmentation and isolation of mesic com- munities, Ata regional level the results indicate that the cooler forests of the southeast (cool temperate lo subtropical) have been separated from the warmer forests of the northeast (tropi- cal}, probably since the late Miocene aridity phases. A dry corridor between Gladstone and St Lawrence, geographically distinguished by a lack of coastal and subeoastal highlands, ensured that the biotas of the two regions developed in isolation While a dry corridor between Townsville and Bowen isolated the Eungella rainfarests from the main massif of northern rainforest. At a provincial level, temperate forests of the cooler Tasmanian and Victorian regions are segregated from the more northern warm temperate, and coo) tn warm subtropical forest while a third drier subtropical forest type occurs subcoastally in southern Queensland. Considering the close ecological ties between FIG. 150. Consensus (ree derived from PAUP analysis. MEMOIRS OF THE QUFENSLAND MUSEUM Charopidae and closed forests, Webb's. rain- forest classification makes it possible to relate past climatic and geological events to present day charopid distribution. CHAROPID DISTRIBUTION Atthe family level distribution of Charopidae in Australia is basically cast coast. Species ore found in moist cool temperate forest of the south- east region (Smith and Kershaw, 1979; Smith and Kershaw, 1985), warm temperate and sub- tropical forests of northern NSW and SEQ, in the structurally subtropical forests near Mackay, MEQ, and the tropical coastal forests between Ingham and Cooktown, NEQ, A small but sig- nificunt radiation of charopids is also present in southwest Australia (Solem, 1983), several species exist across the north of the continent (Solem, 1984), and a presumably relict species 'Pillomena™ aenmula (Tale. 1894), is found in the renmanl mesophylic forest of Central Australia, AJl indications are that the distribution of the group in Australia Was much more extensive jn the past. Of the 50 species reviewed herem 32 are en- demic to the main area of subtropical rainforest between Dorriga, NSW (30°30'S) and Rock- hampton, MEQ (23°30'S), Three species viz, Hedleyocancha delta, Coenacharopa multi- radiata and Cralapa sfroudensis extend further south, while 14 species occur outside this core region, Discocharepa aperta trom the drier cain- forests of SEQ ranges to northern and central Australia, the Bismarcks, Philippines, Indonesia, New Hebrides, Kermadec, Fiji, Samoa, Austral and Society Islands (Solem, 1983, pp. 74-81) The small, semiarboreal Sinployea intense has a long, narrow, coastal distribution which encom- passes a number of sublittoral rainforest patches- Wide ranging species also include the litter dwelling Omphaloropa varicosa which is found from the McPherson Ranges on the NSW/Qld border, to west of Rockhampton, MEQ; Setomedea seticostata and Ngairea dorrigoensis which range fram Dorrigo, NSW, to just north of Brisbane, SEQ; and the litter dwelling Coenocharopa parvicestata with a range from Dorrtigo, NSW, to just north of Gympic, SEQ. The north-south linear ranges of these species ure between 350-700 km and are the upper ex- iremes where the majority of species have ranges of less than 200km, However in general the SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 231 yessabahensis sordidus macromphala alata parvicostata multiradiata levicostata verax ew corticicola dorrigoensis canaliculata murphyi fal ey Fes ailaketoae fallax protoscrobiculata nautilodea sericatatula genithecata paucilamellata [| convoluta vinitincta curtisiana omicron recava concinna cuntortus peregrinus kempseyensis consobrinus minulissimus colmani elizabethae bairnsdalensis aperta globosa annabelli kessneri densilamellata kaputarensis stroudensis carlessi nudicostata janae monteithi seticostata cochlidium intensa varicosa ANCESTOR to "ed bat ranges do not approach Solem’s predicted value for all land snails of less than 50 km (Solem, 1984. p. 14). With the exception of Rhaphodon colmant, whose distfibution is considered to be poorly known the smallest linear ranges of subtropical endemic specics are those of Rhaphodon elfeeberhae and Setomedew nudicastata which Oceur in the relatively small and unusual moist subtropical refugium in the Bobby Range, south- west of Gladstone, SEQ, These examples sug- gest thal species ranges may be determined by the size uf particular rainforest patches and hence subject ta considerable pressure from changing climatic conditions. Coenocharopa yessabahensts, Rhophodan Kempseyensis and Letamala contortus (rom the Yessabah-Currai limestone serits, west of Kempsey, NSW, are further examples of environmentally restricted taxa. Coenoeharopa has four endemic, often over- lapping Species in subtropical forests with com- paratively large linear ranges (180-486 km) indicating that the genus has undergone a long period of environmental sifting. tn Gyracachlea, linear ranges are relatively short (54-108 km) and patterns of overlap are minimal with only Gyrococklea vinitineta and G. tancr/amellara showing minor sympatric occur- rence. It is probable that the Gyrecocklea radia- tion is a relatively recent phenomenon in the subtropical region. Geographic distribution within a genus is generally north-south, RAophedon, witha single Species in the southeast region and six species from Kempsey, NSW, to the Bobby Range, SEQ, is illustrative. Semnedea and Hed- leyeconeha have more striking disjunct north- south distributions with relict species in central and north Queensland. Ngairea has its main radiation in the subtropical forests but also has an Outlier species in the Illawarra region of southern NSW, The southern limits of Elsothera are not known but ils occurrence in the subtropi- cal fures| is also considered to be relict. Asa Tule, genera tend to have relicts in the north rather than vice versa indicating that some genera which are present in the south were also common in the north during cooler times, A major exception to the typical north-south pattern ts Cralopa which has a single widespread species, C. srraudensts, in the coastal and sub- coustal sublropical rainforests; a closely related species, C. kKaputurensis, on Mt Kaputar, West of Narrabri, NSW, and a further widespread, but MEMOIRS OF THE OUEFENSI AND MUSEUM Beugraphically disyunet species, C. carlessi, on the limestone outcrops of the Great Dividing Range. Unlike other genera now confined to the easl, Cralopu has been able to survive in largely snail depauperate areas further west, Although patterns of geographic distribution are usually north-south, large wet ureas such as the McPherson Ranges provide sufficient width in the rainforest massif to allow for localised speciation in un cast-west direction, ¢.g. Ngairea and Gyrocochlea, There is limited evidence tor altitudinal dif- ferentiation. Evilomen cochlidium and Coendcharepa alata show preference for mon- tane habitats in contrast to the foothill/lowland distribulion of their congeners. Neairea cor- licicala is microsympatrie with WV. dorrigoensis in the Great Dividing Range, and extends to the foothills whereas N, dorrigoensis occurs anly in the highlands. Other examples are seen in Gyrocochlea and Rhaphodon. Coherent distribution patterns among closely related genera are less obvious but this 1s probab- ly due to gaps in phyletic knowledge. Where patterns emerge they appear to be similar to those discussed above, Gyrococh!ea which is an endemic subtropical group has ils nearest rela- tive, Biomphalopa, in the rainforests of central and north Queensland. In contrast the closely related Mussonula, Lenwebbia and Negairea have overlapping distributions in SEQ, PATHWAYS OF DIFFFRENTIATION 1 consider that the environmental changes as- sociated with mid to late Tertiary aridity were (he mayor factors affecting the present makeup and distribution of the Australian Charopidae. Two levels of chinges are seen as particularly sig- nificant- those which produced generic differen- ation and major mtrageneric disjunctions, and those which produced fine grain distribution ad- justments within larger areas of rainforest. The first level probably dates from the onset of aridity inthe Miocene, while the second is most likely related to drastic short (erm climatic oscillations of the Quaternary, Th the north the formation of dry corridors between St Lawrence and Rockhampton, and Bowen and Townsville (Fiz, 1) correlate with the current disjunct distributions of Hedlevoconcha and Setomedea, Persumably these groups onee ranged along the entire moist coustil region from north Queensland to northern NSW. As mesic habitats retreated, the ancestral range be- came severely fragmenied, and mid-cast SYSTEMATICS OF AUSTRALIAN CHAROIIDAE Queensland and northeast Queensland popula- tions would have been isolated from each other ind from more southern subtropical rainforests, Arboreal Hedlevoconcha responded more dramatically disappearing from the notophyll vine forests of mid-east Queensland, In the north, H. ailaketoae survived only in the al- titudinal refugia of Mt, Bellenden Ker, while in the south, persistence of suitable habitat enabled H. dela to flourish in a wide area of moist sublropical rainforest in northern New South Wales and southern Queensland. Setomedea would have endured similar en- vironmental pressures to Hedleyoacencha but, probably because of greater adaptability or peculiar microhabitat requirements, managed to maintain S, Janae in mid-east Queensland, and at least (wo other species in north Queensland. Inthe north Setontedea was mostlikely restricted to montane refugia at some time in the drier past, hut subsequently has been able to disperse into the foothills and lowlands (S. monteithi). In southern Queensland and northern New South Wales, 8. seticostata (Hedley, 1924) occupies upland regufia as well as lowland notophyll vine forests also indicating dispersal from high to low altitudes. in wetter times. Further examples al large-scale intrageneric disjunctions which can be related to mesic habitat regression include the separation of Biomphalopa concinna and B. recuva, and the isolation of Ngairea murphiye in \he Hlawarra region, southern New South Wales, from the main mass of Ngairee species further north, Creation of these major dry corridors probably also led to generic differentiation between north and south. This is best illustrated by the disjunct distribution of the closely related Gyrocochlea (southeast Queensland) and Biomphalopa (mid- eas! and north Queensland), In Southeast Queensland, regression of com- plex mesic habitats to upland refugia and their replacement by less complex microphyll vine forests in the lowlands between Gympie and Rockhampton, correlate with disjunctions in several groups. Rhophodan elizabethae, Gyro- cochlea curtisiana, Setomedea nudicostata and Neairea canaliculata occur within this region in wetter montane refugia and are gcographically isolated from congeners further south. This isolation has been severe in R. elizabethae and S. nudicostala Which are restricted to refugial notaphyll vine forests in the Bobby Range, SW of Miriam Vile, and NV, caneficulara which in- habits only the moist forests of the mountain summits between Colosseum Creek and Mt Lar- com. In contrast G. curtisiana is found in both moist and dry forests, Il probably survived climatic attrition in scattered refugia and sub- sequently dispersed into the dricr microphyll Vine forests and [ringing open forest, In com- parison with R. elizabethae, S. nudicostata and N. canaliculata, the latger G. curtestana has less specific microhabitat requirements, It has more gencralised pallial structures indicating that it may have had less difficulty in adapting to drier environments. The fragmented distribution of Rotacharopa is also probably due to these local environmental changes. Although Vealrea has not colonised these drier microphyll vine forests, Lenwebbia protescre- biculaia, which is considered to be a derivative of the Ngatrea group, has done so successfully. Within the main humid subtropical rainforest block, which includes the Border Ranges, Great Dividing Range (from Dorriga, NSW to the Mis- take Mountains, SEQ) and their foothills and lowlands, charopid evolution has been much more complex. The long-term stability of these subtropical forests was ensured by the favourable voleanic soils and above average rainfall, even at lower altitudes. Hence while there was fragmentation and retreat of moist forest into favourable retugia such as gully heads, riverine alluvia and mountain summits in dry periods, alternate periods of higher rainfall would have Jed to many episodes of expansion and amalgamation. In some areas such as the drier western edges of the Border Ranges and Great Divide, and in places where soil types Were poor, periods of lower rainfall did cause a shift invegetation type to drier microphyll vine torest. In higher peaches e.g. Lamington Plategu, ‘temperate’ rainforest persisted in selected refugia. These events would have provided cnn- siderable opportunity for local speciation, dis- persal and isolation. The results have been “fine-grain” speciation patterns and evolution of several endemic genera, The ancestral stock of Gyracochlea presumab- ly had a distribution which covered the uplands and lowlands of the castem Border Ranges. Fol- lowing climate-induced habitat fragmentation, the heavily ribbed Gyrocochlea vinitincta would have been isolated in higher parts of the Lamington Plateau from the sparsely ribbed forms of the foothills; and with the shift to drier forest tvpes in western areas, G. convoluta be- tame sopurated from more casterly &. paicfemelia whieh lives in the palm- 234 dominated forests around the Darlington Range. In wetter times, G. vinitincta dispersed into the range of G. paucilamellata. Similarly evolution of local Rhophodon probably involved frag- mentation of a wide ranging ancestral population into foothill (R. consobrinus) and upland (R. peregrinus) species. Subsequently R. peregrinus expanded its range to become sympatric with R. consobrinus. Aridity episodes probably isolated the weakly ribbed Ngairea corticicola in the subtropical notophyll vine forests of the uplands and foothills from the heavily ribbed stock in more restricted temperate montane refugia. In the western highlands of the Border Ranges the development of drier vine forests effectively iso- lated N. /evicostata from the widespread mon- tane species, N. dorrigoensis. As aridity phases were replaced by periods of higher rainfall, N. dorrigoensis was able to disperse from montane tefugia and gradually occupy the ranges of its two congeners. It is more than coincidental that the expansion of rainforest during wetter periods favoured those species restricted to moist montane refugia. These communities were able to expand into lower areas where the soil type was suitable, and into areas with less suitable soils where rainfall was adequate. On the other hand the lowland and foothill rainforest communities isolated in gully heads would have expanded to produce a more contiguous block, but would not have expanded to higher altitudes. In contrast drier rainforests were environmentally cornered, and consequent- ly, the charopids in them, limited in their dispersal e.g. N. levicostata and G. convoluta. The ability to relate past events of climatic, geological and vegetational changes to present- day distribution patterns in the Charopidae varies from group to group. In Coenocharopa, which contains six species with widely differing microhabitat preferences, these events have produced a very complex pattern of species over- lap. Indications are that the group has undergone a long period of environmental sifting that has blurred historical pathways. There is evidence for segregation of a mesic-adapted highland species (C. alata) and a species which became adapted to the more easterly warm moist notophyll vine forests of the foothills of the McPherson Ranges (C. parvicostata). In the north, C. macromphala and C. sordidus probably developed from a common ancestor In response to environmental change which isolated mesic communities in the D’Aguilar Range from the MEMOIRS OF THE QUEENSLAND MUSEUM araucarian forests of the Blackbutt Range and Mt. Mudlo region. Most likely more than one instance of dispersal has brought about the com- plex sympatric patterns evident today. Nautiliropa and Mussonula are endemic to this region and are probably recent local derivations of more generalised stock, viz. Gyracochlea and Ngairea respectively. Restriction of charopids to wetter coastal habitats during the latter half of the Tertiary did not necessarily mean complete extinction of the family from drier western, sclerophyll- dominated areas. Where conditions were favourable i.e. where moist refugia could exist, charopids persisted e.g. Cralopa. As mesic habitats retreated eastward in the face of increas- ing aridity, C. kaputarensis was isolated in the moist montane refugia of Mt Kaputar, near Nar- rabri, NSW, and a little further east, C. carlessi survived in scattered limestone refugia on the western edge of the Great Divide, albeit reduced to a series of cornered disjunct populations. Limestone habitats are extremely important refuge zones for land snails. Along the Great Divide, they typically support remnants of once more prevalent wet-adapted vegetation. The rocks trap available moisture in crevices and also provide protection from wildfires thus enabling small island communities, quite different from those in the surrounding open forest, to be main- tained. Land snails which inhabit these excep- tional refugia benefit from an ample supply of calcium. Because of long term isolation and need to adapt to a lifestyle among and on rocks, endemicity and specialization among snails is usually high, Limestones of the Yessabah-Car- rai block, west of Kempsey, are a case in point. Their isolation from the main subtropical rain- forest block, presumably due to mesic habitat regression, has resulted in separation of several charopid species from northern congeners e.g. Rhophodon kempseyensis and Coenocharopa yessabahensis. The monotypic Letomola, which may be a derivative of the Gyrocochlea lineage of subtropical charopids, is also endemic to these limestones. All these species have highly dif- ferentiated shells and habitat-related radular modifications. CONCLUSIONS Origins of the eastern Australian subtropical charopid fauna appear to be diverse. Groups with primitive pallial configurations such as Negairea, Hedleyoconcha and Setomedea are SYSTEMATICS OF AUSTRALIAN CHARGPIDAE probably remnants of older stock with origins in the widespread Gondwanan biota. This group seems to have suffered extensively from mesic habitat regression although Ngairea has managed to secondarily diversify in the subtropi- cal forests. Mussonu/a and Lenwebbia are most likely recent, local derivatives. Cralopa and El- sathera have shelland anatomical features relat- ing them ta New Zealand Charopidae and are probable descendants of southern based groups that have altenualed distributions in subtropical Tegions. Coenocharopa can be considered an early offshoot from this stem that radiated into diverse habitats in the subtropical region, Rhophadon, with widely separated species along the east coast is probably of southern origin representing an early attempt at habitat specialisation and like Coenocharopa and Neairea has secondarily diversified in the sub- tropical forests, Gyrocochlea, Biomphalopa, Nautilirepa and Letomola are possibly part of the same lineage as Rhophodon, bul more recent in origin. In the case of Letamola and Nautiliropa generic differentiation has been ac- companied by dramatic changes in conchologi- cal and anatomical features. Of other genera, Sinployea is a remnant of a larger Pacific Island radiation; Egilomen is of uncertain affinity but has anatomical features indicating only distant relationship with other subtropical groups; Omphaloropa has a conflict- ing combination of characters and unusual dis- tribution pattern which suggest that its telationships may be with species well to the north or south; Discocharapa remains problematic; and Rotacharopa is related \o Pacific Basin and north American species. ACKNOWLEDGEMENTS I owe special thanks to Mr Darryl Potter, Queensland Museum, for his assistance in many aspects of this project. I am most grateful to the Jate Dr Alan Solem, Field Museum of Natural History, Chicago, USA, for his critical com- ments and constant encouragement. For the Joan of material | thank staff of the Australian Museum, Sydney; Museum of Vic- totia, Melbourne; South Australian Museum, Adelaide; and British Museum of Natural His- tory, London. Other material was generously provided by Graeme Annabell, Vince Kessner, and Terry Carless. Numerous specimens used in this study were collected during field work funded by the ra tee Ley Australian Biological Resources Survey (1981— M3), } wish to thank the following staff of the Queensland Museum - Dr Robert Raven (Arach- nology) for his help with the SEM and computer analysis; Dr Geoff Monteith (Entomology) for providing material and important information on collecting sites; Ms Gudrun Sarneés (Technician, Queensland Museum) for help with lhe SEM: Mr Bruce Cowell and Mr Gary Cranitch (Photog, raphy) for their assistance with all aspects of photography, In the early stages of the project Miss Alison Green, formerly Technician, Queensland Muscum assisted with SEM and field work. For typing the manuscript, T am particularly grateful ta Mrs Peta Woodgate, Queensland Museum, This wark is based on a thesis which was submitted im fulfilment of the requirements for the degree of Doctor of Philosophy at the University of Queensland (1988) and I would like to thank Professor Barrie Jamieson, Depart- ment of Zoology, for his guidance and criticisms. Finally, | wish to thank a special friend, Jan, who has been most supportive during the course of this project. LITERATURE CITED ANCEY, C.F 1888. Nouvelles contributions malacologiques Bull. Soc. Malac, France, $: 341-375, BAKER, H,B, 1927. Minule Mexican land snails, Pree. Acad, Nat. Sci Phila,, 79, 223-246, pls. 15-20, BAUR, G.N 1959, Nature und distribution of rain- forests in New South Wales. Aust. J. Bol, 5: 199233, BISHOP. MUI. 1981. The bisveography and evolution of Australian land snails, Inv A. Keast (ed.), Ecologleal Biogeography of Australia, pp. 924— 954. W. Junk Publ, The Hague, Netherlands BOYCOTT, A.E. 1934. The habitats of land Mollusca in Britain. J. Eool,, 22: 1-38, BROADBENT, J, AND CLARK, S, (eds). 1976, 4 faunal survey of east Australian reinforeses. Australian Museum, Sydney. BURCH, J.B, 1976, Outline of classifications of Ausiralian terrestrial molluscs (native and intro- duced), J; Malav, Soe. Aust, 3: 127-156. CLIMO, F.M. 1969a, Classification of New Zealand Anonacea (Mollusca: Pulmonata), |, The higher classification. Rec. Dominion Mus, Wellingron, 6(12): 145-158, 5 figs. 236 MEMOIRS OF THE QUEENSLAND MUSEUM 1969b. Classification of New Zealand Arionacea (Mollusca: Pulmonata). II. A revision of Charopa Subgenus Ptychodon Ancey, 1888. Rec. Dominion Mus. Wellington, 6(14) : 175- 258, 34 figs., pls. 1-11. 1970. Classification of New Zealand Arionacea (Mollusca: Pulmonata). II]. A Revision of the genera Charopa Albers, 1860 (excluding sub- genus Prychodon Ancey, 1888), Phenacharopa Pilsbry, 1893, and Flammocharopa n. gen. (En- dodontidae: Endodontinae), Rec. Dominion Mus. Wellington, 6(18) : 285-366, 22 figs., pls. 1-9, 1971a. Classification of New Zealand Arionacea (Mollusca: Pulmonata). IV. A revision of the subfamily Otoconchinae Cockerell (Punctidae Morse). Rec. Dominion Mus. Wellington, 7(6) : 43-49, 2 figs. 1978. Classification of New Zealand Arionacea (Mollusca: Pulmonata). A review of the New Zealand charopine snails with lamellate aper- tures. Rec. National Mus. N.Z., 1(12): 177-201. 1980. Classification of New Zealand Arionacea (Mollusca : Pulmonata). VII. The genera Aeschrodomus Pilsbry and Pulchridomus n.gen. (Charopidae). Rec. National Mus. N.Z., 1(18): 293-303, figs 1-5. 1981. Classification of the New Zealand Arionacea (Mollusca : Pulmonata). VUI. Notes on some charopid species, with description of new taxa (Charopidae). Rec. National Mus. N.Z., 2(3): 9-15, figs. 1-2, 1983. Classification of the New Zealand Arionacea (Mollusca : Pulmonata). IX. The new genus Paracharopa (Charopidae). Rec. National Mus. N.Z., 2(14): 151-161, figs 1-6. 1984. Classification of the New Zealand Arionacea (Mollusca : Pulmonata). X. Montaropa macsweeneyi n.gen., n.sp. (Charopidae) and the identity of Flammutlina alpina Suter, 1904. Rec. National Mus. N.Z., 2(19): 207-210, fig. 1. COTTON, B.C. 1939. Mollusca. In: The Sir Joseph Banks Islands. Reports on the expedition of the McCoy Society for Field Investigation and Re- search. Proc. R. Soc. Vict.. 31(1): 159-176, pl. 7 COX. J.C. 1864. Catalogue of specimens of Australian land shells in the collections of J.C. Cox. J.A, Engel, Sydney, Aust. 1866a. Description d’especes nouvelles provenant d’Ausiralie et des iles Solomon et Norfolk. Journ, de Conchyl., 14: 4548. }866b. Characters of six new Australian land- shells. Proc. Zool. Soc. Lond., 1866, pp. 373— 375. 1868. A monograph of Australian land shells. 110 pp. + 22 pls. W. Maddock, Sydney, 1909. An alphabetical list of Australian land shells. Part I, 84 pp. and corrections. F.W. White, Syd- ney. COX, J.C. and Hedley, C. 1912. An index to the land shells of Victoria. Mem. Nat. Mus, Melb., 4: 5-15, pls 1-3. CUMBER, R.A. 1961. A revision of the genus Phenacohelix Suter, 1892 (Mollusca : Flam- mulinidae) with description of new species, and studies on variation, distribution and ecology. Trans. R. Soc, N.Z., 1(13): 163-196. GABRIEL, C.J. 1930, Catalogue of the land shells of Victoria. Proc. Roy. Soc. Vict., 43(1): 62-88, pls. 2-3. 1947. Additions to and alterations in the catalogue of the land shells of Victoria (including descrip- tions of new species). Mem. Nat. Mus. Vic., 15: 109-125, pls 9-10. GABRIEL, C.J. and Macpherson, J.H. 1947. Mollus- ca. In: A preliminary report on the biology and ecology of the Snowy River area in northeastern Victoria. Mem. Nat, Mus. Vic., 15: 146-171, pls 11-13. GALLOWAY, R.W. AND KEMP, E.M. 1981. Late Cainozoic environments in Australia. In: A. Keast (ed.), Ecological Biogeography of Australia, pp. 52-80. W, Junk Publ., The Hague, Netherlands. HEDLEY, C. 1889. Anatomical notes on the Helicidae, Part II. Proc. Roy. Soc. Queensland, 6: 249-251, pls 14-15. 1892. Observations on the Charopidae. Part I. Proc. Linn. Soc. NSW, 7: 157-169, pls 1-2. 1899. Descriptions of new land shells with notes on known species. Rec. Aust. Mus., 3: 151-154, pl. 28. 1912. Onsome land shells collected in Queensland by Mr Sidney W. Jackson. Proc. Linn. Soc. NSW,, 37: 253-270, pls 4-10. 1924. Some notes on Australian land shells. Aust. Zool., 3: 215-222, pls 29-32. HEDLEY, C. AND MUSSON, C.T. 1892. On a col- lection of land and freshwater shells from Queensland. Proc. Linn. Soc. N.S. W., 6(3): 551- 564. “8 IREDALE, T. 1933. Systematic notes on Australian land shells. Rec. Aust, Mus., 19: 37-59, 1937a, A basic list of the Jand Mollusca of Australia, Aust, Zool., 8(4): 287-333. 1937b. A basic list of the land Mollusca of Australia. Part IH. Aust. Zool., 9(1): 1-39, pls 1-3. 1937c, An annotated checklist of the land shells of SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 237 south and central Australia, §. Aust. Nat, 18: 6-56, pls 1-2, 1939. A review of the land Mollusca of Western Australia.J. Roy. Soe. West. Aust., 25: 1-88, pls i-5. 19414. Guide to the land shells of New South Wales. Part IL. Ause. Nar, 10(8): pp. 262-269, figs. 4-6. 194th, Guide to the land shells of New South Wales. Part TL. Aust. Nat. 11(1): pp. 1-8, figs, 7-8. |942, Guide to the land shells of New South Wales, Part IV, Awst, Nat., bl: 33-40, 1944, The land Mollusca of Lord Howe Island. Aust. Zoal., (3): 299-334, pls 17-20. KEMP, E.M, 1981]. Tertiary palaeogeogruphy and the evaluation of Australian climate. Inr A. Keast (ed.), Ecological Biogeography of Australia, pp. 3249, W. Junk Publ. The Hague, Netherlands. KERSHAW. A.P. 1980. Long term changes in north- édst Queensland rainforest. In: J. Wright, N. Mitchell, P. Watling (Eds), Reef, Rainforest, Mangroves, Man, pp. 32-38. Wildlife Preserva- lion Society of Queensland publ., Brisbane, Aust. 198], Quaternary vegetation.and environments. In: A, Keast (ed.), Ecological Biogeography of Australia, pp. 81-101. W. Junk Publ. The Hague, Netherlands. KERSHAW, R.C, 1954. Studies on Australian Charopidae (Mollusca, Pulmonata) Part 1 -The shell, ils distribution and habitation. Vie, Nat, 71; 95-96. 1955, Studies on Australian Charopidae, Part 2 - Some genera. Vie. Nat, 72: 26-30. 1956a, Studies on Australian Charopidae, Part 3 - Planate genera, Vie. Nat, 72: 137-143, 1956b. Studies on Australian Charopidae. Part 4 - Convex genera. Vic. Nat,, 73: 7-9. MARTIN, H.A, 1978. Evolution of the Australian flora and vegetation through the Tertiary: evidence from pollen. Alcheringa, 2: 181-202. MARTIN, H.A, 1981. The Tertiary flora. In: A, Keast (ed.), Ecological Biogeography of Australia, pp. 391-406. W, Junk Publ,, The Hague, Nether- lands. MONTEITH, G.B. 1980, Relationships of the genera of Chinamyersiinae. with description of a relict species from mountains of North Queensland (Hemiptera : Heteroptera : Aradidae), Pacific Insects, 21(4); 275-285. ODHNER,N.H. 1917. Mollusca. XVII. Results of Dr E, Mjobergs Swedish Scientific Expedition lo Australia 1910-1913. Kungl, Svenska, Verenskap Handl,, 52( 16), 1-115. PETTERD. W.F. and Hedley, C. 1909. A revised census ol the terrestrial Mollusca of Tasmania. Rec, Aust. Mus,, 7, 283-304, pls 82-87. PFEIFFER, L, 1850, Descriptions of twenty-four new species of Helicea from the collection of Hugh Cuming, Esq. Proc. Zool. Sec. Lond., 1849, pp. 126-131. 1851. Neve Molluskengaltungen. Zeitschr. fur Malak,, 8; 126-131. 1854, Die Helicacea. In: Martini and Chemnitz (Eds), Svstematisches Conchylien Cabinet, Bd I. Ab, 12(2): 1-562, pls 67-167. 1857. Descriptions of thirty-three new species of land shells fram the collection of H. Cuming, Esq. Prac. Zool. Soc. Lond., 1856, pp, 385-392, PILSBRY, HA. 1893, Manual of Conchalogy, 2nd series, 9; 1-366, pls 1-71, plus index 126 pp, REEVE, L, 1851-57. Cenchologia Iconica or il- lustrations of the shells of malluscous animals. Vol. 7, parts J and 2, species 1-1495, SHIRLEY, J. 1921, Land shelis of the Queensland National Park, Qld. Nat., 3(2); 32-37. SINGH, G. 1982, Environmental upheavah vegetu- tion of Australasia during the Quaternary. In: I.M.B. Smith (ed,), A History of Australasian Vegeiation, pp. +108, McGraw-Hill, Sydney, Aust, SMITH, BJ. AND KERSHAW, R.C. 1979, Field guide to the non-marine molluscs of South East- ern Australia, 285 pp. ANU Press, Canberra. 1981. Tasmanian land and freshwater molluscs. Fauna of Tasmania Handbook, 5, pp.J—I48. Hobart, Tasmania. 1985. Description of three Tasmanian charopid genera (Pulmonata : Mollusca) with notes on their type species. Ree, Queen Victorta Mus. Launceston, $8: 1-14, pls 1-3, SOLEM, A, 1970. The endodontid Jund snail genera Pilsbrycharopa and Paryphantopsis (Mollusca; Pulmonata). Veliger, 12(3); 239-264. 1972, Malacological applications of scanning eleciron micrascopy. fH. Radular structure und functioning. Veliger, 14(4): 327-336, 4 pls. 1973. Apertural barriers in Pacific Island land snails of the families Endodontidae and Charopidae. Veliger, 15(4); 300-306, 24 figs, 1976. Endodontoid Jand snails from Pacific Is- lands. Part |. Family Endodontidae. Fieldiana> Zoal, Spec. Publ., pp. 1-508, 208 figs, 1977, Shell microsculpture in Striatura. Puncrues, Radiodiscus and Planogyra (Pulmonata). The Nautilus, 91(4); 149-155, figs 1-18. 1978, Classification of the land Mollusca. Inj V, Fretter and J. Peake (Eds), Palmonates Vol. 24, 238 pp. 49-97.Academic Press, London and New York. 1983. Endodontoid land snails from Pacific Islands (Mollusca : Pulmonata : Sigmurethra). Part IT. Families Punctidae and Charopidae, Zoogeog- raphy. Field Museum of Natural History, Chicago, 336 pp. 1984, Small land snails from northern Australia, III : Species of Helicodiscidae and Charopidae. J. Malac. Soc. Aust., 6(3/4): 155-179, figs 1-45. SOLEM, A. AND CLIMO, F.M. 1985. Structure and habitat correlations of sympatric New Zealand land snail species. Malacologia, 26(1-2): 1-30, SOLEM, A., CLIMO, F.M. AND ROSCOE, D.J. 1981. Sympatric species diversity of New Zealand land snails. N.Z. Journ. Zool., 8: 453— 485. STANISIC, J. 1987. Studies on the Charopidae of tropical and subtropical Australia. I. Oreokera: a primitive genus from the high mountains of North Queensland (Mollusca : Pulmonata : Charopidae). J. Malac. Soc. Aust., 8: 1-21, figs 1-5, pls 1-2. 1988. A reevaluation of the relationships of Helix (Thalassia) gayndahensis Brazier, 1875 (Mol- lusca : Pulmonata : Helicarionidae). Mem. Qd Mus., 25(2): 475-479. TRYON, G.W. 1885. Manual of Conchology, 2nd series, 1: 1-364, pls. 1-60. 1886. Manual of Conchology, 2nd series, 2; 1-265, pls. 1-64. 1887. Manual of Conchology, 2nd series, 3: 1-313, pls. 1-63. VAN BRUGGEN, A.C. 1980. Gondwanaland connec- tions in the terrestrial molluscs of Africa and Australia. J. Malac. Soc. Aust., 4(4): 215-222, figs 1-4. VaN DYCK, S. 1982. The status and relationships of the Atherton antechinus, Antechinus godmani (Marsupialia : Dasyuridae), Aust. Mammal, 5: 195-210. WEBB, L.J. 1959. A physiognomic classification of Australian rain forests. J. Ecol., 47: 551-570. 1968, Environmental relationships of the structural types of Australian rainforest vegetation. Ecol- ogy, 49(2): 296-311. WEBB, L.J. AND TRACEY, J.G. 1981. Rainforests: patterns and change. In: A. Keast (ed.), Ecologi- cal Biogeography of Australia, pp. 606-694. W. Junk Publ., The Hague, the Netherlands. WELLMAN, P. AND MacDOUGALL, I. 1974. Cainozoic igneous activity in eastern Austraila. Tectonophysics, 23: 49-65, MEMOIRS OF THE QUEENSLAND MUSEUM APPENDIX LIST OF CHARACTERS AND CHARACTER STATES USED IN PAUP ANALYSIS 1. Tail differentiation (absent,0; present, 1) 2.Foot shape (short broad, 0; bluntly rounded,1; slender tapered,2) 3.Kidney shape (bilobed,0; moderately bilobed,1; weakly bilobed,2; almost unilobed,3) 4.Configuration of kidney apex (simple,0; reflexed,1) 5.Primary ureter configuration (simple,0; reflexed,1; coiled,2) 6.Length of secondary ureter (incomplete,0; com- plete,1) 7.Number of lobes in ovotestis (one,0; two,1; several,2) 8.Ovotestis orientation (parallel,0; right angles,1) 9.Ovotestis shape (palmately clavate,0; finger-like, 1; bilobed,2; unilobed,3) 10.Length of talon stalk (long,0; medium,1; very short,2) 11.Talon shape (circular,0; finger-like, 1) 12.Hermaphroditic duct kinking (unikinked,0; kinked,1) 13.Vas deferens shape (thin,0O; weakly expanded,1; strongly expanded,2) 14.Nature of epiphallus - vas deferens junction (simple,0; complex,1) 15.Differentiation of epiphallus (strong,0; weak,1) 16.Presence of epiphallus (present,0; absent, 1) 17.Presence of epiphallic sheath (absent,0; present, 1) 18.Epiphallus length (short,0; elongate,1) 19.Epiphallic pilaster pattern (longitudinal,0; transverse, 1) 20.Epiphallus configuration (ascending arm longer,0; ascending/descending arms equal,1) 21.Nature of epiphallus - penis junction (simple,0; complex,1) 22.Position of epiphallus - penis junction (apical,0; subapical,1) 23.Nature of epiphallic entry into penis (through pore,0; through verge, 1) 24.Position of penial retractor muscle insertion (penis,O; penis/epiphallus junction,1; epiphallus near penis,2; epiphallus far from penis,3) 25.Presence of epiphallus - penis binding (absent,0; present, 1) 26.Presence of penis sheath (absent,0; present,1) 27.Penis shape (pear shaped, 1; tubular, 1) 28.Penis expansion (slender,0; moderately ex- panded, 1; strongly expanded,2) 29.Presence of penial verge (absent,0; present, 1) 30.Penial pilaster pattern (longitudinal,0; modified, 1) SYSTEMATICS OF AUSTRALIAN CHAROPIDAE 31.Presence of internal penial collar (absent,0, present,1) 32.Shape of genitalia (expanded,0; elongate.1) 33.Length of atrium (short.0; moderately long, 1: very long.2) 34.Length of vagina (short,0; moderately long, 1; very long,2) 35.Presence of preputial tube (absent,0; present,1) 36.Presence of penial stimulatory pad (absent,0: present, 1) 37.Presence of flared extension on radular teeth (ab- sent,0; present, 1) 38.Configuration of radular row (transverse,O; v- shaped,1) 39.Nature of central tooth (strongly tricuspid,0; weak- ly tricuspid,1) 40.Nature of lateral teeth (tricuspid,0; bicuspid, 1) 41.Presence of lateromarginals (present,0; absent,1) 42.Nature of marginal teeth (multicuspid with variable-sized cusps.0; other, 1) 43.Shell cviling pattern (loose,0; normal,]; tight,2 44,Nature of sutures (impressed.0; strongly im- pressed, 1; channelled,2) 45.Presence of apertural barriers (absent,0: present, 1) 46.Presence of supraperipheral sulcus (absent,0; present,1) 47.Shape of umbilicus (closed,O: narrow-u, !: narrow- v,2; wide-v,3; cup-shaped,4; saucer shaped,5) 48,Whorl profile (keeled,O; angulate.l; flattened above and below.2; rounded,2: rounded below.4; laterally compressed,5) 49.Number of whorls (3,0-3.5,0; 3.5-4.0,1; 4.0- 4.5,2; 4.5-5.0,3; 5.0-5.5,4; 5.5-6.0,5; 6,0-6.5,6) 50.Height of shell (0-1 mm,0; 1-2 mm,1: 2~3 mm,2; 3—4 mm,3; 4-5 mm,4; 5-6 mm,5; 6-7 mm,6) 51.Shell diameter (1-2 mm,0; 2-3 mm,1; 3-4: mm,2; 4—5 mm,3; 5-6 mm,4; 6-7 mm,5; 7-8 mm,6) §2.H/D ratio (0.40-0.50,0; 0.51—0.60,1; 0.61—0.70,2; 0.71-0.80,3; 0.81-0.90,4) 53.Spire protrusion (depressed,G; 0.01-0.20 mm, 1; 0.21-0.50 mm,2; 0.50-1,00 mm,3; 1.00-2.00 mm,4) 54,Body whorl width (0.01-0.50 mm,0; 0.51—1.00 mm,.1; 1.01-1.50 mm,2; 1.51-2.00 mm,3; 2.01— 2.50 mm,4; 2.51-3,00 mm,5; 3.01-3.50,6) 55.SP/BWW ratio (0.00-0, 10,6; 0,1 1-0.20,1; 0.21- 0.30,2; 0.31-0.40,3: 0.41-0.50,4, 0.51-0.60.5; 0.61-0.70,6) 56.Number of nbs on body whorl (25-50,0; 51-75,1: 76-100.2; 101-125,3; 126-1504; 151-175.5; 175-,4) 57.Ribs/mm (0.01-5.00,0; 5.01-10.00,1; 10.01— 15.00,2; 15.01-20.00,3; 20,01-25.00,4; 25.01- 30.00,5; 30.01-35.00,6) 58.Umbilical width (closed,0; 0.01-0.50,1: 0.51- 1.00.2; 1.01-1.50,3; 1.51-2.00,4) 59.D/U ratio (0.01—2.50,0; 2.51-5,00,1; 5.51—7.50,2; 7,51-10,00,3; 10.00-closed,4) 60.Nature of apical sculpture (spiral,(; predominantly radial, 1; other,2) 61.Presence of radials on terminal part of protoconch (absent: present, 1) 62.Presence of regular pits on protoconch (present,0; absent.1) 63.Presence of irregular pits on protoconch (present,O; absent,1) 64.Presence of apical spiral grooves (present,0; ab- sent,|) 65.Presence of apical spiral segments (present,0; ab- sent, 1) 66.Presence of post nuclear spiral grooves (present,0; absent, 1) 67.Presence of post nuclear spiral furrows (present,0; absent, 1) 68.Presence of two-part sculpture on adult whorls (present.0; absent,1) 69,Presence of post nuclear microradials (present,0); absent, 1) 70,Presence of periostracal blades on major ribs (present,0; absent,1) 71.Presence of post nuclear periostracal selae (present,0; absent,1) 72.Shell colour (monochrome,0; flammulated,1) 73.Presence of large major radial ribs (present.0; ab- sent,1) 74.Presence of ‘elongate—bead’ microsculpture (present,0; absent, 1) 75.Nature of beaded microsculpture (elongale,0: cir- cular.1) DATA FOR PAUP ANALYSIS | ,OPTION SETTINGS INGOTS « ppt cee oe a ee 50 NCHAR ...-.. 1% User-tress) . iuek) sa opal we NO HYPANG tt, . ct ves eaeoh atta ta » SH ADDSEQ. 2). ee pe oe CLOSEST | (07 ic aaa a aie Cs ne a 1 SWAP ere tbe tet ees en FE GLOBAL MULPARS: © py ectfhiee) item elt adds s a YES OF e ce fe cae vet tees 2 Ss FARRIS ROOF 26 ef oe ANCESTOR Weightsapplied ....,,.,4-, net NO OUTWIDTH ~o:. emp pre (bpp cea 80 Missing datacode - 2.2.2.2... 204. 9 MAXTREE 100 2. STATISTICS DERIVED FROM CONSENSUS TREE Consensus fork index (component count) = 42 CF (normalised) = 0.875 Term information = 400 Mickevich’s consensus information (Cl) - 0.290 Levels sum - 6473 Rohlf's C11) = 0.813 Rohlf’s - log Cl(2) = 0.16566E+ 05 Weighted consensus fork = 0.361 240 MEMOIRS OF THE QUEENSLAND MUSEUM PAUP DATA MATRIX Character states for species in order of listing above. dorrigo; 0020001107710000000000020001001000001011001000213351343??7141011111111011117 Llevicos: 75030002101 10000001000100000100110200100100200020434133422311011311111010112 ~°502000110101000000000002000100100000101100100021335134377232011010111011117 crepap pitoz obopepesoogel LeDsrotaooododaverso02Bss1Sa2>2za0KnELOMAIONIET? #7 50201022010000000000000100010000000000001011002422322417722201011010021011? *{92000220111000000000000000100112200100010100020666345622232010111101110112 **102000220111000000000001000100110200100010100020445344622232010111101110112 * 7021 20200200000000001102000100000101000000110034322122111310011111010101012 50212020020000000000000300011000010000000021002331112122121001112111012101? ¥#90212020020000000000100200010000010000000011003431211211131003111201010101? 73021 20200200000000000100000101000100000000110034423223210320011111010101012 3223222 2272777227272722227227727272277227721000211444335311421111101111 10117 *7920001000200000000000020000101700107000000110033232011122210011111110110000 7%910001100000000000000010010100001100000000110044211111200200011111110011001 ¥922001110101100001000110111000012200000000110043246106030411011111110110017 *°9220011007701000010001001110000122000000001300453341 0501041101111 rLOLLOO1? P332200110000010000100010011100001220000000011004523420400041101 1111110110012 oN22001100000100001000100111000012200000000110043224114031412001111110111017 7)22001100100000000000100001010012200000000210045423103021402001111110111017 °922001100100000000000100001000012200000000210045312102021312001111110110012 722001 100000000000000000000100001101011011110033224104031312001111110011017 *°910001001200000000000001001000011201001001211152101001027212010111110110112 “ort an Ml GdabobovoonndndoshantooddosdLomdnobrsadsasiadMitorstieLilidiloLiond: at510091 1004 G009909099N00S0001 00012 20L0NDANDAIDOD3a113z7IELIE4LONETTEIZORIHOOL a FTL O1LOOLOZARADADAOMOGHOLIGUNON10000NG00T200232411411 29x310111 1 110110072 err Lao povastavuongsoeZUDIeodDouLGONGOtREredonsaveLEarasos cect it moneDcle *#311101100110261000000000001200002100000000120023212121311221021111110110017 pe312101100100000000000001010100012201000000211054300001053201011111110111000 rr TaTiNOLOANSDEPOOODAnADAOAMAOoOdKODONdheTapAdsoiorrodaessorad saasd1140be SYSTEMATICS OF AUSTRALIAN CHAROPIDAE kempsey: 012101100100000001000001010100012200000000211054411001066301011111110111000 minutis: 012101100100000000000001000100010200000000211054300011023201011111110110000 colmani: elizabe: 012101100100000000000001000100012200000000211054311011111301011111110111000 bairnsd: 012101101000000000000001000100011200000000211054301011001201011111110110000 nautilo: *522101100100010000000001000100100200000000110003223123152041011111110110012 #°522101100110010011000003000100100200000000020103224103041041011111110110012 nro Prods muneteaygountousueexeunewoaca mals vunadegse ra re ONL UNE ptaGane 912001 10020000000000000111010000110000000011003221112120021101111111001011? p91?22?100?0000000000000200010000010000000011003221111127721101111111001011? a Tad naa dovwnesnodabiousewepioianesennnorontdadssiaratsesa 1191117149108? multira: Mr ReLaSh ovo vaiebeaNnomDtonmuns edgendaTnuaer bios seraenveint epilds needa LLoeevocinaagianeadguaToritus uompeioet on iaenl errinsieaer rie) ioone sen *922001102100000000110003000110010210000000210013411211122132011101110010000 v2 700 LO BUOOSgMbEd1ANHOSDULNhG102 boDadbHstOD IsEOS si edTi FoMAOTdAHtODOD aperta: 241 CONTENTS STANISIC, J. Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pulmonata) from the subtropicalinainforests ae... ctyspeRoseetes ic, sect ic evlcistet ose eeranape oie mve # cleo lo alloc at tae 1