Memoirs OF THE Queensland Museum VOLUME 34 1994 PUBLISHED BY ORDER OF THE BOARD VOLUME 34 IS COMPLETE IN THREE PARTS © Queensland Museum PO Box 3300, South Brisbane, Q. 4101, Australia. Phone (07) 840 7555. Fax (07) 846 1918 National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Memoirs of the Queensland Museum may be reproduced for scientific research, individual study or other educational puiposes. Properly acknowledged quotations may be made but queries regarding the republicalion of any papers should be addressed to the Director. Copies of the journal can be purchased from the Queensland Museum Bookshop. A Queensland Government Project Typeset at the Queensland Museum. Printed by Merino Lithographies, 18 Baldock Street, Moorooka, Queensland 4105. CONTENTS ^ART 1 (Issued 24 December, 1993) iTANISIC,J. The identity of Helicarion semoni Martens, 1894: a large semi-slug from the Wet Tropics, northeastern Queensland (Pulmonata: Helicarionidae) 1 5TANISIC, J. Danielleilona gen. nov., from the Wet Tropics, northeastern Queensland (Pulmonata: Charopidae) 11 5TANISIC, J. Lenwebbia paluma sp. nov., from the Wet Tropics, northeastern Queensland (Pulmonata: Charopidae) . . 21 5TANISIC, J. Eungarion mcdonaldi gen. et sp.nov., a montane semi-slug from mideastem Queensland rainforests (Pulmonata: Helicarionidae) 27 3RAILOVSKY, H. A revision of the Tribe Colpurini from Australia (Hemiptera-Heteroptera-Coreidae) 35 5HORT, J.W. Caridina zebra^ a new species of freshwater atyid shrimp (Crustacea; Decapoda) from northeastern Queens- land rainforest 61 SHORT, J.W. & DAVIE, P.J.F. Two new species of freshwater crayfish (Crustacea: Decapoda: Parastacidae) from northeastern Queensland rainforest 69 RAVEN, R.J. The biodiversity of Australian mygalomorph spiders. 1. Two new species of Namirea (Araneae:Dipluridae) 81 RICHARDS, S.J. Functional significance of nest construction by an Australian rainforest frog: a preliminary analysis .... 89 COUPER, P.J., COVACEVICH, J.A. & MORITZ, C. A review of the leaf-tailed geckos endemic to eastern Australia: a new genus, four new species, and other new data 95 WHITTIER, J.M. Ecological notes on Carlia rostralis in rainforest and associated habitat in the southern Wet Tropics ... 125 CUNNINGHAM, M. Reproductive biology of the Prickly Forest Skink, Gnypetoscincus queenslandiae, an endemic species from northern Queensland 131 SADLIER, R.A., COLGAN, D.J. & SHEA, G.M. Taxonomy and distribution of the scincid lizard Saproscincus challengeri and related species in southeastern Australia 139 COVACEVICH, J.A., COUPER, P.J. & JAMES, C. A new skink, Nangura spinosa gen. et sp.nov., from a dry rainforest of southern Queensland 159 JAMIESON, B.G.M, & SCHELTINGA, D.M. The ultrastructure of spermatozoa of Nangura spinosa (Scincidae, Reptilia) 169 INGRAM, G.J. & COVACEVICH, J.A. Two new species of striped blindsnakes 181 COVACEVICH, J.A., COUPER, P.J. & INGRAM, G.J. New reptile records from rainforests of south and mideastem Queensland 185 COVACEVICH, J.A. & MCDONALD, K.R. Distribution and conservation of frogs and reptiles of Queensland rainforests 189 JOSEPH, L., MORITZ, C. & HUGALL, A. A mitochondrial DNA perspective on the historical biogeography of mideastem Queensland rainforest birds 201 HORSUP, A., JAMES, C. & PORTER, G. Vertebrates of dry rainforest of south and mideastem Queensland 215 WERREN, G.L. Conservation strategies for rare and threatened vertebrates of Australia’s Wet Tropics Region 229 NOTES RICHARDS, S.J., MCDONALD, K.R. & INGRAM, G.J. Recognition of Litoria eucnemis (Lonnberg) in Australia 94 MCDONALD, K.R. & STORCH, D.R. A new reproductive mode for an Australian hylid frog 200 WERREN, G.L. & TRENERRY, M.P. Size and diet of Bufo marinus in rainforest of northeastern Queensland 240 WHITTIER, J.M. & MOELLER, D.R. Varanus prasinus (the Emerald Goanna) on Moa Island, Torres Strait, Australia 130 TRENERRY, M.P. & WERREN, G.L. Possum assemblages in rainforest of the Carbine Uplands, NEQ, with special reference to Hemibelideus lemuroides 188 PART 2 (Issued 1 March, 1994) LAWRENCE, D. Customary exchange across Torres Strait 241 PART 3 (Issued 1 August, 1994) BABINCHAK, J.A., MOELLER, P.D.R., VAN DOLAH, F.M., EYO, P.B. & RAMSDELL, J.S. Production of ciguatoxins in cultured Gambierdiscus toxicus 447 BAGNIS,R. Natural versus anthropogenic disturbances to coral reefs: comparison in epidemiological patterns of ciguatera 455 BENOIT, E. & LEGRAND, A.M. Gambiertoxin-induced modifications of the membrane potential of myelinated nerve fibres 461 BLYTHE, D.G., FLEMING, L.E., AYYAR, D.R., DE SYLVA, D., BADEN, D. & SCHRANK, K. Mannitol therapy for acute and chronic ciguatera fish poisoning 465 DALZELL, P. Management of ciguatera fish poisoning in the south Pacific 471 DICKEY, R.W., GRANADE, H.R. & McCLURE, F.D. Evaluation of a solid-phase immunobead assay for detection of ciguatera-related biotoxins in Caribbean finfish 481 HOKAMA, Y., ASAHINA, A.Y., TITUS, E., ICHINOTSUBO, D., CHUN, S., HONG, T.L.W.P., SHIRAI, J.L., ASUNCION, D.A. & MIYAHARA, J.T. Assessment of ciguateric fish in Hawaii by immunological, mouse toxicity and guinea pig atrialassays 489 HOLMES, M.J. & LEWIS, R.J. The origin of ciguatera 497 HOLMES, M.J., LEWIS, R.J., SELLIN, M. & STREET, R. The origin of ciguatera in Platypus Bay, Australia 505 ICHINOTSUBO, D., ASAHINA, A.Y., TITUS, E., CHUN, S., HONG, T.L.W.P., SHIRAI, J.L. & HOKAMA, Y. Survey for ciguatera fish poisoning in west Hawaii 513 KALY, U.L. & JONES, G.P. Test of the effect of disturbance on ciguatera in Tuvalu 523 LANG, R.J., VOGALIS, F., HOLMES, M.J. & LEWIS, R.J. Maitotoxin induces muscle contraction and a non-selective cationic current in single smooth muscle cells of the guinea-pig proximal colon 533 LEWIS, R.J. Immunological, biochemical and chemical features of ciguatoxins: implications for the detection of ciguateric fish 541 LEWIS, RJ- Impact of a validated, cost effective screen for ciguateric fish 549 LEWIS, R J. & BRERETON, I.M. Inverse-detected NMR of ciguatoxin: quaternary carbon locations confirmed in CTX-1 555 LEWIS, R.J., HOLMES, M.J. & SELLIN, M. Invertebrates implicated in the transfer of gambiertoxins to the benthic carnivore Pomadasys macidatus 561 LEWIS, R.J., SELLIN. M., tGILLESPIE, N.C., HOLMES, M.J., KEYS, A., STREET, R., SMYTHE, H., THAGGARD, H. & BRYCE. S. Ciguatera and herbivores: uptake and accumulation of ciguatoxins in Ctenochaetus striatus on the Great Barrier Reef 565 MANGER, R.L., LEJA, L.S., LEE, S.Y., HUNGERFORD, J.N. & WEKELL, M.M. Cell bioassay for the detection of ciguatoxins, brevetoxins, and saxiloxins 571 MOLGO, J., JUZANS, P. & LEGRAND, A.M. Confocal laser scanning microscopy: a new tool for studying the effects of ciguatoxin (CTX-IB) and D-mannitol at motor nerve terminals of the neuromuscular junction in situ 577 PARK, D.L. Reef management and seafood monitoring programs for ciguatera 587 PAYNE, J. Ciguatera poisoning: current issues in law 595 PEARN, J. Ciguatera: dilemmas in clinical recognition, presentation and management 601 PEARN, J. & LEWIS, R. Ciguatera: risk perception and fish ingestion 605 RUFF, T.A. & LEWIS, R.J. Clinical aspects of ciguatera: an overview 609 TERAO, K., ITO, E., OHKUSU, M. & YASUMOTO, A. Pathological changes in murine hearts induced by intermittent administration of ciguatoxin 621 VERNOUX, J.P. The mouse ciguatoxin bioassay: directions for use to control fish for consumption 625 VERNOUX, J.P. & LEJEUNE, J. Ciguatera in the french West Indies 631 ABSTRACTS BROCK, J.A., JOBLING, P., McLACHLAN, E.M. & LEWIS, R.J. Effects of Ciguatoxin-1 on electrical activity recorded intracellularly from rat tail artery in vitro 454 CAPRA. M.F., CAMERON, J., FLOWERS, A.E. & PURCELL, C.E. Responses of vertebrate nerves to Ciguatoxin 454 CHALOUPKA, M.Y., LEWIS, R.J. & SELLIN, M. The changing face of ciguatera prevalence 554 HAHN, S.T., CAPRA, M.F. & MILLER, D.M. Oral and intraperitoneal administration studies of toxins derived from fish tissues and extracts of cultured G. toxicus in the humbug {D. aruanus). Damsel-fish (P. wardi) and the Stripey (L. carponotatus) 554 HALLEGRAEFF, G.M. On the global increase of harmful algal blooms 560 HAMBLIN, P., McLACHLAN, E.M. & LEWIS, R.J. Ciguatoxin-1 induces spontaneous synaptic activity in isolated sympathetic ganglia of guinea pigs .... 560 LEGRAND, A.F. & LOTTE, C.J. Detection of ciguatoxic fish by using the binding property of ciguatoxins to voltage-dependant sodium channels 576 PALAFOX, N. Evaluation of intravenous mannitol for treatment of actute ciguatera fish poisoning 576 PURCELL, C.E., CAMERON, J. & CAPRA, M.F. Modification of nerve conduction in the rat by brevetoxin (PBTX~3) 586 SCHEUER, P.J. Ciguatera research - an historical perspective 586 YASUMOTO, T., SATAKE, M., MURATA, M. & NAOKI, H. Structures of maitoloxin and ciguatoxin congeners isolated from cultured Gambierdiscus toxicus 600 2 4 ) THE IDENTITY OF HEL1CARION SEMONI MARTENS, 1894: A LARGE SEMI-SLUG FROM THE WET TROPICS, NORTHEASTERN QUEENSLAND (PULMONATA: HELICARIONIDAE) JOHN STANISIC Stanisic, J. 1993 12 24: The identity of Helicarion semoni Martens, 1894: a large semi-slug from the Wet Tropics, northeastern Queensland (Pulmonata: Helicarionidae). Memoirs of the Queensland Museum 34(1): 1-9. Brisbane. ISSN 0079-8835. The identity of the semi-slug Helicarion semoni Martens, 1894, is established. The name is applied to a large species from the northern region of the Wet Tropics, NEQ, and the type locality is redesignated as the Big Tableland, south of Cooklown, NEQ. On the basis of conchological and anatomical features the species is referred to Thularion gen.nov. and redescribed. Some aspects of its biogeography and relationships are discussed. □ Pul- monata, Helicarionidae, Thularion semoni {Martens, 1894), new genus, semi-slug, sys- tematics, biogeography, Wet Tropics. John Stanisic, Queensland Museum, PO Box 3300, South Brisbane, Queensland, 4101, Australia. Semi-slugs belonging to the family Helicarionidae are particularly diverse in eastern Queensland. Iredale (1937) listed 8 species while Smith (1992) listed 7 with a further 2 as incertae sedis. Based on material acquired by the Queensland Museum over the past 1 3 years these are underestimations. Among the described forms several, including Helicarion semoni Mar- tens, 1894, are taxonomic enigmas. Although listed under Parmacochlea Smith, 1884, by Iredale (1937) the species was considered incer- tae sedis by Smith (1992). Long efforts by the author to locate the type were fruitless until a chance discovery of two syntypes by Dr Rudolf Kilias in the collections of the Zoological Museum, Humboldt University, Berlin. Ex- amination of the types revealed that they were conspecific with a large helicarionid semi-slug from the northern region of the Wet Tropics, NEQ. The type locality given by Martens (1894) was Burnett R., SEQ (Fig. 1), which may explain the confusion surrounding this species even though the original descriptions of both shell and animal were excellent. Marten’s introduction of H. semoni was included in a listing of molluscan species (land and freshwater) collected from east- ern Queensland in 1891-1893. Although the Bur- nett R. featured in several species descriptions other Queensland localities were visited by the expedition and these included Hammond Is., Tor- res Strait, and Cooktown, NEQ. Since both the original description and the types agree with the NEQ animal, I consider that there was a labelling error during either collection or study of the specimens. Cooktown is situated in the far north of the Wet Tropics and would have provided ready road access to the area where H. semoni occurs. The black form of the animal (Fig. 3a), which was referred to in the original description, comes from the Big Tableland area, near Helenvale, NEQ. The species is placed in Thularion gen.nov. Studies of Australian helicarionid semi-slugs are sparse. Baker (1941) figured the anatomy of a specimen of 'Helicarion* leucospira (Pfeiffer, 1857) and R. Kershaw (1979, 1981, 1983) and Dartnall and R. Kershaw (1978) completed studies on some southeastern Australian species. Most references consist of brief single species descriptions and mentions in faunal checklists. Hence it is not surprising that generic relation- ships and even species identities are poorly un- derstood. Although a comprehensive revision of the group is clearly required the author has dis- sected a number of other eastern semi-slug taxa in order to place the following observations in a wider context. I am grateful to Ron C. Kershaw for supplying many personal notes and drawings on eastern semi-slugs. They have facilitated this project. The following abbreviations are used:- Q-Queensland, NSW-New South Wales, ZMB-Museum fiir Naturkunde der Humboldt Universitiit, Berlin, Germany, QM-Queensland Museum, CMVF-complex mesophyll vine forest. DG-prostate, E-epiphallus, EC-epiphallic caecum, EF-epiphallic flagellum, EP-epipallic pore, G-ovotestis, GD-hermaphroditic duct, GG- 2 MEMOIRS OF THE QUEENSLAND MUSEUM albumen gland, LL-left shell lappet, LML-lefl mantle lobe, P-penis, PPM-main penital pilaster, PRM-penial retractor muscle, PS-penial sheath, RL-right shell lappet, RML-right mantle lobe, S-spermatheca, SH-shell, SP-spermatophore, SS- spermathecal stalk, UT- uterus, UV- free oviduct, V-vagina, VD-vas deferens, Y-atrium, Z-diges- tive gland. SYSTEMATICS Family Helicarionidae Thularion gen.nov. Diagnosis Shell degenerate with incomplete early whorls and membranous base, depressed, and with reduced whorl numbers. Shell sculpture of fine crowded spirals on the protoconch becoming ob- solete on the adult whorls; body whorl with wide- ly spaced, spiral furrows. Animal large with shell lappets prominently united posteriorly; left mantle lobe forming a prominent cephalic shield. Genitalia with epiphallic caecum and flagellum. Penis with sheath, internally with a large lamel- late pilaster and pustular wall ornamentation; no verge or papilla present. Etymology Latin thule north; referring to the distribution of the genus in northern Australia. Type Species Helicarion semoni Martens, 1894; herein designated. Comparisons Thularion differs most markedly from Fas- tosarion (-^Vercularion) (type species: Vitrina superba (Cox, 1871) by its degenerate, depressed shell which has lower whorl numbers. The exter- nal animal characters of Thularion, particularly the enlarged shell lappets and mantle lobes, are additional discriminating features. Par- macochlea (type species: Parmacochlea fischeri Smith, 1884) has an even more reduced shell and greatly enlarged shell lappets and mantle lobes. The penis oi Thularion is pustulose with a lamel- late pilaster apically and no vergic structures. In contrast the penial wall sculpture of Fastosarion consists of obliquely arranged, longitudinal pilasters complemented by a large central pilaster and either an apical verge or papilla (R. Kershaw, pers. comm.). Parmacochlea has a strongly papil- lose penis interior with a large central papillose pilaster and a verge. Thularion semoni (Martens, 1894) comb. nov. (Figs. 1-7) Helicarion semoni Martens, 1894, p.87, pl.4, fig.8a-e; Smith 1992,p.244. Parmacochlea semoni (Martens); Iredale, 1937, p.lO. Comparative Remarks Thularion semoni can be distinguished from Fastosarion brazieri (Cox, 1873) (type locality: Mt Bellenden Ker, NEQ) by its depressed, degenerate shell (Fig. 2a-b), lower whorl num- bers and enlarged shell lappets which are strongly fused posteriorly (Fig. 5a). Parmacochlea spp. are smaller and have a simple, plate-like shell. Anatomically the very long penis with apical pilaster (Fig. 5b, c) immediately differentiates T. semoni from other sympatric semi-slugs (Stanisic, unpubl.). Previous Studies Martens (1894) compared T. semoni with Peloparion helenae (Godwin-Austen, 1883) from the Sydney region, NSW, on the basis of overall similarity of T. semoni to Godwin- Austen’s drawings. However, the Sydney semi- slug is much smaller and is only grossly like the northern species (R, Kershaw, pers. comm.). Ire- dale (1937) referred T. semoni to Parmacochlea Smith, 1 884, without giving reasons. Presumably the allocation was based on the degenerate nature of the shell. However, although the shell of T semoni is reduced it is not plate-like as in Par- macochlea spp. The superficial nature of this conchological similarity is highlighted by major differences in anatomy (external and internal) between Parmacochlea and T semoni. Smith (1992) listed the species as incertae sedis and indicated that the type was probably lost. Type Locality Big Tableland, c. 15k S of Helenvale, NEQ - herein designated. Material examined Lectotype: ZMB46231, Burnett River (error), Queensland. Collected by R.W. Semon. Height of shell 7.(X)mm, max. diameter 24.92mm, min. diameter 16.58mm, H/D ratio 0.28, whorls 2V8 - herein desig- nated. Paralectotype: ZMB46231, I specimen, same col- lection data as lectotype (damaged). Other MaterialiNEQ; Lorna Doone, c. 15k SE HELICARION SEMONI 3 4 em^ant c/lM/r, Zoolog. Museum Berlin. ^6 i 3 y ■X’yJc c- ocycx^o-T^ C7<2^2x^> CJ^ ■<^ 2^^'7no €-(X^lc/^ 5^ ^a:^i ^ 52'r>^ eJLy FIG. 1 .Provenance of type material Helenvale, (15°47’S, 145“I7’EX CMVF/Palms, under Logs and discarded palm fronds (9, QMM023769, 4 Oct 1988» J. Stanisic, D. Potter); Mt Finnigan, summit, 1050m (15°48’S, 145°17’E) (4, QMM044296, 3-5 Dec 1990; 5, QMM044295 16-17 Jan 1991, G.B. Monteith); Big Tableland, via Helenvale, rainforest, under logs (1. QMM014166, 17 Dec 1983, J. Covacevich); Mt Misery, via Shiptons Flat (15°53’S, 145^4* E), summit, 850m (I, QMM044297, 6 Dec 1990, G.B. Monteith); Big Tableland (15°43’S, 145°17’E), 700m (3, QMM044299, 19-20 Dec 1990, G.B. Monteith); Mt Sorrow, via Cape Tribulation, 300-800m, rainforest (1, QMMO43308, 15 Oct 1980, G.B. Monteith); Table Top, Mt Finnigan, Shipton’s Flat, rainforest, in litter (1, QMM04697, 1 1 Jun 1971, D. Douglas). Diagnosis Shell (Fig. 2a, b) large, maximum diameter 22. 00-25. 30mm (mean 24.14mm), minimum diameter 16.58-^8. 82mm (mean 17.49mm), very thin, lenticular, homy, poorly calcified. Whorls 2 V 2 + - 2^8+ (mean 2%), rapidly expanding. Body whorl greatly inflated and descending slightly in front, with lower margin membranous. Spire and apex flattened. Height of shell 6.49-1 1.69mm (mean 9.51mm), H/D ratio 0.27-0.53 (mean 0.40). Protoconch (Fig. 4a) of 1 V 4 whorls, sculp- tured with fine, crowded, incised spiral lines. Post-nuclear sculpture (Fig. 4c-e) of fine radial growth lines and sparse spiral furrows on the body whorl. Microsculpture (Fig. 4e-f) of very fine, almost obsolete, periostracal, spiral wrinkles. Sutures (Fig. 4b) shallowly impressed. Whorls flattened above and rounded below the periphery. Internal walls of early whorls (Fig. 2b) membranous and incomplete. Lip thin, membranous below. Colour yellow with whitish apex; interior chalky white. Animal (Figs. 3a,b; 5a) large, body length in preservative 45.9-59.4mm (mean 53.0mm, n=7). Foot moderately broad, tripartite, rounded anteriorly, tapered posteriorly. Tail relatively long, high, sharply keeled mid-dorsally. Caudal horn vestigial; caudal foss a long vertical slit in the tail. Pedal grooves typically aulacopod. Colour (in life) black and reddish-brown with FIG. 2. Lectolypeof777w/anc»n.y^mo/7/ (Martens 1894), ZMB46231. Scale in mm. 4 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3.Colour moqjhs of Thularion semoni. a, black, Big Tableland, NEQ, QMM014166; b, brown, Mt Finnegan, NEQ* QMM04697. brown markings to black (Fig. 3a,b), always with varying degrees of paler creamy-yellow markings on the body ornamentation; pustules on mantle lobes and shell lappets also with paler markings. Foot black to reddish-brown with paler edges and paler central region. Right mantle lobe small, with short, lobate anterior extension. Left mantle lobe large, with a triangular, tapered lobe posteriorly, expanded into a cephalic shield anteriorly, and fused with left shell lap anteriorly. Right shell lap fused with right mantle lobe at lower margin and with left shell lap posteriorly. In life, right shell lap covering apex and about one-third of shell; left shell lap covering remainder of shell. Genitalia (Fig. 5b) with mullilobate ovoteslis buried in the digestive gland. Talon a blind spherical pouch on a short peduncle embedded in the surface of the albumen gland. Hermaphroditic duct long and convoluted; carrefour embedded in the albumen gland. Prostate a band of creamy alveoli appressed to the upper three-quarters of uterus. Uterus with a short, narrow basal section, giving rise to a larger, highly sacculated apical portion. Vas deferens arising from the prostate- uterus as a highly convoluted, thin tube, descend- ing to the penioviducal angle then ascending the penis as a straight narrow tube to enter a wider, muscular epiphallus. Epiphallus shorter than penis, apically with a long thin flagellum which is weakly connected to the side of the penis, internally with longitudinal pilasters basally, be- coming transverse apically. Penial retractor muscle inserting sub-apically onto epiphallus at a point where a short, club-shaped caecum arises. Vagina short with irregularly arranged, thin, lon- gitudinal thickenings. Free oviduct expanded, in- ternally with longitudinal thickenings that become transverse apically, entering vagina sub- apically. Spermatheca with a long, thin stalk and an ovately elongate head, insert^ apically on vagina. Penis (Figs. 5b, c) long, reflexed centrally; apical region with simple entrance of epiphallus and short, crowded, transversely elongate pus- tules that become less crowded in some areas and a long, slender, lamellate pilaster; basal region with a sheath, internally with short, narrow, crowded, transverse pustules, tending to be more crowded centrally. Atrium short, without unusual features. Spermathophore (Fig. 5d) elongately ovoid with a long, thin chitinous tail. Based on three (two dissected) specimens (QMM023769, QMMOI4166). Radula (Fig. 6a-0 having a tricuspid central tooth with lanceolate mesocone and reduced lateral cusps; laterals strongly bicuspid with en- docone reduced to a tiny point on the edge of the mesocone, ectocone short and pointed, anterior edge irregularly swollen and grooved; marginals bicuspid with a long curved shaft, long and curved mesocone, short ectocone, no endocone, rounded cusps. Basal plates of central and laterals short, squarish, asymmetrical in laterals with an elevated lateral ridge. Interrow support provided by interlock of anterior edge with basal plate of tooth in front. Radular formula 83.23.1.23.83. Distribution and Habitat Tropical rainforests of the coastal region be- tween Cape Tribulation and Cooktown, NEQ (Fig.7); mainly in the uplands and highlands; probably an opportunistic riparian species in the foothills where circumstances allow. Its apparent absence from the Thornton Peak area, south of Cape Tribulation needs confirmation. HELICARION SEMONI 5 FIG. 4.Shell details of Thularion semoni. a-c, Mt Finnegan, NEQ, QMM044296; d-f, Loma D^ne, NEQ, QMM023769. a, spiral furrows on body whorl; b, protoconch sculpture; c, penostracal scratches on body whorl, d, radial growth ridges on body whorl; e, microsculpture on body whorl; f, radial periostracal wrinkles due to drying. Scale lines as marked. Remarks Thularion semoni differs from species of Fas- tosarion and Parmacochlea in external anatomy, shell features and genitalia and appears to repre- sent a level of semi-slug development inter- mediate between these two genera. For these reasons it has been given separate generic status. 6 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5.Anatomy of Thularion semoni. a, Big Tableland, NEQ, QMM014166; b-d, Loma Doone, NEQ, QMM023769. a, whole animal showing shell lappets and mantle lobes (body ornamentation not shown); b, reproductive system; c, penis interior; d, spermatophore. Scales lines = 5mm. DISCUSSION Establishing the identity of T. semoni provided an opportunity to study some features of the east coast semi-slug fauna. Although most species have a superficial similarity this gross resemblance overlays a complex evolutionary history. Concentration of helicarionid semi-slugs in the coastal and subcoastal mountain ranges of eastern Australia is not accidental. These mountains are mostly volcanic and their eastern slopes lie in the HELICARION SEMONI 7 HG. 6.Radular details of Thulanon semoni, Loma Doone, NEQ, QMM023769. a, cental and lateral teeth showing irregular anterior edge and elevated ridges on basal plates; b, laterals; c, laterals showing interrow support mechanism; d, laterals and marginals; e-f, marginals. Scale lines as marked. path of prevailing winds which provide plentiful rain. The derived soils are acidic and there is relatively little calcium available for shell manufacture. Under these conditions the need for a shell to resist desiccation is reduced and the energy budget required to produce a shell is in- creased providing an ideal environment for slug evolution. The Helicarionidae is considered a recent addi- tion to the Australian land snail fauna having migrated from the Indo-Malayan area following collision of the Australian plate with the Sunda 8 MEMOIRS OF THE QUEENSLAND MUSEUM Island arc in the mid-Miocene (Bishop, 1981). The post-Miocene drying of the continent, which led to the retreat of mesic habitats elsewhere, would have been ameliorated along the east coast by the presence of elements of the Great Dividing Range. They maintained moist forests along their length and, most likely, provided a corridor for movement and colonisation of semi-slugs in east- ern Australia. Drying events of the late Quater- nary occurred in more rapid succession than those of the Miocene and Pliocene (Galloway and Kemp, 1981) and would have had severe impacts on moisture-sensitive animals such as semi-slugs. Many groups would have survived only in upland refugia and the present-day distribution patterns of Mysticarion (highlands of the Great Dividing Range from SEQ to central NSW) and 'Helicarion' australis in the uplands of the Bor- der Ranges, SEQ (alt>800m), are probable ex- amples of the results of this climatic attrition. The move to slugdom (limacization) has oc- curred more than once and probably in more than one way, and in the Wet Tropics region three distinct grades of semi-slug development are evi- dent. TT»ese correlate with degrees of shell reduc- tion - least in Fastosarion (relatively high whorl count, fully formed whorls, spire elevation) and greatest in Pannacochlea (very low whorl count, shell flat and plate-like). Thularion with a depressed shell that has a relatively low whorl count, incomplete internal whorls and a membranous base is intermediate between these extremes. Elsewhere along the east coast many semi-slugs e.g. Helicarion s.s. show the Fas- tosarion shell form but in the wetter parts of the eastern ranges (Border Ranges, SEQ; New England - Barrington Tops area, NSW; Clarke Range, MEQ) more extreme examples of shell reduction are evident. The distribution and habitat of Thularion semoni indicate that the high mountains of the northern Wet Tropics were probably crucial in its survival, especially during more recent drying events (Kershaw, 1981). Compared with Fas- tosarion brazieri, which has been recorded from many upland (Atherton Tableland) and lowland localities in the Innisfail-Caims area, T. semoni is known almost exclusively from upland and high- land localities. The greater shell reduction in T. semoni and concommilantly greater area of ex- posed animal tissue (including expanded shell lappets and mantle lobes) makes this species potentially more sensitive to climatic fluctuation than F. brazieri. This provides an ecological basis for explaining not only its current distribution pattern but also past restrictions of the species. The role of montane refugia in Wet Tropics land snail biogeography has been previously high- lighted for the Gondwanan family Charopidae (Stanisic, 1987). Their importance to ‘modem’ families highlights the significant effects of Late Tertiary climatic fluctuations on all east coast terrestrial molluscs. The relationships of Thularion are problematic, due mainly to the lack of adequate revisionary studies on related taxa. The conchological and anatomical features of Thularion can be derived from the more generalised Fastosarion and it is probable that it is a localised derivative of a Fastosarion - like ancestor. Interestingly no Fas- HELICARION SEMONI 9 tosarion grade semi-slug is found north of the Daintree R., indicating that Thularion may have colonised this region in the absence of Fas- tosarion. The restriction of T. semoni to the north- ern part of the Wet Tropics again exposes the role of the Daintree R. as a possible barrier to land snail dispersal in the region (Stanisic, 1987). Whether or not Parmacochlea is a direct descen- dant of a Thularion - like ancestor or whether it is derived independently from a more generalised ancestor has yet to be determined. ACKNOWLEDGEMENTS I wish to thank Ron Kershaw (Queen Victoria Museum and Art Gallery, Launceston) for sup- plying essential notes on the Australian Helicarionidae which contributed to the success of this project. I would also like to thank Kylie Williams (SEM illustrations); staff of the Queensland Museum Photographic Section; Darryl Potter for assis- tance with the preparation of illustrations; and Jennifer Mahoney and Vedette McGuire for typing the manuscript. LITERATURE CITED BAKER, H.B. 1941. Zonitid snails from Pacific Is- lands. Pt III and IV. Genera other than Microcys- tinae. Bulletin of the Bernice P. Bishop Museum 166: 205-370. BISHOP, M.J. 1981. The biogeography and evolution of Australian land snails. Pp. 924-954. In Keast, A. (ed.), ‘Ecological biogeography in Australia’. (W. Junk Publ: The Hague, Netherlands). DARTNALL, A.J. & KERSHAW, R.C. 1978. Description of a new species of Helicarion (Stylommalophora: Helicarionidae) in Tas- mania. Records of the Queen Victoria Museum Launceston 62: 1-18. GALLOWAY, R.W. & KEMP, E.M. 1981. Late Caniozoic environments in Australia. Pp. 52^80. In Keast, A. (ed.), ‘Ecological biogeography in Australia’. (W. Junk Publ: The Hague, Nether- lands). IREDALE, T. 1937. A basic list of the land Mollusca of Australia. Australian Zoologist 9: 1-39. KERSHAW, A.P. 1981. Quatemaiy vegetation and environments. Pp. 81-101. In Keast, A. (ed.), ‘Ecological biogeography in Australia’ . (W. Junk Publ: The Hague, Netherlaixls). KERSHAW, R.C. 1979, Redescripiion of Helicarion cuvieri from southern Tasmania and Helicarion freycineti from New South Wales (Pulmonata: Helicarionidae). Journal of the Malacological Society of Australia 4; 145-156. 1981. Redescription of the genus Helicarion and of Helicarion niger (Quoy and Gaimard, 1832) from Victoria (Pulmonata: Helicarionidae). Jour- nal of the Malacological Society of Australia 5: 17-31. 1983. The identity of Helicarion freycineti Ferussac (Mollusca: Pulmonata). Journal of the Malacological Society of Australia 6: 59-61. MARTENS, E.C. VON 1894. Die Mollusken. In Semon, R.W. (ed.), ‘Zoologische Forsch- ungsreisen in Ausiralien und dem Malayischen Archipel mit unterslulzung des Herrn Dr Paul von Ritter ausgefiihrt in der Jahren 1891-1893’. Denschriften der Medizenisch-Naturwis- senschaftlichen GeselLschafl zu Jena 8: 85-96, SMITH, B.J. 1992. Non marine Mollusca. In Houston, W.W.K. (ed.), ‘Zoological catalogue of Aus- tralia’. 8; 1-405, (Australian Government Print- ing Service: Canberra). SOLEM, A. 1966. Some non-marine mollusks from Thailand, with notes on classification of the Heli- carionidae. Spolia Zoologica Musei Hauniensis 24: 1-108. 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: Char- opidae). Journal of the Malacological Society of Australia 8: 1-21 . 10 MEMOIRS OF THE QUEENSLAND MUSEUM DANIELLEILONA GEN.NOV., FROM THE WET TROPICS, NORTHEASTERN QUEENSLAND (PULMONATA:CHAROPIDAE) JOHN STANISIC Stanisic, J. 1993 12 24: Danielleilona gen.nov., from the Wet Tropics, northeastern Queensland (Pulmonata:Charopidae). Memoirs of the Queensland Museum 34(1): 1 1-20 Brisbane. ISSN 0079-8835. A new genus of charopid, Danielleilona gen.nov., is described from the summits of the high mountains in the Wet Tropics, NEQ. Two species, D. marycolliverae sp.nov. and D. multicostata sp. nov., occur in geographically proximate, yet isolated, mountain ranges south of Cairns. The distribution panem is novel in relation to previously described charopids from the region and its biogeographic significance is discussed. The unusual conchological and anatomical features of the genus are related to other Australian charopids. □ Pulmonata, Charopidae, Danielleilona, new genus, D. marycolliverae, D. multicostata, new species, systematics, biogeography. Wet Tropics. John Stanisic, Queensland Museum, PO Box 3300, South Brisbane, Queensland, 4101, Australia. Stanisic (1987) reviewed the status of the charopid genus Oreokera and briefly referred to the diversity of the Charopidae in the Wet Tropics. Traditionally this family has been re- garded as having a temperate distribution with only a few species living in tropical Queensland (Iredale, 1937; Smith, 1984). Field work by staff of the Queensland Museum has revealed a sig- nificant number of species in both subtropical and tropical eastern Queensland (Stanisic, 1990). Danielleilona gen.nov. includes two new, large species of Charopidae from the tops of high mountains south of Cairns, NEQ. These moun- tains are renowned for the restricted floral and faunal elements near their peaks (Monteith 1980). The two land snails are similarly restricted and add to knowledge of this area. This study ex- amines the conchological and anatomical charac- ters of Danielleilona and places them in a biogeographic context. The following abbreviations are used; Q- Queensland; QM-Queensland Museum, Bris- bane; AM- Australian Museum, Sydney; A- anus; DG- prostate; E- epiphallus; EC- epiphal- lic caecum; G-ovotestis; GD-hermaphroditic duct; GG-albumen gland; GT-talon; H-heart; HG-hindgut; HV-hindvein; I-intestine; K-kid- ney; KD-ureter; KX-ureteric pore; MC-mantle collar; P-penis; PC-penial caecum; PCP-penial caecum pore; PP-penial pilaster;PPM-main penial pilaster; PRM-penial retractor muscle; PS-penial sheath; S-spermatheca, SS-sper- mathecal stalk; UT-uterus; UV-free oviduct; V- vagina; VD-vas deferens; X-carrefour; Y- atrium; Z-digestive gland. SYSTEMATICS Family Charopidae Danielleilona gen.nov. Etymology For my eldest daughter Danielle Ilona TYPE SPECIES Danielleilona marycolliverae sp.nov. Diagnosis Shell large, with about 4 V 2 tightly coiled whorls, the last expanding more rapidly. Apex and spire flat to very slightly elevated Apical whorls with low, rounded, rugosely sculptured spiral cords. Postnuclear sculpture of prominent, very strongly protractively sinuated, moderately crowded (multicostata) to widely spaced (marycolliverae), broad radial ribs; apices with radial riblets. Microsculpture of fine, crowded radials and spirals crossing in a regular lattice. Umbilicus deep, narrow U to V-shaped, margins rounded. Sutures strongly channelled. Lip simple with a prominent apertural sinus; aperture round- ly lunate. Genitalia with ovoteslis consisting of two clumps of finger-like lobes oriented parallel to the plane of coiling. Talon with long, coiled shaft. Epiphallus partially enveloped in the penial sheath, with a short caecum and entering penis apically. Penis long with a lateral outpocket (marycolliverae) or a long muscular caecum (multicostata)', internally with longitudinal 12 MEMOIRS OF THE QUEENSLAND MUSEUM pilasters; sheath present. Penial retractor muscle inserting on epiphallus at junction with penial sheath. Free oviduct long. Pallial cavity with complete sigmurethrous ureter. Kidney with large pericardial lobe and rectal lobe reduced to a vestige. Radula (Fig. 8a-f) with long mesocone on tricuspid central and lateral teeth; marginal teeth almost bicuspid with endocone lengthened and minor ectoconal splitting. Comparisons Danielleilona has a number of features which clearly separate it from other genera of Charopidae occurring in the Wet Tropics. These include the large, almost discoidal. flammulated shell, the channelled sutures and apertural sinus together with penial diverticulum and sheath. Oreokera Stanisic, 1987, has a trochoid shell; Biomphalopa Stanisic, 1990, has a biconcave shell; Setomedea Hedley, 1924, has a depressed helicoid shell with periostracal hairs; Hed- leyoconcha Pilsbry, 1893, has a whitish, trochoid shell; and 'Roblinella" intermedia Odhner, 1917 is very small, with a monochrome brown shell. All lack channelled sutures and an apertural sinus. Danielleilona and Oreokera both have spirally sculptured protoconchs but this superficial similarity is not reflected in anatomy, in particular the difference in pallial structures. Unlike Daniel- leilona, Oreokera has an incomplete secondary ureter. Oreokera also shows a right angle orien- tation of the ovotestis, a feature common to few east coast charopids (Stanisic, 1990) and in con- trast to the widespread, parallel condition also seen in Danielleilona. In addition, the optically observed similarity in apical spiral cords is shown to have structural inconsistency when viewed with the SEM. In Oreokera the spirals are much higher than wide and appear as periostracal addi- tions to the protoconch whereas in Danielleilona they are low, wider than high, elevations of the protoconch surface. Conchologically Danielleilona bears a striking resemblance to Roblinella mathinnae (Petterd, 1879) and to a lesser extent Robinella curacoae (Brazier, 1871) from Tasmania (Fig. 9a-0. Fea- tures which these species have in common with Danielleilona include coiling pattern, very strongly protractively sinuated radial ribs, chan- nelled sutures, apertural sinus and spiral protoconch. No soft parts are available in collec- tions to allow rigorous assessment of kinship. Danielleilona marycolliverae sp.nov. (Figs. 1-4) Etymology Named for Mary Colliver in recognition of her voluntary contribution to the Queensland Museum Malacology Section. Comparisons D. marycolliverae differs from D. multicostata in having a shell with more widely spaced radials and larger umbilicus. Anatomically the short penial outpocket of D. marycolliverae contrasts with the long, muscular penial caecum of D. multicostata. Type Material Holotype: QMMO 15803, Mt Bellenden Ker, NEQ. Summit TV Stn, 1560m, rainforest (17°16’S, 145°5rE), 29 Apr 1983, G.B. Monteith, D. Yeales. Height of shell 3.83mm, diameter 6.38nim, DAJ ratio 4.69, H/D ratio 0.60, whorls 4 ^ 2 . Paratypes: Northeastern Queensland: Mt Bartle Frere, summit centre pk, ca 1 540m, SMVFT (1 7°23’S, I45°48’E) (1 dead adult, QMM016473, 7-8 Nov 1981); Ml Bartle Frere, summit north-w'est pk, 1440m (I dead adult, AMCl 50086, 7 Oct 1980, G.B. & S.R. Monteith); Ml Bellenden Ker, summit, 1560m, SMVFT (1 live adult, QMMOl 1263, 25-31 Oct 1981, Earthwatch/QM); Mt Bartle Frere, summit north-west peak, 1440m, SMVFT (I7'*23’S, 145M8’E) (I live adult, QMMOl 1419, 8 Nov 1981, Earthwatch/QM); Mt Bellenden Ker, summit, 1560m, SMVFT (4 live adults, QMMOl 6474, AMCl 50087, 17-24 Oct 1981, Earthwatch/QM); Mt Bellenden Ker, just below sum- mit, approx. 1500m, under bark of rotting logs (6 live adults, QMM014877, 4 Jul 1983, J. Stanisic, D. Pot- ter). Other Material: Mt Bellenden Ker, summit, 1560m, SMVFT (2 dead sub adults, QMMOl 1251, 17-24 Oct 1981, Earthwatch/QM); Mt Bartle Frere, summit centre peak, c.l540m, SMVFT (17°23’S, 145°48’E) (1 broken dead adult, QMMOl 1398, 7-8 Nov 1981, Earthwatch/QM); Ml Bellenden Ker, sum- mit TV Stn. 1560m, SMVFT (17^16'S, 145^51 ’E)(l dead adult, QMMOl 1893, 17-24 Oct 1981, Earthwatch/QM); Ml Bellenden Ker, summit centre peak, litter, (1 dead juvenile, QMMOI6475, 9 Jun 1980, G.B. Monteith): Mt Bellenden Ker, summit TV Stn, 1560m, rainforest (17"16’S, 145^5 rE)(l broken dead sub adult, QMMO15806, 1-7 Nov 1981, Earthwatch/QM); Mt Bellenden Ker, summit TV Stn, 1560m, rainforest (1 specimen, QMMO 16476, 29 Apr 1983, G.B. Monteith, D. Yeates). Diagnosis Shell large (for family), 6.38-7 .23mm (mean 6.64mm) in diameter, with 4V2 tightly coiled whorls; body whorl expanding more rapidly. DANIELLEILONA, A NEW CHAROPID GENUS 13 FIG. 1. Shell of Danielleilona marycolliverae sp. nov. Mt Bellenden Ker, NEQ, QMMO15803, holotype. Scale line = 2mm. Apex and spire flat, last whorl descending more rapidly. Height of shell 3.49-3. 86mm (mean 3.71mm); H/D ratio 0.53-0.60 (mean 0.56). Protoconch (Fig. la) of whorls, 850 pm in diameter at iVi whorls. Apical sculpture (Fig. 2a-c) of 20-25 regularly spaced, nigosely sculp- tured, low, broad, spiral cords. Postnuclear sculp- ture (Fig. 2d-f) of very prominent, high, broadly rounded, rugosely and radially sculptured. strongly protractively sinuated radial ribs, 45-68 (mean 57) on the body whorl, without periostracal blades, with intertices 5-10 times their width. Ribs/mm 2.24-3.39 (mean 2.78). Microsculpture (Fig.2e) of fine radial riblets, 15-30 between each pair of major ribs, crossed by fine, similarly spaced, spiral cords; microsculpture not reduced below the periphery and also continued onto the sides of the major radials. Umbilicus (Fig. lb) narrow V-shaped, regularly decoiling, D/U ratio 4.30-4.81 (mean 4.62). Sutures channelled (Figs la, 2a); whorls rounded above and below the periphery (Fig. Ic). Colour yellowish-light brown with prominent, irregular, darker reddish- brown flammulations. Aperture roundly lunate. Lip simple, retracted at the parieto-palatal margin to form a deep, apertural sinus. Columella weakly reflected; parietal callus weak. Based on 4 measured adults (QMMO15803, QMM016473, AMC 150086, QMMO 16474). Foot and tail broad, bluntly rounded posterior- ly. Body colour (in preservative) creamy white with no obvious markings on foot or neck region; irregular black pigmentation on pallial roof and apical whorls of visceral hump, Ommatophores black. Mantle collar well developed, without mantle gland. Pallial region (Fig. 30 about V 2 whorl apically; kidney twice as Ihng as broad, with prominent pericardial lobe and vestigial rec- tal lobe. Ureter sigmurethrous with complete secondary ureter and widely diverging arms. Ovotestis (Fig. 3e) two clumps of finger-like lobes embedded in the apical whorls of the diges- tive gland and oriented parallel to the plane of coiling. Talon (Fig. 3c) with long coiled stalk, embedded in the albumen gland. Prostate-uterus typical. Vas deferens a thin tube entering epiphal- lus apically. Epiphallus, thick, muscular, reflexed, partially enveloped in penial sheath; with a small diverticulum at flexure; entering penis apically through a simple pore; internally, ascending branch with longitudinal pilasters, des- cending branch with small pustules. Penis (Fig. 3b, d) large, sub-cylindrical, with a sub-apical out- pocket containing a terminal, pocket-like, fleshy pilaster and central, longitudinal pilaster which continues into the penis proper. Penis interior (Fig. 3b) with main corrugated longitudinal pilaster and many secondary, low, longitudinal thickenings. Free oviduct longer than vagina. Vagina short, internally with simple longitudinal thickenings. Spermalheca typical. Based on four dissected specimens (QMM014877, QMMO- 11419, QMM016474). 14 MEMOIRS OF THE QUEENSLAND MUSEUM Range and Habitat Known only from the summits of Mt Bellen- den Ker and Mt Bartle Frere (Fig. 4) in the Bellenden Ker Ra., in simple microphyll vine- fern thickets and forests. Collected above the ground under the bark of rotting logs suggesting semi-arboreality. Remarks D. marycolliverae has been collected FIG. 2. Shell features of Danielleilona marycollherae sp. noy. Mt Bellenden Ker, NEQ, QMMO 1 6476, paratype. a, protoconch showing channelled suture; b-c, details of protoconch sculpture; d, postnuclear sculpture; e-f; details of postnuclear microsculpture and major rib structure. Scale lines as marked. DANIELLEILONA, A NEW CHAROPID GENUS 15 FIG. 3.Anatomy of Danielleilona marycolliverae sp. nov. Mt Bellenden Ker, NEQ, QMM014877, paratype. a, reproductive system; b, penis interior; c, details of talon and carrefour region; d, penis; e, ovotestis; f, pallial cavity. Scale lines = 1mm. microsympatrically with Oreokera cumulus (Odhner, 1917) on Mt Bellenden Ker (QMMO- 14877). Danielleilona multicostata sp.nov. (Figs. 4-8) Etymology From the Latin multi many; costa - rib; refer- ring to the adult sculpture of crowded, radial ribs. Comparisons D. multicostata bears an overall similarity to D. marycolliverae but is marginally smaller with narrower umbilicus, and has more crowded ribs on the body whorl (Figs. 1,5). Type Material Holotype: QMM033817, North Bell Pk, Malbon Thompson Ra., NEQ, 800-900m ( 17°05’S, 145°53T). 19-22 Nov 1990, G.B . Monteith, G. Thompson. Height of shell 3.78mm, diameter 5.71mm, D/U ratio 6.80, H/D ratio 0.66, whorls 4^/$-, Paratypes: North Bell Pk, 20Ian S Cairns, NEQ, 900-1000m (2 dead adults, 1 dead sub-adult, QMMO15490, 15-16 Sept 1981, G.B. Monteith, D. Cook). Diagnosis Shell large (for family), 5.7 1-5. 96mm (mean 5.88mm) in diameter with 4V4-4V8- (mean 4*4-i-) tightly coiled whorls; body whorl expanding more rapidly. Apex and spire flat, last whorl descending rapidly in front. Height of shell 3.49- 3.78mm (mean 3.61mm); H/D ratio 0.59-0.66 (mean 0.62). Protoconch (Fig. 6a) of P/4 whorls, 850|im in diameter at 1 V 2 whorls. Apical sculp- ture of 18-22 regularly spaced, rugose, spiral cords. Postnuclear sculpture of prominent. FIG. 4.Distribution of Danielleilona marycolliverae sp. nov. (dots) and D. multicostata sp. nov. (diamond) in the Wet Tropics region. 16 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. S.Shell of Danielleilona multicostata sp. nov. North Bell Peak, NEQ, QMM033817, holotype. Scale in mm. crowded, strongly protractively sinualed, radial ribs, 93-108 (mean 101) on the body whorl, with interstices 5-7 times their width. Ribs/mm 5.18- 5.77 (mean 5.47). Microsculpture of fine radial riblets, 12-20 between each pair of major ribs, crossed by fine spiral cords to form a reticulate pattern; sculpture continuous on base. Micros- culpture (Fig. 6b) continuous on sides of major radials. Umbilicus (Fig. 5b) very narrow, U- shaped, barely decoiling. D/U ratio 5.38-6.80 (mean 6.00). Sutures channelled; whorls rounded above and below the periphery. Colour light yel- low-brown with irregular, darker, brown suf- fusions. Aperture roundly lunate. Lip simple, retracted at the parieto-palatal margin to form a conspicuous apertural sinus. Columella slightly reflected over the umbilicus. Parietal callus present. Based on 3 measured adults (QMMO- 33817, QMMO15490). External animal features similar to D. rnarycol- liverae. Pallial cavity with typical kidney and complete sigmurethrous ureter. Apical genitalia not studied. Terminal genitalia (Fig. 7a) similar to that of D. marycolliverae except for penis morphology. Penis (Fig. 7b) with long, muscular caecum; internally with simple, low longitudinal pilasters. Penial caecum entering penis subapical- ly about V'i of the way down through a mus- cularised pore. Penis sheath present. Radula (Fig. 8a-0 with tricuspid central and lateral teeth; marginals muiticuspid with enlarged endocone, ectocone split into minor cusps. Based on one dissected specimen (QMMO- 33817). Range and Habitat Known only from 4 specimens collected from the top of North Bell Pk on the Malbon Thompson Ra., NEQ (Fig. 4), in simple microphyll vine- fern forest. No microhabitat data is available. FIG. 6.Shell sculpture of Danielleilona multicostata sp. nov. North Bell Peak, NEQ, QMMO15490, paratype. a, apical sculpture; b, postnuclear micros- culpture. Scale lines as marked. DANIELLEILONA, A NEW CHAROPID GENUS 17 FIG. T.Anatomy of Danielleilona multicostata sp. nov. North Bell Peak, NEQ, QMM033817, holotype. a, terminal genitalia; b, penis interior. Scale lines = Imm. Remarks Although this species is known from very limited material, shell and anatomical characters readily separate it from D. marycolliverae. Discussion The discovery of Danielleilona highlights the unusual nature of the mountain top communities in tropical northeastern Queensland. Previously, Stanisic (1987, 1990) provided examples of land snails from these environments which displayed both narrow and widely disjunct generic distribu- tions. This is the first example involving the Bellenden Ker and Malbon Thompson Ranges. Summits of their higher peaks are characterised by the occurrence of simple microphyll vine-fern forest (SMVFF). This grades into thicket (SMVFT) on the higher mountains of the Bellen- den Ker Ra. Presumably these cool moist, largely temperate mountain top communities would have had a more widespread distribution involving the lowlands in cooler, wetter times. Climatic change, particularly in the Plio-Pleistocene, had a dramatic impact on ‘temperate* communities (Galloway & Kemp, 1981) leading to their present-day restriction in the north. Temperate communities remained only on those peaks with the high altitudes necessary to preserve them, and extinctions undoubtedly occurred on lower peaks. A key ingredient in the survival of Daniel- leilona appears to be the presence of SMVFF on the Bellenden Ker and Malbon Thompson Ran- ges. However the absence of SMVFT from the latter could be the crucial factor in the disparate distribution of Oreokera Stanisic, 1987, which is sympatric with Danielleilona in the Bellenden Ker Ra., but missing from the Malbon Thompson Ra, The morphological gap between species of Danielleilona is greater than that recorded for species of Oreokera which may indicate a more stringent environmental and climatic selection regime on North Bell Pk montane communities. Some species of Charopidae, (e.g. Setomedea monteithi Stanisic, 1990), are found in the warm humid mesophy 11 vine forests of the lowlands and foothills of the Wet Tropics, but most are con- fined to the uplands and highlands. Those now found in lower areas also would probably have depended on montane refugia for survival in the past, but have been able to disperse rapidly in recent times (Stanisic, 1990). Significantly, Setomedea has species in central and southern Queensland, indicating wide environmental 18 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. S.Radula of Danielleilona multicostata sp. nov. North Bell Peak, NEQ, QMM03381 7, holotype. a, central and inner lateral teeth; b. laterals and lateromarginals; c, lateromarginals showing squarish basal plate; d, laterals and first lateroniarginal showing ectoconal reduction; e-f, lateromarginals and marginals showing ectoconal reduction and ectoconal splitting. Scale lines as marked. tolerance. The specialised microhabilat of and anatomical characters which place it outside Danielleilona (semi-^boreal) probably reflects a mainstream of charopid evolution in tropical higher degree of environmental sensitivity than , * • i ^ , , rr^ , for generalist ‘under-the-log’-dwelling species subtropical eastern Queensland. The large such as Setomedea. shell with channelled sutures, apertural sinus and displays a suite of conchological very strongly, protractively sinuated radial ribs. DANIELLEILONA, A NEW CHAROPID GENUS 19 FIG. 9.a-c, Roblinella mathinnae (Petterd, 1879). Tasmania, AMC 103794. a, shell; b, proloconch sculpture; c, postnuclear sculpture, d-f, Roblinella curacoae (Brazier, 1871). Tasmania, AMC 1035 18. d, shell; e, protoconch sculpture; f, postnuclear microsculpture. Scale lines as marked. together with the penial sheath and penial diver- ticulum are major differentiating features. Tht occurrence of a number of very similar shell features in Tasmanian taxa (Fig. 9a-0 may have important biogeographic implications. Southern relationships have been demonstrated in a number of Wet Tropics charopid genera by Stanisic (1990), but a Tasmanian connection would span a geographic gap shared by plants {O rites), the Peloridiidae (moss bugs) and the stag-beetle genus Lissapterus. These are all ‘antarctic' relicts (Monleith, 1980) occurring in montane refugia in subtropical and tropical areas. Stanisic (1990) showed that protoconch sculp- 20 MEMOIRS OF THE QUEENSLAND MUSEUM ture and adult microsculpture were useful shell characters for indicating relationships among the Charopidae. A spirally Urate protoconch is rare in tropical and subtropical Australian species. Most have radially ribbed or reticulate protoconchs. The presence of spirally Urate protoconchs (Fig. 9b, c) and similar shell microsculpture (Fig. 9c, 0 in R. tnathinnae and R. curacoae may not be sufficient to indicate possible relationships. How- ever, combined with the co-occurrence of an apertural sinus, strongly protractively sinuated radial ribs and channelled sutures (Fig. 9a,d) there is a tantalising prospect that these taxa are more closely related than they are to others in the family. ACKNOWLEDGEMENTS Part of the material was collected on fieldwork sponsored by the Earthwatch organisation. This support is gratefully acknowledged, as are the collecting efforts of Dr Geoff Monteith, Queensland Museum and his many co-workers. Thanks are due also to Kylie Williams, for the excellent SEM illustrations which were processed by the Queensland Museum’s Photog- raphy Section; Darryl Potter, who assisted with preparation of illustrations; and Jennifer Mahoney and Vedette McGuire, who typed the manuscript. LITERATURE CITED GALLOWAY, R.W. & KEMP, E.M. 1981. Late Cainozoic environments in Australia. Pp.52-80. In Keast, A. (ed.), ‘Ecological biogeography in Australia*. (W. Junk PubI: The Hague, Nether- lands). IREDALE, T. 1937. A basic list of the land Mollusca of Australia. Australian Zoologist 8(4): 287-333. 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 In- sects 21(4): 275-285. SMITH, B.J. 1984. Regional endemism of the south eastern Australian land mollusc fauna. Pp. 178- 188. In Solem, A. & Van Bruggen, A.C. (eds.), ‘World wide snails. Biogeographical studies on non marine Mollusca’. (E.J. Brill: Leiden). 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). Journal of the Malacological Society of Australia 8: 1-21. 1990. Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pulmonata) from subtropical rainforests. Memoirs of the Queensland Museum 30(1): 1-241. LENWEBBIA PALUMA SP. NOV., FROM THE WET TROPICS, NORTHEASTERN QUEENSLAND (PULMONATA: CHAROPIDAE) JOHN STANTSIC Stanisic, J. 1993 12 24:Lenwebbia paluma sp. nov., from the Wet Tropics, northeastern Queensland (Pulmonata: Charopidae). Memoirs of the Queensland Museum 34(1): 21-26. Brisbane. ISSN 0079-8835. A new species of charopid, Lenwebbia paluma sp. nov., is described from the Paluma Range, in the south of the Wet Tropics, NEQ. This discovery establishes a widely disjunct north-south distribution for the genus. The biogeographic significance of this pattern is discussed in relation to rainforest evolution and the specific microhabitat preference of the species. □ Charopidae, Lenwebbia paluma, systematics, biogeography, Wet Tropics. John Stanisic, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; 10 November, 1993, The charopid fauna of the Wet Tropics, NEQ, is poorly known. Only six species have been described (Odhner, 1917;Hedley, 1924; Stanisic, 1987, 1990). Their small size and limited vagility, and relatively large numbers in eastern Australia make them a potentially powerful tool in biogeo- graphy, Stanisic (1987, 1990) has highlighted a number of unusual aspects of the distributions and relationships of several Wet Tropics char- opids. The discovery of a new species of Lenwebbia from Mt Spec in the Paluma Range, NEQ, provides an additional opportunity to examine charopid distribution patterns in relation to the evolution of mesic communities in eastern Aus- U-alia. The following abbreviations have been used in the text: NEQ, northeastern Queensland; QM, Queensland Museum. SYSTEMATICS Family Charopidae Lenwebbia Stanisic, 1990 Lenwebbia Stanisic, 1990, p.52. Type Species Lenwebbia protoscrobiculata Stanisic, 1990, by original designation. Remarks Lenwebbia is a distinctive genus of charopid characterised by its greatly reduced shell sculp- ture, and is unlikely to be confused with other nordieastem Queensland members of the family. It could be mistaken for some of the Wet Tropics Helicarionidae, such as Tarocystis Iredale, 1937, and Expocystis Iredale, 1937, which also have macroscopically featureless shells. However, these genera differ microscopically from Len- webbia in having incised spirals on the protoconch and notched incised grooves on the postnuclear whorls. They also lack the impressed sutures of Lenwebbia. The features of L. paluma sp. nov. are consistent with the generic diagnosis given by Stanisic (1990). Lenwebbia paluma sp. nov. (Figs. 1-4) Etymology For the type locality, Paluma, NEQ. Comparative Remarks The shell of L. paluma is grossly similar to that of L. protoscrobiculata Stanisic, 1990, from southeastern Queensland. Differences in L. paluma include a smaller umbilicus (D/U ratio 10.5-14.9 compared with 5.34-5.62 for proto- scrobiculata) and finer microsculpture that does not include spiral grooves. The shell of L. paluma is also slightly smaller and flatter, due mainly to a smaller body whorl width. Anatomically, the unequal length of the penial pilasters in L paluma contrasts with the condition in L. proto- scrobiculata (Stanisic, 1990, Fig. 30b). Type Material Holotype: QMM043311, Mt Spec NP, c. 0.5k E Paluma, NEQ (19°0rS, 146°13’E), rainforest, 800m, under bark of trees. Collected by J. Stanisic, D. Potter, 6 May, 1990. Height of shell 2.68mm, diameter 4.30mm, H/D ratio 0.61, D/U ratio 14.6, whorls 4V$. 22 MEMOIRS OF THE QUEENSLAND MUSEUM Paratypes: 2 adults, 5 subadults, QMMO29085, same collection data as holotype. Diagnosis Shell (Fig. la,b) small, glossy, turbinate, diameter 4. 30-4. 47mm (mean 4.38mm) with 4^8 evenly coiled whorls, the last descending slowly in front. Apex and spire moderately elevated, SP/BWW ratio 0.29-0.44 (mean 0.38), height 2.68-3. 06mm (mean 2.82mm). H/D ratio 0.61-0.69 (mean 0.64). Body whorl width 1.45- 1.53mm. Apical sculpture (Fig. 2a) of crowded dimples. Protoconch of whorls, mean dia- meter 939|J-m. Post-nuclear sculpture (Fig. 2b-d) of weak radial growth ridges, fine crowded threadlike radial ridgelets and irregular spiral cords; sculpture reduced on the base. Umbilicus (Fig. lb) very narrow, partially covered by reflec- tion of the columella, diameter 0.3 1-0.4 1mm (mean 0.38mm), D/U ratio 10.5-14.9 (mean 13.3). Sutures impressed. Whorls rounded above and below the periphery. Lip simple, columella dilated. Parietal callus prominent. Colour brown. Based on three measured adults (QMMO29085, QMM043311). Animal colour (in life) grey, becoming darker on the mantle collar, neck, top of foot and in the region between the pedal grooves. Foot short and broad. Pallial cavity (Fig. 3c) with well developed mantle collar (MC) and prominent mantle gland (MG). Pallial roof with black speckling. Kidney (K) subtriangular, orangy-brown with an incom- plete sigmurelhrous ureter (KD), Ureteric pore (KX) situated in the angle between the kidney and the rectum (HG). Hindvein (HV) prominent. Genitalia (Fig, 3a) with enlarged albumen gland (GG). Ovotestis (Fig. 3b, G) consisting of several white clumps of alveoli located in the apical whorls of the digestive gland (Z) and oriented at right angles to the plane of coiling. Epiphallus (E) short, stout, entering penis apically through two fleshy lips (EP). Penis (P) cylindrical, tapering toward atrium (Y), internally (Fig. 3d) with a large fleshy longitudinal pilaster and a shorter, apical, secondary pilaster (PP). Vagina (V) short. Free oviduct (UV) long. Spermatheca (S) typical, stalk (SS) long. Radula with tricuspid central and lateral teeth (Fig. 2e) in which the mesocone is long and lanceolate; anterior flare present. Mar- ginal teeth (Fig. 2f) multicuspid with endocone and ectoconc split into several minor cusps. Basal plates squarish, with a central ridge that articu- lates with the anterior flare of the tooth behind. Based on two dissected specimens (QMMO- 29085). FIG. 1. Shell of Lenwebbia paluma sp.nov. Ml Spec NP, 0.5km E Paluma, NEQ, QMM0433 1 1, holotype. Scale line = 3mm. Range and Habitat Known from only the type locality near Paluma, NEQ (Fig. 4). Cooler peaks of this area support rainforest in contrast to the drier eucalypt forest of the foothills. The peaks are not high when compared with mountains of the Wet Tropics, and the altitudinal stratification of rain- forest communities is less pronounced. Upper reaches support simple notophyll vine forest [ which grades into mesophyll vine forest in warmer sheltered gullies. L. paluma lives under the bark of standing live or dead trees. Remarks L. paluma has a specialised microhabitat com- pared with most eastern Australian charopids (in- | eluding L. protoscrohiculata) which live under I, logs. The flatter shell (when compared with L. protoscrohiculata) is probably an adaption for this lifestyle. This selective microhabitat may account for the comparative rarity of the species , in collections. However, considerable collecting [ effort in the moister parts of the Wet Tropics, north of the Herbert R., makes it unlikely that the genus has representatives in tho.se forests. Con- j versely its presence in areas such as the Seaview Ra., Bakers Blue Mtn and Bluewater Ra. cannot I be discounted until additional collecting is under- | taken. | DISCUSSION ’ The north-south, widely disjunct distribution of I Lenwebbia has parallels among other charopid j genera {Setomedea Iredale, 1941; Hedleyocon- cha Pilsbry, 1893; Biomphalopa Stanisic, 1990). AH have species in the Wet Tropics with nearest relatives either in mideastem or southeastern Queensland. However, in contrast to Lenwebbia^ their northern representatives occur in the north- ern, moister parts of the Wet Tropics, between NEW RAINFOREST SNAIL, LENWEBBIA PALUMA 23 FIG. 2. Shell and radular detai Is of Lenwebbia pahima sp. nov . Mt Spec NP , 0.5km E Paluma, NEQ, QMMO29085 , paratype.a, protoconch sculpture; b, adult sculpture; c-d, post nuclear microsculpture; e, central and inner lateral teeth; f, marginal teeth (note maIformation).Scale lines as marked. Ingham and Cooktown, NEQ. The restriction of altitude and closer proximity to the sea, and L. paluma to the southern, drier end is unusual in higher areas have a mean annual rainfall greater this context. than 2000mm. The main vegetation type is simple The Paluma Ra. is a southern outlier of the main notophyll vine forest on the summits and Wet Tropics rainforest massif. It is wetter than the mesophyll vine forest at lower elevations (Nix, more northerly SeaviewRa. because of its greater 1991). The ‘temperate summit communities 24 MEMOIRS OF THE QUEENSLAND MUSEUM MG HV E PRM MC FIG. 3. Anatomy of Lenwebbia palutna sp.nov. Mt Spec N.P., 0.5k E Paluma, NEQ. QMMO29085, paratype. I a, reproductive system; b, ovotestis; c, pallial structures; d, penis interior. Scale lines = 1mm. found on mountains further north are absent and Nix (1991) considered that the southern outliers of the Wet Tropics would not have provided significant refuges during the last glacial. Rainforest in eastern Australia experienced its major retreat in the Pliocene when plant genera such as Eucalyptus gained ascendancy and oc- cupied the drier areas vacated by rainforest vegetation (Galloway & Kemp 1981). Major north-south disjunctions evident today in eastern Australian biota probably date from this epoch. The establishment of the Fitzroy and Burdekin dry corridors (Nix, 1991) fragmented populations of formerly widespread mesic communities. Land snails would have suffered considerable range attrition during the drying of the late Ter- tiary and Quaternary because of their extreme ■ moisture sensitivity and low vagility. In the Wet Tropics region, temperate and subtropical com- | i munities were isolated in upland refugia and, in 1 1 spite of Holocene ‘wet-phases’ (Kershaw & Nix, 1988), did not re-emerge because of potent NEW RAINFOREST SNAIL, LENWEBBIA PALUMA 25 bioclimatic barriers operating at lower elevations (Nix, 1991). Lenwebbia is a significant element of these environmentally-cornered assemblages and the restriction of Lenwebbia in the north is inextricably linked with the fragmentation and reduction of rainforest communities since the mid-Miocene. Nix (1991) has shown a strong association be- tween bioclimates and rainforest structural types. The bioclimatic parameters used (radiation, temperature, moisture and rainfall) translate directly to the critical factors in land snail sur- vival, So it is not surprising that the distribution of rainforest structural types (Webb, 1968) and land snail assemblages (communities) show great coincidence (Stanisic, unpubl.). As indicators of past and present climatic and edaphic profiles rainforest types provide a framework for under- standing land snail evolution in eastern Australia. The drying events of the Pliocene, and their more radical effects in the Pleistocene (Galloway & Kemp, 1981) most likely caused the initial frag- mentation of Lenwebbia. Climatic changes in the last 100,000 years which also caused major vegetation changes in eastern Australia (Ker- shaw, 1981) probably attenuated the morphologi- cal disparities between the northern and southern species. In the south, L. protoscrobiculata Stanisic, 1990 is associated with comparatively drier sub- tropical araucarian microphyll vine forests. It is absent from the wetter refugia of the Border Ran- ges, which suggests long-term adaptation to a subtropical seasonally moist/dry rainforest type where mean annual rainfall is 900-1 100mm (Webb & Tracey, 1981). The araucarian forests which flourish in the south are now only relicts in the north. Pollen sampling on the Atherton Tableland indicates that they were replaced by drier sclerophyll vegetation during the last glaci^ (380(X)- 26000 BP) (Kershaw, 1981). Dry condi- tions reached a climax at 17000 BP and persisted until KKXX) BP. Prior to this the araucarian forests persisted under a rainfall regime of about 1200mm per annum which is not unlike their present-day requirements in the south. The elimination of this forest type in all but refugia in the north reflects a climatic shift which probably caused significant changes in land snail com- munities including numerous extinctions. The isolation of L. paluma in the Paluma Ra. from the main mass of rainforest further north probably occurred during this period. It is likely that the distribution of Lenwebbia in the north was more extensive. (dot). Although some recolonisation of areas outside the Paluma Ra. by Lenwebbia was possible when major rainforest units and outliers were united during the cool, wet phase of the Holocene (7500- 6000 BP) (Kershaw & Nix, 1988; Nix, 1991) this does not appear to have occurred. L. paluma shows a greater degree of micro- habitat specialisation than L. protoscrobiculata . Its flatter shell with smaller body whorl width is an expression of this major shift to life under bark of trees. This microhabitat specialisation is a like- ly contributing factor to the current restricted distribution of Lenwebbia in the north. Relatively exposed microhabits and specialised microhabit- ats presumably have greater probability of pertur- bation by climatic change. Hedleyoconcha, Lenwebbia and Setomedea all have widespread species in subtropical southeastern Queensland. Hedleyoconcha lives in exposed situations on the 26 MEMOIRS OF THE QUEENSLAND MUSEUM leaves, trunks and branches of shrubs and trees. Significantly, Hedleyoconcha survives only in the very wet, cloudy moist summits of the Wet Tropics and is absent from the mideastern Queensland rainforests. In contrast, Setomedea, which lives under logs where ambient conditions are considerably more stable, has managed to persist in mideastem Queensland, even in the somewhat drier habit of Mt Diyander. The Wet Tropics sj^ies, S. monteithi Stanisic, 1990, is comparatively widespread in the uplands, foothills and lowlands (mountain-mass effect?) between Innisfail and Cooktown. In comparison, Lenwebbia is absent from mideastem Queens- land and has a narrow distribution in the Wet Tropics, a situation not unlike that seen in Hed- leyoconcha, Microhabitat specialisation in L. paluma may have resulted from the need to adapt to habitat change. However, in the process the species has probably increased its sensitivity to climatic changes. As an environmentally-cornered species, L paluma must be regarded as a key indicator species of the Paluma biogeographic unit of Nix (1991). ACKNOWLEDGEMENTS Thanks are due to Kylie Williams for the SEM illustrations, to the Queensland Museum Photog- raphy Section for processing and to Jennifer Mahoney for typing the manuscript. LITERATURE CITED GALLOWAY, R.W. & KEMP, E.M. 1981. Late Cainozoic environments in Australia. Pp.52-80. In Keast, A. (ed.), ‘Ecological biogeography of Australia*. (W. Junk PubI: The Hague, Nether- lands). HEDLEY, C. 1924. Some notes on Australian land shells. Australian Zoologist 3: 215-222. KERSHAW, A.P. 1981. Quaternary vegetation and environments. Pp. 81-101. In Keast, A. (ed.), ‘Ecological biogeography of Australia’. (W. Junk PubI: The Hague, Netherlands). KERSHAW, A.P. & NDC, H.A., 1988. Quantitative palaeoclimatic estimates from pollen data using bioclimatic profiles of extant taxa. Journal of Biogeography 15: 589-602. NIX, H.A. 1991. Biogeography: pattern and process. Pp. 1 1-42. In Nix, H.A. & Switzer, M.A. (eds), ‘Rainforest animals’ Kowari 1. (Australian Na- tional Parks and Wildlife Service Publication: Canberra). ODHNER, N.H. 1917. Mollusca. XVII. Results of Dr E. Mjobergs Swedish scientific expedition to Australia 1910-1913. Kungliga Svenska Ven- tenskapsakademiens Handlingar Bd 52, No 16, 1-115. STANISIC, J. 1987. Studies on the Charopidae of tropical and subtropical Australia. I. Oreokera: A pri mitive genus from the high mountains of north Queensland (Mollusca : Pulmonata: Charopidae). Journal of the Malacological Society of Australia 8: 1-21. 1990. Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pulmonata) from subtropical rainforests. Memoirs of the Queensland Museum 30(1): 1-241. WEBB, L.J. 1968. Environmental relationships of the structural types of Australian rainforest. Ecology 49(2): 296-311. WEBB, L.J. & TRACEY, J.G. 1981. Rainforests: pat- terns and change. Pp. 606-694. In Keast, A. (ed.), ‘Ecological biogeography of Australia’. (W. Junk PubI: The Hague, Netherlands). EUNGARION MCDONALDI GEN. ET SP.NOV., A MONTANE SEMI-SLUG FROM MIDEASTERN QUEENSLAND RAINFORESTS (PULMONATA: HELICARIONIDAE) JOHN STANISIC Stanisic, J. 1993 12 24: Eungarion mcdonaldi gen. el sp.nov., a montane semi-slug from mideastem Queensland rainforests (Pulmonata: Helicarionidae). Memoirs of the Queensland Museum 34(]):27-34. Brisbane. ISSN 0079-8835. Eungarion mcdonaldi gen. et sp.nov. is described from the montane rainforests of mideastem Queensland. £ mcdonaldi has a suite of unusual features which are discussed in relation to other eastern Australian semi-slugs and eastern Australian rainforest biogeography. □ Pulmonata, Helicarionidae, Eungarion mcdonaldi, semi-slug, systematics, biogeography, rainforest. John Stanisic, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; 8 November, 1993. The rainforests between Sarina and Proserpine, mideastem Queensland (MEQ) are significant land snail habitats. Many species are endemic to the region, and some are restricted to the moist uplands. Indications are that these montane habitats were important refugia for land snails in drier times (Stanisic, 1990). Snails are particular- ly moisture sensitive animals and utilise their shells primarily for protection from desiccation. Semi-slugs have reduced shells which have developed in response to stable, high-moisture levels, and a comparatively low calcium environ- ment (Solem, 1978). Eastern Australia provided favourable habitats for semi-slug evolution during the late Tertiary and Quaternary and, today, helicarionid semi-slugs are particularly diverse in this region. The species display varied degrees of shell reduction, ranging from a reduced shell which is complete and still provides shelter for the animal, e.g. Helicarion rubicundus of Tasmania (Dartnall & Kershaw, 1978) to a flat, plate-like, almost internalised shell (e.g. Parmacochleafischeri Smith, 1884, of northeastern Queensland). Between these i extremes are a number of species with highly membranous, lenticular shells in which the in- ternal walls of the early shell whorls are incom- plete, Eungarion mcdonaldi gen. et sp.nov. from the summit refugia of the Clarke Range, MEQ, is one of these. This paper details the morphological features of E. mcdonaldi and relates them to other east- ern Australian semi-slugs. It also places the species in biogeographic context. The study is based on specimens in the Queensland Museum (QM). SYSTEMATICS Helicarionidae Eungarion gen. nov. Previous Studies The only semi-slug previously described from MEQ is Fastosarion superba (Cox, 1871). It was described from Mt Dryander but, according to Queensland Museum records, also occurs in the Clarke Ra. Iredale (1933) created the monotypic Fastosarion for this species on the basis of shell features. Smith (1992) considered that Fas- tosarion included Vercidarion Iredale, 1933 (type species: Helicarion bullaceus Odhner, 1917) and also added to it several subtropical and tropical species. The extent of the Fastosarion radiation needs rigorous testing by revisionary studies, but unpublished work (R.C. Kershaw, pers. comm.) shows that at least F. superba and F. bullaceus may be closely related. DIAGNOSIS Shell small, glossy, poorly calcified with membranous base and incomplete internal whorls. Shell sculpture obsolete. Animal with long and slender foot; caudal horn prominent. Shell lappets and mantle lobes well developed. Genitalia with epiphallic caecum and flagellum. Penis with sheath, internally pustulose and with conical verge. Spermatophore with a thin chitinous tail, otherwise without ornamentation. Radula with increased numbers of marginal teeth. Etymology From Eungella (the township near the type locality) and helicarion (the epithet most com- monly applied to this group of semi-slugs). 28 MEMOIRS OF THE QUEENSLAND MUSEUM Type Species Eungarion mcdonaldi sp.nov. Comparisons Reduced shell size, absence of shell sculpture and the degenerate nature of the shell (poor cal- cification, incomplete early shell whorls, membranous shell base) are features which separate Eungarion from Fastosarion Iredale, 1933. The pustulose internal penial sculpture of Eungarion (Fig. 4c) contrasts with the pattern of obliquely arranged, thin longitudinal pilasters seen in Fastosarion superba (Cox, 1871) (Ker- shaw, pers.comm.). Eungarion shows a similar level of shell reduction to that seen in the north- eastern Queensland Thularion Stanisic, 1 993, but differs in key shell features such as sculpture, size and degree of calcification. Anatomically, the much stronger development of shell lappets and mantle lobes, lack of a prominent caudal horn and absence of a penial verge in Thularion are key differentiating features. Eungarion mcdonaldi sp.nov. (Figs 1-5) Etymology Named for Keith McDonald, Conservation Strategy Branch, Queensland Department of En- vironment and Heritage, for his invaluable assis- tance in providing specimens and field data on Queensland land snails. Comparative Remarks Eungarion mcdonaldi (Fig. 2) is readily distin- guished from the sympatric Fastosarion superba (Cox, 1871) by its smaller size and degenerate shell. F. superba is one of the largest semi-slugs in eastern Australia (approx, animal length = 50- 70mm) and has a less reduced shell with prominent spiral sculpture. In the juvenile state the shell features are also useful discriminating features. Material Examined Holotype: QMMO43309, Daliymple Heights, Eun- geIlaNP,MEQ.2r02’S, 148°36’E. 1000m,nolophyll vine forest. M.J. Bishop, Nov 1976. Height of shell 4.81mni, max. diameter 1 1.30mm, min. diameter 8.31mm, H/D ratio 0.43, whorls 'PA-. Paratypes: MEQ: QMMO43310, 35 juveniles and adults, same collecting data as holotype; slopes of Mt Dalrymple, W of Mackay, 900m, on dead leaves, simple notophyll evergreen vine forest (2T04’S, 148°35’E) (5, QMMO36087, 21 May 1990, D. & N. Potter, J. Stanisic); slopes of Mt Macartney, SW Proserpine, 900m, in discarded palm fronds, notophyll vine forest/tree ferns (20°50’S, 148°33.5’E) (1, QMM035628, J. Stanisic, D. & N. Potter. 18 May 1990); Mt Dalrymple, Eungella NP, in cave in rain- forest (1, QMMO43307, V. Hansen, 19 Aug. 1975). Diagnosis Shell small, glossy, maximum diameter 9.74- 11. 30mm (mean 10.46rTun), minimum diameter 7.14-8.31mm (mean 7.76mm), lenticular, very thin, poorly calcified. Whorls 2 V%, rapidly ex- panding. Body whorl expanded, not descending in front, with basal margin membranous. Spire and apex flat. Height of shell 3.25-4.8 1mm (mean 4.26mm), H/D ratio 0.33-0.44 (mean 0.41). Protoconch (Fig. 3a) of 1 V 4 whorls, smooth ex- cept for ve^ faint growth ridges. Post-nuclear sculpture (Fig. 3b) of weak, arcuate, radial growth ridges. Sutures weakly impressed to flat. Whorls rounded above the periphery, with internal walls, membranous and incomplete. Colour yellow to golden, apex creamy-white. Based on 4 measured adults (QMMO43309, QMMO43310). FIG. 1. Holotype of Eungarion mcdonaldi sp.nov., QMM0433()9. a, top view; b, bottom view. Scale in mm. A NEW MONTANE SEMI-SLUG 29 Animal (Fig. 4a) small, body length (in preserv- ative) 24.l-33.7mm (mean 29.9mm, n=5). Foot long, slender, tripartite, broadly rounded anterior- ly, tapered posteriorly. Tail long, keeled mid-dor- sally, with a prominent caudal horn. Caudal foss a long vertical slit in tail. Pedal grooves typically aulacopod and united above the tail. Colour (in live) radish-brown on foot, neck, tentacles, and mantle lobes; visceral hump yellow; shell lappets pale with darker reddish-brown circular mark- ings, also present in preserved specimens. Body ornamentation strongly develop^ on tail and in neck region; shell lappets with widely spaced subcircular pustulations, mantle lobes smooth. Right mantle lobe (RML) small, subtriangular, fused to right shell lappet posteriorly; left mantle lobe (LML) large, taper^ posteriorly, weakly developed anteriorly and fused to left shell lappet. Right shell lappet (RL) large, subcircular, weakly fused with left shell lappet posteriorly; left shell lappet (LL) semi-circular, fused to narrow mantle collar anteriorly. Genitalia (Fig. 4b) with multilobate, creamy- coloured ovotestis (G) embedded in the apical whorls of a dark-brown digestive gland (Z). Her- maphroditic duct (GD) swollen and convoluted. Talon (GT) with spherical head and short stalk. Carrefour (X) sub-ovoid, white. Albumen gland (GG) a multilobate mass of creamy-brown acini. Prostate (DG) a band of creamy-white alveoli appressed to the surface of a sacculate uterus (UT). Vas deferens (VD) arising from the prostate - uterus as a thin tube, descending to the penioviducal angle then ascending a short dis- tance before entering a much thicker, muscular reflexed epiphallus (E). Epiphallus longer than penis, with a long, apical flagellum (EF) and a large epiphallic retractor caecum (ERC) arising mid-length; internally with longitudinal rib-like thickenings below level of caecum, pustulose above. Penial retractor muscle (PRM) short, at- tached to epiphallus adjacent to caecum. Penis (P) relatively short, cylindrical, muscular with a thick sheath (PS) surrounding the lower two-thirds; internally (Fig. 4c) with bold, nodulose pustules apically, becoming transversely elongate basally and two pustulose longitudinal pilasters (PP). Epiphallus entering penis through a smooth, con- ical verge (PV) that has a terminal pore (EP). Male terminal genitalia enveloped in a thin con- nective sheath. Vagina (V) short with sub-apical entrance of free oviduct; internally with fleshy longitudinal thickenings. Free oviduct (UV) twice as long as vagina, internally with transverse spongy folds. Spermatheca (S) with a short ex- FIG. 2.Animal of Eungarion mcdonaldi sp.nov. Mt Macartney, MEQ, QMM035628. panded stalk (SS) inserted at the base of the free oviduct and subovate head attached about halfway along the uterus. Atrium (Y) short. Spermatophore (Fig. 4d, SP) elongately ovoid with a long chitinous tail. Radula with a tricuspid central tooth (Fig. 3d) that has a broadly lanceolate mesocone and two small, pointed lateral cusps; lateral teeth (Figs 3d,f) strongly bicuspid with endocone reduced to a small point high up on the edge of the mesocone, ectocone short and pointed, anterior edge ir- regularly grooved and ridged; marginals (Fig. 3e) bicuspid with two, equally short, pointed cusps and a long sinuate shaft. Latero-marginal transi- tion abrupt. Basal plates of central and lateral teeth short, squarish with lateral ridges. Interrow support (Fig. 30 provided by interlock of basal plates with anterior edge of tooth behind. Jaw (Fig, 3c) arcuate with low, concentric growth ridges. Radular formula. 140.1 1.1.1 1.140. Based on 4 dissected specimens (QMMO- 35628, QMMO43309, QMMO36087). Range and Habitat Known from only simple notophyll vine forests of the summits and near summits (9(X)- 1 200m) of the Clarke Ra., MEQ (Fig. 5). The absence of the species from the Conway Ra. and Mt Dryander to the north, is probably real. Considerable coHect- ing has been undertaken in these areas. Kmcdonaldi has been collected from inside dis- carded palm fronds and on leaves on the ground. Remarks Relationships of E. mcdonaldi are not clear. A number of other semi-slugs in both northeastern and southeastern Queensland are altitudinally restricted and have degenerate shells. These will need to be dissected in order to determine whether ‘very reduced-shell’ forms are derivatives of local semi-slugs with more complete shells or whether they are in fact closely related species 30 MEMOIRS OF THE QUEENSLAND MUSEUM that have been separated by climatically-induced habitat fragmentation. DISCUSSION The eastern Australian semi-slug fauna (family Helicarionidae) is largely undescribed. Based on the collections of the Queensland Museum, it is likely that the number of species, currently 25 (Smith, 1992), will at least double. Of the named species, only a handful have been studied to a standard acceptable in modern systematics. Generally there has been undue emphasis on gross shell characters, in a group displaying a diverse range of shell reduction, to the detriment of anatomical studies. The distribution of the eastern Australian semi-slugs is mainly focussed around the archipelago of rainforest refugia oc- curring along the eastern edge of the continent. This is not surprising in view of their moisture sensitivity, and suggests a complex evolutionary history since their ancestors reached the continent from the Indo-Malay region in the mid-Miocene (Bishop, 1981). Sympatry involves 2, rarely 3-4 species, and there is considerable microhabitat diversification among the sympaters. Microsym- patry has yet to be observed. There are no iden- tifiable large, local radiations (Stanisic unpubl.) so that ordering of character states and identifying direction of character change probably hinges on revision of the entire semi-slug fauna. Against this background attempts to draw relationships between E. mcdonaldi and other described helicarionid semi-slugs need to be tempered with caution. The use of shell characters in a group most remarkable for degrees of shell reduction requires a focus on fundamental features (Fig. 1) such as sculpture and coiling pattern rather than gross similarity. Hence, although the shell of Thularion semoni (Martens, 1894) from the Wet Tropics, NEQ, also has incomplete early whorls, poor calcification and membranous shell base (Stanisic, 1993) these alone are not sufficient to warrant close association with E. mcdonaldi. The shell of E. mcdonaldi is rounded, and, apart from its flattened spire and degenerate nature, more closely resembles the shell of Helicarion s.s. and Fastosarion sensu Smith (1992). Sculpture (Fig. 3a-b) is obsolete, but this should be viewed in the context of shell reduction and the concomitant loss of key features. Although reduced, the shell of T. semoni retains typical helicarionid spiral sculpture on both protoconch and post-nuclear whorls and has a much more expanded final whorl which gives the shell an elliptical aspect. The animal of E. mcdonaldi (Figs. 2,4a) has retained the basic features (reasonably discrete shell lappets and mantle lobe, prominent caudal horn) seen in temperate species (Dartnall *& Ker- shaw, 1978; Kershaw, 1979, 1981) and some subtropical Fastosarion. In contrast T. semoni has the lappets and lobes more fused and greatly expanded anteriorly to form a cephalic shield. Stanisic (1993) considered this exuberant development of the accessory respiratory sur- faces in T. semoni as an evolutionary experiment intermediate between the Helicarion s.s. and Par- macochlea Smith, 1884. E. mcdonaldi differs from southern species as- signed to Helicarion s.s. such as H. nigra (Quoy andGaimard, 1832),//. rubicundus (DoxinaU and Kershaw, 1978) and //.cnv/cr/Ferussac, 1821, in genital anatomy (Fig. 4b-d), by possessing an epiphallic retractor caecum and lacking an epiphallic gland. The penis of E. mcdonaldi has a verge and simple sculpture of discrete pustules contrasting with the pattern of penial papilla and complex penial sculpture seen in the southern species (Dartnall & Kershaw 1978; Kershaw 1979, 1981). The relatively simple sper- matophore of E. mcdonaldi is also quite distinct from the heavily ornamented (spined) sper- matophores seen in H. nigra, H. rubicundus and H. cuvieri. 1 consider these to be fundamental differences above the level of changes associated with species interactions. In contrast, E. mcdonal- di shares some basic anatomical features with T. semoni (e.g. shape of spermatophore and presence of an epiphallic retractor caecum). T. semoni lacks a penial verge but retains pustular sculpture in the penis chamber (Stanisic, 1993). The highly modified Parmacochlea from far north Queensland possesses pustular penial sculpture, penial verge, epiphallic caecum and has a spermatophore not unlike that of E. mcdonaldi and T. semoni in shape. It differs only in having fine spination on the chitinous section (Simroth, 1898). Hence there are several basic differences which separate the tropical and temperate semi-slugs. At the same time, I do not necessarily consider E. mcdonaldi and T. semoni close relatives. Whereas T. semoni appears to represent a distinct lineage of semi-slug development (Stanisic 1993), E. mcdonaldi can be readily derived from species with less reduced shells by simple but more ex- tensive reduction in shell. The radulae of eastern Australian semi-slugs A NEW MONTANE SEMI-SLUG 31 FIG. 3. Eungarion mcdonaldi sp.nov. Mt Dalrymple, MEQ, QMMO36087. a, protoconch; b, adult shell sculpture; c, jaw; d, central and inner lateral teeth; e, marginal teeth; f, lateral teeth showing interrow support. Scale lines as marked. appear to be fairly generalised (Baker, 1941; Dartnall & Kershaw, 1978; Hedley 1893; Ker- shaw, 1979, 1981; Semper, 1885; Stanisic, 1993) in form and function (interrow support). Minor differences should probably relate to niche spec- ialisation due to sympatry. E. mcdonaldi shows some radular differences from both the temperate species {H. nigra, H. cuvieri, H. rubicundus) and the tropical T. semoni. In H, nigra, H. cuvieri, H. rubicundus and T. semoni the marginal teeth have subequal, somewhat elongate cusps. In E. mcdonaldi the cusps (Fig. 3e) are very short and 32 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Eungarion mcdonaldi sp.nov. a, Dalrymple Heights, MEQ, QMMO43309, holotype; b-d, Mt Macartney MEQ, QMM035628, paratype. a, whole animal; b, reproductive system; c, penis interior; d, spermatophore Scale lines = 2mm. A NEW MONTANE SEMI-SLUG 33 FIG. 5. Distribution of Eungarion mcdonaldi sp.nov. (dots). equal. In addition, the number of marginal teeth in E. mcdonaldi is greatly increased (140) when compared with H. nigra (38), H. cuvieri (30), H. rubicundus (80) and H. semoni (83). While it is possible that these features may be phylogeneti- cally significant it is also probable that they are merely related to feeding specialisation by E. mcdonaldi in response to its sympatry with Fas- tosarion superba {Cox, 1871). The distribution of E. mcdonaldi (Fig. 5) in the summit refugia of the Clarke Ra. suggests that it has been restricted in drier times and has become environmentally-cornered. Hence, its nearest relatives may not survive in MEQ. These refugia are also home to other restricted endemic land snails, e.g. Setomedea janae Stanisic, 1990; Biomphalopa recava (Hedley, 1912) and the slug-like caryodid, Pandofella whitei (Hedley, 1912). With the exception of B. recava, which also occurs at lower altitudes at Finch Hatton Gorge, they are confined to these upland refugia. Relationships of these relict species are both with the south (P. whitei, S. janae), or north (B. recava, S. janae) and not locally, providing evidence of past connections between regions now separated by tracts of dry, snail-poor countryside. It is pos- sible that the nearest relatives of E. mcdonaldi will be similarly located. At a subregional level these restricted taxa are key elements of a largely endemic MEQ snail subfauna (Stanisic, unpubl.). The Charopidae and Caryodidae are Gondwanan groups and occurrences of family relicts in the MEQ montane communities high- light the importance of these refugia to the per- sistence of ancient land snail faunal elements in the region. The discovery of the restricted E. mcdonaldi extends their significance to modem groups. ACKNOWLEDGEMENTS I am grateful to R. Kershaw for supplying per- sonal notes; to Kylie Williams for producing the SEM illustrations; and to the Queensland Museum Photography Section. Thanks are due also to Jennifer Mahoney for typing the manuscript. LITERATURE CITED BAKER, H.B. 1941. Zonitid snails from Pacific Is- lands. Pt in and rv. Genera other than Microcys- tinae. Bulletin of the Bernice P. Bishop Museum 166: 205-370. BAKER, M.J. 1981. The biogeography and evolution of Australian land snails. Pp. 924-954. In Keast, A. (ed.), ‘Ecological biogeography in Australia’. (W. Junk PubI; the Hague, Netherlands). BISHOP, MJ. (1981). The biogeography and evolu- tion of Australian land snails. Pp. 924-954. In Keast, A. (ed.), Ecological biogeography of Australia’. (W. Junk Publ: The Hague, Nether- lands). DARTNALL. A.J. & KERSHAW. R.C. 1978. Description of a new species of Helicarion (Stylommalophora: Helicarionidae) in Tas- mania. Records of the Queensland Victoria Museum Launceston 62: 1-18. HEDLEY, C. 1 893. On Parmacochlea fischeri. Smith. Proceedings of the Linnean Society of New South Wales, Macleay Memorial Volume, 201- 204, pl.27. IREDALE, T. 1933. Systematic notes on Australian 34 MEMOIRS OF THE QUEENSLAND MUSEUM land shells. Records of the Australian Museum 19: 37-59. KERSHAW, R.C. 1979. Redescription of Helicarion cuvieri from southern Tasmania and Helicarion freycineti from New South Wales (Pulmonata: Helicarionidae). Journal of the Malacological Society of Australia 4: 145-156. 1981. Redescription of the genus Helicarion and of Helicarion niger (Quoy and Gaimard, 1832) from Victoria (Pulmonata: Helicarionidae). Jour- nal of the Malacological Society of Australia 5: 17-31. ODHNER, N.H. 1917. Mollusca. XVII. Results of Dr E. Mjoberg’s Swedish scientific expedition to Australia 1910-1913. Kungliga Svenska Ven- tenskapsakademiens Handlingar Bd 52, No 16, 1-115. SEMPER, C.G. 1885. Reisen im archipel der Philip- pinen. Landmollusken. 3(2): 1-327. SIMROTH, H. 1898. Uber die gattungen Par- macochlea^ Parmarion und Microparmarion. Zoologische Jahrbucher (Abteilung Systematik) II: 151-172. SMITH, B.J. 1992. Non marine Mollusca. In Houston, W. W.K. (ed.), Zoological Catalogue of Australia 8: 1-405. (Australian Government Printing Ser- vice: Canberra). SOLEM, A. 1978. Classification of land Mollusca. Pp. 49-97. In Fretter, V. & Peake, J. (eds.), ‘Pul- monates Vol. 2A, Systematics, evolution and ecology’. (Academic Press: London). STANISIC, J. 1990. Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pul- monata) from subtropical rainforests. Memoirs of the Queensland Museum 30(1): 1-241. 1993. The identity of Helicarion semoni Martens, 1894: a large semi-slug from the Wet Tropics, northeastern Queensland (Pulmonata: Helicarionidae) Memoirs of the Queensland Museum 34(1): 1-10. A REVISION OF THE TRIBE COLPURINI FROM AUSTRALIA (HEMIPTERA-HETEROPTERA-COREIDAE) H. BRAILOVSKY Brailovsky, H. 1993 12 24: A revision of the Tribe Colpurini from Australia (Hemiptera- Heteroptera-Coreidae). Memoirs of the Queensland Museum 34(l):35-60. Brisbane. ISSN 0079-8835. Four new genera {Grosshygia, Grosshygioides, Pachycolpuroides and Woodwardhygia); one new subgenus, Hygia {Australocolpura); and eight new species (Grosshygia lobatula, G. monticeps, G, nigra, Grosshygioides numdibularis, H. (A.) sandaracine, P. monteithi, W. bifida, and Sciophyrus australicus are added to the Colpurini (Coreidae) of Australia. These, along with previously known Australian coreids, are described, illustrated and keyed. Acanthotyla fasciata and Sciophyrus sortita, long known from New Guinea, are recorded on northern Cape York Peninsula, the first Australian record. □ fnsecta, Hemiptera, Heterop- tera, Coreidae, Colpurini Australia, taxonomy, rainforest. H. Brailovsky, Instituto de Biologi'a UNAM, Departamento de Zoologia, Apdo. Postal # 70153, Mexico 04510 D.F; 1 September 1993. This contribution summarizes knowledge of the tribe Colpurini (Hemiptera, Heteroptera, Coreidae) for the Australian region, bringing together information from the literature and from previously unstudied collections. A number of species now known to be present in Australia were either undescribed or were previously known only from New Guinea or adjacent islands to Australia’s north. The Australian Colpurini have never been revised. The first species recorded were by Breddin (19(X)) who described one new genus and two new species from the region under the binomial names, Pachycolpura manca and Acantholybas brunneus. The third known species was Acantholybas kirkal- dyi (Bergrolh, \9Q9\ described from Tasmania’. The fourth and last previously known Australian species was Agathyma praecellens Stal (1861) described from Aru Islands and later recorded from New Guinea, D’Entrecasteaux Islands. Bismarck Archipelago, Waigeo, Ternate, Misool and Australia (Dolling, 1987). The Australian members of the tribe Colpurini are usually black or dark coloured. Among the unique aspects of this tribe are the striking struc- tural differences in the hemelylra, as well as the great diversity in the male genital capsule and in the female genital plates, in contrast to other tribes of coreids (Brailovsky, 1990). They are typically species of the rainforest where species occur on the ground associated with piles of freshly dead leaves from recently fallen trees. This revision brings the known Australian fauna to 15 species ip 13 genera. Queensland has the richest fauna with 13 species. There are 5 species in northern New South Wales. The only record from a state other than Queensland and New South Wales is of the problematic Acantholybas kirkaldyi from Tasmania. This locality needs con- firmation. In the following text I include all the species of Colpurini now known from Australia and describe four new genera, one new subgenus and eight new species. The following abbreviations are used to iden- tify institutions where types are deposited or which generously lent material for this paper: ANIC- Australian National Insect Collection, Canberra; BPBM Bernice P. Bishop Museum, Honolulu, Hawaii; BMNH- The Natural Histoiy Museum, London, England; CAS- California Academy of Sciences (Golden Gate Park); DEI- Deutsches Entomologisches Institut, DDR, Ger- many; IBUNAM- Coleccion Entomologica del Instituto de Biologfa, Universidad Nacional Autonoma de Mexico; QMBA- Queensland Museum, Brisbane, Australia; SAMA-’ South Australian Museum. Adelaide, Australia; UQIC- University of Queensland Insect Collection, Bris- bane, Australia; ZIL- Zoological Institut, Leningrad, USSR; ZMUH- Zoological Museum, University of Helsinki.Geopraphical abbrevia- tions are: FNEQ, NEQ, MEQ, SEQ - far north- eastern, northeastern, mideastem, southeastern Queensland; NENSW - northeastern New South Wales. All measurements are given in millimetres. The name of Dr G.B. Monteith is shortened to GBM in the lists of specimens. 36 MEMOIRS OF THE QUEENSLAND MUSEUM KEY TO THE KNOWN AUSTRALIAN COLPUREMI 1. Each side of head immediately in front of eyes with a long, pointed spine (Figs 1 2A,B) Pachycolpura manca Breddin Sides of head in front of eyes unarmed 2 2, Tyius projecting as a single, large, acute spine 3 Tylus globose, truncated or bifid 5 3. Femora unarmed; callar region of pronotum con- spicuously convex; pronotal disc behind middle line with a transverse wrinkle; male genital capsule with a small median projection Acanthotylafasdata (W alker) Femora strongly armed with long, sharp ventral spines; callar region of pronotum weakly convex to flat; pronotal disc without transverse wrinkle; posteroventral edge of male genital capsule without median projection 4 4. Mandibular plate armed with a short projection; posteroventral edge of male genital capsule con- vex, obtusely rounded Agathyrna praecellens Stal Mandibular plate unarmed; posteroventral edge of male genital capsule elongate and bifurcate Woodwardhygia bifida sp. nov. (in part) 5. Antenniferous tubercles armed 6 Antenniferous tubercles unarmed 10 6. Bucculae armed with an obvious spine near the middle third; scutellum longer than wide Acantholybas brunneus Breddin* Bucculae uniformly rounded 7 7. Micropterous; hemelytral membrane reduced to a small flap; ocelli inconspicuous; head with dorsal region convex to globose in lateral view; ab- dominal stemite VO of female with plica and fissura 8 Macropterous to submacropterous; hemelytral membrane well developed; ocelli clearly developed; head with dorsal region practically flat in lateral view; abdominal stemite VII of female without plica or fissura Pachycolpuroides monteithi sp. nov. 8. Head wider than long; apex of scutellum globose; hemelytra reaching anterior third of abdominal segment II Grosshygia nigra sp. nov. Head longer than wide; apex of scutellum subacute; hemelytra reaching median third of abdominal segment HI 9 9. Antenniferous tubercles armed with long lobes (Fig. HA); antennal segment II longer than 1.96mm Grosshygia lobatula sp. nov. Antenniferous tubercles armed w'ith a short lobe (Fig. IIB); antennal segment II shorter than 1.73mm Grosshygia monticeps sp. nov. 10. Abdominal stemite VTI of female without plica or fissura: frontal angles of pronotum rounded, blunt, not produced 11 Abdominal stemite VII of female with plica and fissura; frontal angles of pronotum produced for- wards as conical teeth 1 3 1 1 . Body length shorter than 1 0. lOmm; posteroventral edge of male genital capsule produced into medium and wide lateral projections, enclosing a deep, U-shaped concavity (Fig. 3B) Sciophyrus diminutus Horvath Body length longer than 10.30mm; posteroventral edge of male genital capsule not as above 12 12. Posteroventral edge of male genital capsule with a small Vshaped concavity, with its lateral arms shorter (Fig. 3C) Sciophyrus sortita (Horvath) Posteroventral edge of male genital capsule, with a very large U-shaped concavity, laterally enclosed by two strong divergent arms (Fig. 3A) Sciophyrus australicus sp. nov. 13. Mandibular plate armed with a large prominent tubercle, pronotal disc with a deep longitudinal furrow along midlinc; hemelytra staphylinoid, membrane absent Grosshygioides mandibularis sp. nov. Mandibular plate unarmed; pronotal disc flat, without midline furrow; macropterous; hemelytral membrane well developed 14 14. Femora armed with two rows of ventral spines; tylus apically bifid Woodwardhygia bifida sp. nov. (in part) Femora unarmed; tylus apically globose Hygia (Australocolpura) sandaracine sp. nov. * = Acantholybas kirkaldyi Bergroth not available. Acantholybas brunneus Breddin (Figs 6E,F, 15A) Acanthoclybas brunnea Breddin, 1900: 40-41. Material Examined SEQ: 3 dd, 39 9 : Brisbane, 20.iii.l930, Liv.l965,C. Hembrow and viii.1971, M.B Malipatil. 49 9 : Mt Glorious, 13.ix.l966, GBM. 1 : Tamborine, A.M. Lea. 19: Central Station, Fraser Is., 14-15.X.1978, GBM. NSW: 1 9, Tooloom Plateau via Urbenville, 31.x. 1970, GBM. ANIC, BPBM, QMBA, UQIC, IBUNAM. Diagnosis Largely dull orange yellow; tylus globose; an- tenniferous tubercles armed with a large, sharp, pointed spine; bucculae with a short, blunt mesal projection; femora unarmed or with small den- ticles along their ventral surface; pronotal mar- gins straight; macropterous. Spermatheca: Bulb elongate, duct heavily coiled, chamber slender. Distribution Lowlands and plateaux from NNSW to SEQ, north to Fraser Is. (Fig. 1 5 A). Breddin described A REVISION OF THE TRIBE COLPURINI 37 FIG. 1 . Dorsal view of Grosshygia lobatula sp. nov., 9 . A. brunneus from New South Wales, without definite locality. Acantholybas kirkaldyi Bergroth (Fig. 15A) Acantholybas kirkaldyi B&T^oth, 1909: 185-186. This species was described from Tasmania without definite locality. I have not been able to locate the type specimen which was a male 10mm long. No additional material was available for this study. Bergroth (1909) distinguished A. kirkaldyi from A. brunneus Breddin because the latter species has the rostrum longer, the pronotum shorter, the abdominal sterna with a median fur- row and the posteroventral edge of the male geni- tal capsule conspicuously sinuate. Acanthotyla fasciata (Walker) (Figs IOC, 15A) Walker, 1871: 196-197. Material Examined FNEQ: 1 d, 2 9 9: Iron Ra., Cape York Pen., 5-10.V.1968 and 30.vi-4.vii.l977, GBM. 3 dd: West Claudie R., Iron Ra., 3-10.xii.1985, GBM & D. Cook. 1 d, 1 9: Cape York, Rocky R., 1958, Darlington. 1 d: Gordon Ck, Iron Ra., 16.x. 1974, M.S. Moulds. QMBA, SAMA, UQIC, IBUNAM. Diagnosis Pale yellow, punctures orange to dark brown red and with a median black red longitudinal stripe located on prosternum, mesosternum, metastemum, abdominal stemite III to VII and between callar region of pronotum; each anten- niferous tubercles armed with a short lobe; tylus projecting in front of jugae. upturned to form a median acute horn at apex; mandibular plates armed with a prominent tubercle; bucculae with small spiny anterior projection; callar region of pronotum globose; pronotal disc with posterior lobe wrinkled; macropterous; abdominal sternite VII of female without plica or fissura. Distribution In Australia from only the lowland rainforests of central Cape York Peninsula (Fig. 15A). This species was originally described from Misool (Walker, 1871) and was later recorded from New Guinea (Stal, 1873). It is here recorded from Australia for the first time. Agathyrna praecellens Stal (Figs lOA, 15A) Agathyrna praecellens S\.k\, 1861: 145. Previous Records Northern Queensland: Cape York Peninsula and Iron Range (Dolling, 1987). Material Examined FNEQ: 9 dd, 19 : West Claudie R., Iron Ra., 3-lO.xii. 1985, GBM Ei D. Cook. 3 d d , 1 9 : Gordon’s Mine area. Iron Ra., 12-1 8. ii. 1976, GBM. IBUNAM, QMBA, UQIC. Diagnosis Largely bright orange red; femora strongly den- ticulate; antenniferous tubercles truncate; tylus projecting in front of jugae, upturned to form a small robust horn at apex; mandibular plates armed with a prominent tubercle; bucculae with small spiny anterior projection; pronotal margins straight; pronotal disc without posterior wrinkle; macropterous; abdominal, sternite VII of female without plica or fissura. 38 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Grosshygia spp. nov.: Grosshygia lobatula 6 genital capsule, posterior view (A), lateral view (B); 9 genital plates, posterior view (C), lateral view (D): G. monticeps S genital capsule, posterior view (E), lateral view (F); 9 genital plates, posterior view (G), lateral view (H). A REVISION OF THE TRIBE COLPURINI 39 no. 3. Sciophyms spp. 6 6 genital capsules; S. australicus sp. nov.(A), S. diminutus Horvath (B), S. sortita (Horvath) nov. comb (C); 9 9 genital plates, S. australicus sp. nov; posterior view(D), lateral view (E); S. diminutus Horvath posterior view (F), lateral view (G). 40 MEMOIRS OF THE QUEENSLAND MUSEUM plates, posterior view (E), lateral view (F), S genital capsule in posterior view (G). A REVISION OF THE TRIBE COLPURINI 41 Distribution In Australia from only lowland rainforests of Iron Ra., FNEQ (Fig. 15A).This is the most com- mon and widespread species of the genus. Described from Aru Islands, and later recorded from Australia, New Guinea, D’Entrecasteaux Islands, Bismarck Archipelago, Waigeo, Temate, and Misool (Dolling, 1987). Pachycolpura manca Breddin (Figs 6C,D, 12A,B, 15B) Pachycolpura manca Breddin, 1900: 39-40. Material Examined NSW: 4 c? (3, 4? 9 Sydney, Lea. 3 66 Parramatta, Macarthur’s Bridge, 29.iii-l2.iv. 1980, R. Patterson. 1 9 : Carlingford, lO.X. 1970, F.H. Ulher Baker. SEQ: 1 9 : Ml Glorious, 24-28.ii.1961, J.L. and M. Gressilt. 1 9: BunyaMts, 19.xi.!967, J.andM Sedlacek. 2 6 6, 19 : Bald Mt. area, via Emu Vale, 16-20.ii.l970 3- 4000’,GBM.3 d(3,79 9 : Brisbane, 17.iii.l957, J.H Martin. 3 6 6,2 99: South Pine River, 17.i.l963, GBM. 19 : Whiteside Xing, N Pine River, 12.vii. 1963, GBM. 1 9 : Ravensboume. 15.ix.l971, B. Cantrell. 1 9: Bald Mtn area, via Emu Vale, 27-31.i.I972 3- 4000’, GBM. 1 9 :Indooroopilly,7.ix.I979,G.Gordh. ANIC, BPBM, BMNH, CAS, IBUNAM, QMBA, SAMA, UQIC, ZIL. Diagnosis Largely dull pale ochre yellow, with brown and orange reflections; sides of head in front of eyes with a long and pointed spine; antenniferous tubercles and femora unarmed; tylus globose; macropterous to brachypterous (sensu Slater, 1 975)bucculae with small spiny anterior projection; abdominal stemite VII of female entirely fissurated. Spermatheca: bulb somewhat elongate, duct rela- tively coiled, chamber more or less globose. Parameres, Figs 6C,D. Distribution Widespread, Sydney, NSW to Brisbane and Bunya Mts, SEQ (Fig. 15B) This species is described from Australia (Breddin 1900, Blote 1936, Kumar 1966). Notes Kumar (1966) described the life history stages and recorded its feeding on stinging nettles (Ur- tica dioica L.), tomatoes, pumpkins and Coreop- sis lanceolatus L. Sciophyrus diminutus Horvath (Figs 3B,F,G, 5C-E, 13 A, 15B) Sciophyrus diminutus Worvdih, 1900: 635. Material Examined FNEQ, NQ, SEQ: 7 d (3, 7 9 9 : Cape York, Locker- bie area, 13-27.iv.l973, GBM. 1 9 : Upper Mulgrave R., I-3.xii.l965,GBM.2 c3d,2 9 9:Cape York Pen., Iron Ra., 28.iv-17.v.l968, GBM, 16-23.xi. 1965, GBM. 4 c3(3.29 9: Kowanyama, 7.i.l977, D.L. Han- cock. 1 3: Mt Webb NP, 50km N Cooktown, 11- 14.vii.l976, G.B. and S. Monteith. 7 3 3, 89 9: Hibberd Pt, Weipa, 3-5.ii.I976, GBM. 1 3: Mt Glorious, 7.V.1959, A.C. Robinson. IBUNAM, QMBA, SAMA. Diagnosis Surface dull, dark brown, with orange or pale ochre yellow reflections; size less than 10.10mm; antennal segments I to III pale orange hazel with basal third pale yellow; antennal segment IV yel- low with basal and apical third pale brown; mac- ropterous; posteroventral edge of male genital capsule extending into medium and wide lateral projections, enclosing a deep U-shaped concavity (Fig. 3B); parameres. Figs 5C-E. The genital plates of the female are straight, with a dorso- ventral enlargement and with external face of gonocoxa I clearly convex (Figs 3F,G). Distribution Widespread on both eastern and western Cape York Peninsula, south to the Mulgrave R., NEQ. One specimen is labelled ‘Mt Glorious’ but this locali^ needs confirmation since it is 1500km beyond the range of the other specimens (Fig. 15B). This species was originally described from Thursday Island and later recorded from Kei Is- lands and New Guinea (Horvath, 1919 and Blote 1936). Sciophyrus sortita (Horvath) nov. comb. (Figs 3C, 5J,K, 13B. 15B) Colpura sortita Horvath, 1900: 635-636. Material Examined FNEQ 1 3: w Claudie R., Iron Ra., 3-10.xii.1985 (50m), GBM & D. Cook 2 33: Iron Ra., Cape York Pen., 5-10.V.1968, GBM. Same locality, 4.vii.l977, GBM. 2 3 3: Cape York, Lockerbie area, 13- 27.iv.1973, GBM. IBUNAM, QMBA. Diagnosis This is a large striking species, similar in colour 42 MEMOIRS OF THE QUEENSLAND MUSEUM and habitus to S. australicus sp. nov., but differ- ing in having the posteroventral edge of male genital capsule with a small ‘V’ concavity, with lateral arms conspicuously shorter (Fig. 3C); parameres like Fig. 5J,K. Anterior and middle femora armed; posterior femora unarmed; ant- erior lobe of pronotum mostly smooth; macro- pterous. Female: Unknown. Distribution In Australia, known only from the rainforests of Lockerbie and Iron Ra., FNEQ (Fig. 15B). This species was originally described from New Guinea and was known only from there until now. Sciophyrus australicus sp. nov. (Figs 3A,C,D, 5A, 13C, 15B) Material Examined Holotype 6 : NEQ: Mt Fisher, 7km SW Millaa Millaa, 27-29.iv.l982 (1050-1 100m), GBM, Yeates and Cook, in QMBA (T12708). Paratypes: NEQ: 3 cJd, 1?: Malanda Falls, 9.xii.l989 (750m), GBM, Thompson & Janetzki, 2 d <3 and 12 in QMBA and 1 S in IBUNAM. 1 2: Bakers Blue Mt.. 17km W Mt Molloy, ll.ix.l981 (900m). GBM &D. Cook, in QMBA. 1 2 : Townsville, 19.i.l9M. J. Sedhicek, in CAS. 4 cJd. Broadwater Park, 35km NW Ingham, 16.xii.l986 (60m). GBM, Thompson & Hamlet, 3 in QMBA and 1 in IBUNAM. 32 2 : Mt Fisher, 7km WNW Cape Tribulation (site 2), 23.ix-7.x.l982 (50m) GBM, Yeates & Thompson, in QMBA. 1 c3, 2 2 2: Kirrama Ra. (Yuccabine Ck. area), 9-12.xii.1986 (600m), GBM, Thompson & Hamlet, I c5 & 1 2 in QMBA and 1 2 in IBUNAM. 1 2 : Kirrama SF, via Cardwell, 17-18.viii.l%6, GBM in UQIC I 2; Mission Beach, 128km S Cairns, 19- 20.1.1964, J. Sedlacek, in CAS. I <3: Cooper Ck, 18m N Daintree R., 14.xi.l969. T. Weir in UQIC. 1 <3: Lacey’s Creek, Mission Beach, 21.iv.l970, GBM, in UQIC. 1 (3: Upper Little Mulgrave R., SW Cairns, 4.ix.l%9,J.E.Tobler.inQMBA.3 <3(3,2 2 2:Upper Mulgrave rd, Kearney’s Falls, 10.xii.l988, GBM & Thompson, 2 66 22 2 in QMBA and 1 6 in IBUNAM. 2 6 6: Kirrama Ra. (Barracks area) 11- 12.xii.l986 (600m), GBM, Thompson & Hamlet in QMBA. 7 3 3, 12 2 2 ; Bellenden Ker Range, Cableway Base Stn, 17-31.X.I981 (100m), GBM, 5 33 & 102 2 inQMBA&2 33&22 2 in IBUNAM. 1 3: Baldy Mt. rd, SW Atherton, lO.x.1980 (1200m), GBM in QMBA. I 3 : Palmerston NP, via Innisfail, 7-8.viii.1968, B. Cantrell, in UQIC. 1 3 Mossman Gorge, 7. xii. 1966, B. Cantrell in UQIC. 1 3,12 :Lake Eacham, 24.ix.1970, GBM, female in UQIC. male in IBUNAM. 1 3: The Boulders, via Babinda, 15.xii.l966, B. Cantrell in UQIC. 1 3, 1 2 : Upper Mulgrave R., 30.iv.l970, GBM, in UQIC. 22 2 : Upper Mulgrave R., 1-3. xii. 1965, GBM, in UQIC. 1 3,12: Wallaman Falls, via Ingham, 7.viii.l968, B. Cantrell in UQIC. 1 3 : Mossman Gorge, via Mossman, 28.xii.1967, B. Cantrell in UQIC. 1 3: Henrietta Ck, Palmerston NP, 29 .xii. 1964, GBM in IBUNAM. 1 3. Kauri Ck, Tinaroo Dam, 24.iv. 1970, G.B. Monteith in UQIC. 1 2 ; Malanda Falls, 1 1 .v.1970, GBM in UQIC. I 3, 32 2 : Upper Mulgrave, via Gordonvale, 26- 27.xii.1967, GBM, 1 3 & 22 2 in UQIC & I 2 in IBUNAM. ! 3: Kirrama Ra. (Douglas Ck rd), 9- 12.xii.l986 (800m), GBM. Thompson & Hamlet in QMBA. 1 3: Shipton’s Flat, via Helenvale, 20- 27. vii. 1974, GBM & D. Cook, in QMBA. 2 33: Cape Tribulation, I2-19.X.1980, GBM in QMBA. 1 3: Fly- ing Fish Pt, 21. i. 1965, E.C. Dahms in QMBA. I 3: Millaa Millaa Falls, 4.xii.l965, GBM in UQIC. 1 3, 22 2: Henrietta Ck, Palmerston NP,5.xii.l965, GBM in UQIC 12 : Rifle Ck, Black Mtn, 18km ESEJulat- ten, 13-14.iv.l982 (400m), GBM, Yeates & Cook in QMBA. 3 ; Hinchinbrook Is ., Gayundah Creek, 7- 15. xi. 1984 (10m), GBM, Cook & Thompson, in QMBA. 1 2 :MossmanBluffTrack,5-10kmW. Moss- man, 1-16. i. 1989 (360m), GBM, Thompson & ANZSES in QMBA. 1 2 ; (Gordon’s Mine Area, Iron R., 12-18.ii.l976, GBM, in QMBA . Description Male: Dorsal coloration: Orange brown, head and anterior lobe of pronotum alternating red brown or black spotting; following areas pale yellow: a longitudinal band running from the antenniferous tubercles to the neck, comprising the space between the eye and the ocelli and the superior area of the postocular tubercle, lateral margins of the pronotal collar, anterolateral bor- ders of pronotum, external edge of humeral angles, an irregular spotting at the middle lobe of the pronotal disc and the greater part of the scutel- lar surface and its apex; antennal segments I and II ochre chestnut brown, with a shining orange red apex; segment III chestnut red and IV ochre yel- low, with its basal third red brown and the apical third chestnut orange; hemelytral membrane amber brown with veins darker; connexival seg- ments red brown, with posterior border yellow and most of the surface with orange reflections; abdominal segments orange red with the greatest part of VII orange brown. Ventral coloration: Chestnut orange on an ochre yellow background; following area chestnut orange to chestnut orange brown: middle head region, prostemum, meso- stemum and metastemum as well as the area neighbouring the lobes of the metathoracic scent glands: rostral segments I to III chestnut orange and IV somewhat paler; neighbouring region of the inferior eye area pale yellow, anterior lobe of the metathoracic scent gland yellow, the posterior lobe chestnut orange; coxae red brown; troch- anters yellow with some chestnut brown spots; femora red brown with two yellow rings, one A REVISION OF THE TRIBE COLPURINI 43 basal, the other one near the middle line, tibiae alternating two yellow rings with three chestnut brown to dark brown rings; tarsi chestnut brown with orange reflections on 11 and HI; propleura and metapleura with a black half moon and mesopleura with an anterior irregular black spot; abdominal stemites with three-coloured pleural margin, upper one red brown, posterior one dark yellow and dark orange the rest. Structure: Eyes medium sized, prominent; ocelli with large diameter and situated on a line such that their upper margin is level with the lower margin of eyes; rostrum almost reaching the posterior third of sternal segment V; rostral segment I reaching anterior margin of proster- num. Pronotum: Middle and posterior lobe den- sely punctate, inconspicuously striate; anterior lobe with small diameter punctures and some smooth areas; frontal angles blunt, slightly produced; anterolateral borders clearly emar- ginated and bilobate, with anterior lobe short and convex, and posterior one higher and convex; humeral angles rounded, not exposed, in lateral view feebly convex; posterolateral borders straight and somewhat sinuate; posterior border straight. Legs: Femora armed with a double row of minute denticles. Scutellum: Abruptly punctate and irregularly striate; apex short and blunt. Hemelytra: Macropterous, reaching the apex of the abdomen clavus; and corium with medium punctures except the posterior third of the corium which is smooth. Ventral surface den- sely punctate. Genital capsule: Posteroventral margin with a pronounced U-shaped concavity, enclosed by two very open and conspicuously robust arms (Fig. 3A). Parameres: Fig. 5A,B. Female: Coloration: Similar to male. Structure: Abdominal stemite VII without plica or fissura; abdominal segment IX trapeziform, short, with a narrow posterior border and a small U-shaped concavity, never deep. Genital plates: Gonocoxae I straight, well developed, with an antero-posterior enlargement and with the external face entire, feebly convex; paratergite VIII small; paratergite IX medium sized and not conspicuously raised (Fig. 3D,E). Spermatheca: Bulb elongate, duct heavily coiled. Measurements: 6 first, then 9: Head length: 1.50, 1.57; interocellar space: 0.45, 0.50; inter- ocular space: 0.85, 0.87; width across eyes: 1.65; 1.70; preocular distance: 0.90, 0.97; length anten- nal segments: 1, 1 .65, 1.70; II, 2.30, 2.50; III, 1 .70, 1.80; IV, 1.52, 1.55. Pronotum: Total length: 2.20,2.32; width across frontal angles: 1.35, 1.45; width across humeral angles: 3.35, 3.60. Scutellar length: 1.75, 2.(X); width: 1.60, 1.70. Total body length: 10.60, 11.56. Diagnosis S. aiistralicus is easily distinguished by the pronounced U-shaped concavity at the posteroventral edge of the male genital capsule, which is laterally enclosed by two strong diver- gent arms (Fig. 3A). In 5. anmdipes Blote, the related species, the posteroventral edge has two long and pointed arms and the U-shaped opening is less pronounced. The female strongly resembles that of S. an- nulipes. The determination is based upon the locality labels, assuming that both sexes were collected at the same time; however the opening of the posterior border of the abdominal segment IX in S. australicus, is narrower than in S. an- nul ipes. Distribution Widespread and common in high and low elevation rainforest of the region between Cook- town and Townsville, NEQ (Fig. 15B), One specimen from Iron Ra. needs confirmation. Etymology Relating to the collecting locality. Grosshygia gen. nov. Description Head: Slightly longer than wide across eyes or wider than long, pentagonal to subquadrate, con- spicuously convex dorsally; tylus unarmed, api- cally truncate, extending anteriorly to the jugae and more raised in lateral view; jugae unarmed, thickened and shorter than tylus; antenniferous tubercles armed, lobes raised, diverging anterior- ly and apically acute or rounded or armed with short and robust lobes (Fig. 1 1 A,B); sides of head in front of eyes unarmed and straight; antennal segment I robust, thickest, slightly curved out- wards and shorter than head; segments II and III terete and slender; segment IV fusiform; antennal segments I, III, and IV subcqual, II longest of all; ocelli not elevated, difficult to see and situated on a line where the upper margin does not reach the lower margin of the eyes; prcocellar pit deep; eyes small, globular; postocular tubercles protuberant, globose; bucculae rounded, short, elevated, not projecting beyond antenniferous tubercles, without teeth and with the external edges thick- ened; rostrum long, reaching the posterior margin 44 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Parameres, Sciophyrus australicus sp. nov. {Kfi)^.diminutus Horvath (C’E),Grosshygia lobatula sp. nov. (F,G), G. monticeps sp. nov. (HJ), Sciophyrus sortita (Horvath) nov. comb. (J,K). of abdominal stemite V or anterior margin of VI; rostral segment I reaching the posterior gular region; rostral segment IV longest, rostral II longer than I and III subequal to I; vertex globose, with slight transverse depression, dividing it into two elevations, the anterior one broader, ir- regularly nodulose and higher than posterior one which is slightly rounded. Thorax: Pronotum quadrate, bilobed and not declivent, anterior lobe longer than posterior lobe, both with lateral margins convexly rounded, moderately elevated and slightly reflexed; anterior collar wide; posterolateral border straight and posterior border slightly concave; callar region transversely nodulose and separated along the middle line by a slightly longitudinal depres- sion. Anterior lobe of metathoracic scent gland globose and reniform, posterior lobe sharp, small. Legs: Unarmed; tibiae terete, with sulcus dif- ficult to see. more slender than femora. Scutellum: Triangular, wider than long, with subacute apex; disc convexly rounded. Hemelytra: Micropterous, reaching anterior third of abdominal segment II or median third of abdominal segment III; wings reduced to small pads separated from each other, leaving the ab- domen exposed mesally; clavus and corium fused with raised veins; membrane absent or repre- sented by a small lobe. Abdomen: Connexival segments higher than body, posterior angles complete, not projected into spines; dorsal segments IV-V and VI-V with two mound-like projections to the sides of the middle line; abdominal stemites with medial ster- nal furrow extending to posterior border of ster- nite VI. Integument: Body surface rather dull. Pronotum, scutellum and hemelytra with scat- tered punctation. Antennae and legs minutely granulate. Head, pronotum, scutellum, legs, hemelytra, thorax and abdomen with long or short decumbent to suberect conspicuous golden or silvery bristle-like hairs. Ventral surface with cir- cular greyish-white farinose punctures. Male genital ia: Genital capsule: Posteroventral margin protruding as a feeble lip with edge com- plete (Fig. 2A,B) or with a protruding margin slightly excavated on the mid-line. Parameres: simple and straight; anterior lobe convex and continuous with body; posterior lobe ending in a sharp and slender projection (Fig. 5F,G) or in a thick projection (Fig. 5H,I); space between posterior lobe and body wide and amply indented (Fig. 5F,G) or feebly concave (Fig. 5H,I). Female genitalia: Abdominal stemite VII with plica and fissura evident; plica narrow and transversely evolved; gonocoxae I squarish, large; paratergite VIII short, square, with visible spiracle; paratergite IX squarish, and larger than former paratergite (Fig. 2C,D,G.H). Spermatheca with bulb globose, duct heavily coiled. Diagnosis: Like Lygaeopharus Stal, with rounded bucculae, short, without sharp mesal projection, femora unarmed, tylus unarmed and apically truncated or globose and hemelytra short. The females of both genera have abdominal ster- nitc VII with plica and fissura evident. The two genera can be separated on the basis of the following combination of characters. In Grosshygia the antenniferous tubercles are armed with either large or short robust lobes, the cephalic dorsum is ncxlule-like, scutellum is wider than long and with a convexly rounded disc; gonocoxae 1 larger, squarish, straight and not protruding past the apex of paratergite IX when viewed laterally; abdominal stemite VII with plica transversely narrowed. In Lygaeo- pharus the antenniferous tubercles are unarmed, cephalic dorsum slightly convex, scutellum longer than wide with disc not globose and gono- A REVISION OF THE TRIBE COLPURINI 45 FIG. 6. Parameres, Woodwardkygia bifida sp. nov, (A,B), Pachycolpura manca Breddin (C,D)» Acan- tholybas brunneus Breddin (E,F), Pachycolpuroides monteithi sp. nov (G,H). coxae I short, triangular, outwardly projecting, reaching further than the apex of paratergite IX; abdominal stemite VII triangular and very broad. Etymology Named for Gordon F. Gross in recognition of his many contributions to the study of Australian Heteroptera. Type Species Grosshygia lobatula sp. nov. Grosshygia lobatula sp. nov. (Figsl,2A-D, 5F,G, llA, 16A) Material Examined Holotype NEQ: Zillie Falls, via Millaa Millaa, 12.viii.l968, B. Cantrell, in QMBA (T12709). Paratypes: NEQ: 1 9 : Downey Ck, 25km SE Millaa Millaa, 7.xii.l988, GBM & Thompson (4(X)m) in QMBA. 1 6: Mission Beach, 7.xii.l965, GBM, in UQIC. 1 cj, 1 9 : Upper Boulder Ck, via Tully, 24- 27.X.1983, GBM, Yeates & Thompson (650-900m), in QMBA. 6 c5‘, 69 9 : Palmerston NP, via Innisfail, 23-25.iv.l968 & 7-8.viii.1968. GBM & B. Cantrell. 4 &49 9 in UQIC & 2 & 29 9 in IBUNAM. 3 : Palmerston NP, Henrietta Ck, 29.xii.1964, GBM, in UQIC. 1 c?, 19 : Palmerston NP, 2.i.l990, GBM in IBUNAM & QMBA. 1 c5, 19 ; Palmerston NP, 2.i.l990 (350-400m), GBM in QMBA. 19 : Upper Boulder Ck, 11km N Tully, 5-7.xii.1989 (1000m), GBM, Thompson & Janetzki in QMBA. Description Coloration: Body dark red brown, with follow- ing areas bright yellow ochre; antennal segment IV (except base), a large spot located near pos- terior edge of connexival segment III to VI, rostral segments I to IV and posterior angle of connexival segment III to VI; coxae bright red brown; trochanters bicoloured, with external side hazel brown and internal side bright yellow; ant- erior and middle femora dark hazel brown; pos- terior femora dark hazel brown with three yellow rings, one subbasal, other almost mesal and the third one subdista! including the greater part of the posterior half; tibiae pale hazel brown; tarsi pale hazel brown with yellow reflections. Structure: Antenniferous tubercle armed, with lobes raised, extremely long, diverging anteriorly and with the apex acutely rounded (Fig. 1 ). Geni- tal capsule: Posteroventral edge complete (Fig. 2A,B). Parameres. Fig. 5F,G. Female: Coloration: Similar to male. Genital plate. Fig. 2C,D. Variation: The type material has some colour variation, present on practically each specimen; 1 . Upper side of postocular tubercle yellow ochre or entirely yellow ochre. 2. Rostral segments I and II pale hazel orange. 3. Posterior margin of connexival segments III to VI entirely yellow ochre. 4. Pleural margin of abdominal sternites III to VI with posterior margin yellow ochre. 5. Abdominal sternites dark red brown and con- spicuously spotted with yellow ochre. 6. Anterior and posterior lobes of melalhoracic scent glands dark orange. 7. Anterior and middle femora dark red brown with pale yellow ochre reflections. 8. Posterior femora yellow with two pale brown rings, one subbasal, other almost mesal. 9. Pos- terior tibiae pale hazel orange with two yellow internal reflections. 10. Tarsi yellow. Measurements: 6 first, then 9 : Head length: 1.72, 1.82; inlerocellar space: 0.58, 0.60; inter- ocular space: 1.12, 1.16; width across eyes 1.64, 1.72; preocular distance: 1.32, 1.40; length anten- nal segments; 1, 1.16, 1 .14; II, 2.00, 1 .96; III, 1.16, 1.16; IV, 1.24, 1.20. Pronotum: Total length of anterior lobe: 1.04, 1 .08; total length of posterior lobe: 0.60, 0.60; total width of anterior lobe: 2.20, 2,36; total width of posterior lobe: 2.52, 2.56. Scutellar length: 0.88, 0.92. Width: 0.96, 0.98. Total body length: 9.66, 10. 15. 46 MEMOIRS OF THE QUEENSLAND MUSEUM Distribution In rainforest at high and low elevations, from southern Atherton Tableland to Mission Beach, NEQ (Fig. 16A). Etymology Referring to the appearance of the antennifer- ous tubercles. Grosshygia monticeps sp. nov. (Figs 2E-H,5H,I, IIB, 16A) Material Examined Holotype S : NEQ: Upper Mulgrave R. via Gordon- vale, 25. iv. 1968, GBM & B. Cantrell, in QMBA (T12710). Paratypes: NEQ: 5 d' d , 1 9 : same data as holotype; 4 d d in QMBA and 1 d and 1 9 in IBUNAM. 2 d d: Upper Mulgrave R., l-3.xii.l%5, GBM, 1 in QMBA & 1 in IBUNAM. 1 d: Graham Ra., 9.iv. 1979, GBM, 100-200m, in QMBA. 2 66: Bellenden Ker Ra., I km 5 Cable Tower 6, 17.x-5.xi. 1 98 1, 500m, Earthwatch/QM, pyrethrum knockdown, in QMBA; I d: Upper Mulgrave R., 30.IV. 1970, GBM, in UQIC. Description Coloration: Body dark red brown, with dark orange reflections and with following areas bright or dark yellow ochre: antennal segment IV (ex- cept base), postocular tubercle, posterior edge of connexival segments III to VII, rostral segments II to IV and posterior edge or posterior angle of abdominal sterna III to VI; rostral segment I hazel orange; anterior and middle femora dark red brown with diffused dark yellow ochre spots; posterior femora mostly pale yellow with two or three incomplete dark red brown rings; tibiae and tarsi pale yellow ochre with orange brown reflec- tions. Structure: Antenniferous tubercles with short and robust lobes (Fig. HB). Genital capsule: Posteroventral edge protruding, indented towards the middle line and laterally excavated (Fig. 2E,F). Parameres. Fig. 5H,I. Female: Coloration: Similar to male. Genital plate. Fig. 2G,H. Variation: The type material has some colour variation, present on practically each specimen: 1. Ail femora pale yellow ochre including the apex and with two or three rings dark red brown. 2. All tibiae dark red brown, with two yellow rings, one subbasal and the other near middle. 3. Posterior edge of abdominal sterna VII dark yel- low ochre. Measurements: 6 first, then 9: Head length: 1.80, 1.68; interocellar space: 0.54, 0.54; inter- ocular space: 1.14, 1.12; width across eyes: 1.68, 1.62; preocular distance: 1.28, 1.22; length anten- nal segments: I, 1.14,0.92; II, 1.72, 1.60,111, 1.04, 0,96; IV, 1.12, 1.08. Pronotum: Total length of: anterior lobe; 0.96, 1 .00; total length of posterior lobe: 0.64, 0.52; total width of anterior lobe: 2.24, 2.02; total width of posterior lobe: 2.72, 2.46. Scutellar length: 0,92, 0.88; width: 1.08, 0.94. Total body length: 10.00, 8.70. Diagnosis This species is similar in colour and general habitus to G. lobatida. In G. monticeps the anten- nal segment II is shorter (1.60 -1.72) and more robust, the antenniferous tubercle is armed with short and robust lobes (Fig. 1 IB) and the shape of the paramenes (Fig. 2A,B,E,F) is quite distinc- tive. In G. lobatula antennal segment II is larger (1.96-2.00) and more slender and the anten- niferous tubercle is armed with extremely long lobes (Fig. 1 lA). Distribution This species occurs at lowland rainforest sites in a restricted high rainfall area around the Bel- lenden Ker Ra., NEQ (Fig. 16A). Etymology Referring to the appearance of the head viewed laterally. Grosshygia nigra sp. nov. (Figs 13D, 14, 16B) Material Examined Holotype 6 : NEQ: Stewart Ck, 4km NNE Mt Spur- geon (Camp 1), 16°24’S 145°13'E, I5-20.X.I991 { 1 250- 1 300m), GBM, Janelzki, Cook & Roberts, in QMBA (T12711). Paratypes: NEQ: 1 6 : same data as holotype, in IBUNAM. 1 6 : 7km N Mt Spurgeon (Camp 2), 16°22’S I45°13’E, i749.x.I991 (1200-1250m), GBM, Janetzki, Cook & Roberts, in QMBA. Description Coloration: Body dark red brown, with follow- ing areas bright yellow ochre: antennal segment IV (except base), posterior edge of connexival segment III to VI and few spots scattered on the pronotal disc and on the abdominal sternites III to VI; coxae bright red brown; trochanters bright yellow with hazel brown reflections; femora pale yellow with four or five incomplete or complete hazel brown rings; tibiae and tarsi pale hazel brown. Structure; Head wider than long; anten- niferous tubercle armed, with lobes, raised, ex- i A REVISION OF THE TRIBE COLPURINI 47 tremely long, diverging anteriorly and with the apex acutely rounded; apex of scutellum globose. Hemelytra: Micropterous, reaching anterior third of abdominal segment II; membrane absent. Genital capsule: Posteroventral margin protrud- ing as a feeble lip with edge complete. Female: Unknown. Variation: The type material has some colour variation. 1. Tibiae pale hazel brown with two yellow rings, one subbasal and the other almost mesial. 2. Tarsi pale yellow. Measurements: S: Head length: 1.45; inter- ocellar space 0.67; interocular space: 1.10; width across eyes: 1.55; preocular distance: 0.93; length antennal segments: 1,0.71; II, 1.14; III. 0.80; IV, 0.93. Pronotum: Total length of anterior lobe: 0.80; total length of posterior lobe: 0.43; total width of anterior lobe: 2.26; total width of post- erior lobe: 2.63. Scutellar length: 0.65; width: 0.67. Total body length: 8.18. Diagnosis Like G. lobatula, has the antenniferous tubercle armed with extremely long lobes, diverging ant- eriorly and with the apex acutely rounded and the male genital capsule has the posteroventral mar- gin protruding as a feeble lip with edge complete. G. nigra can be recognized by its head wider than long, apex of scutellum globose and hemelytra extremely micropterous just reaching anterior third of abdominal segment II. In G. lobatula the head is longer than wide, apex of scutellum sub- acute and hemelytra longer, reaching median third of abdominal segment III. Distribution From only higher, western parts of the Carbine Tableland, NW Mossman, NEQ (Fig. 16B). Etymology From the latin nigra, black. Grosshygioides gen. nov. Description Head: Longer than wide (across eyes), pen- tagonal, non declivent and dorsally flat; tylus unarmed, apically globose, extending anterior to jugae and more raised when viewed laterally; jugae unarmed, thickened and shorter than tylus, mandibular plates, directly below apices of jugae with a large prominent tubercle; antenniferous tubercles unarmed; sides of head in front of eyes unarmed, straight and longer than total length of eye; antennal segment I shortest, robust, thickest, slightly curved outwards and shorter than length of head; segments II and HI cylindrical, slender, IV fusiform; antennal segment II the longest and segment IV longer than III; ocelli absent; preocel- lar pit obliquely deep; eyes small, globular; pos- tocular tubercle protuberant, globose; bucculae rounded, short, elevated, not projecting beyond antenniferous tubercle, angulate, without teeth and with the external edges thickened; rostrum long, reaching anterior margin of abdominal ster- nite VI; rostral segment I shortest, reaching the posterior gular region: rostral segment IV longest and rostral segments II and III subequal. Thorax; Pronotum wider than long, trapeziform, non declivent; anterior collar wide; frontal angles projecting forward as rounded teeth; anterolateral borders weak and obliquely convex; humeral angles rounded, not exposed; posterolateral borders straight; posterior border concave; callar region convex, separated along the midline by a deep longitudinal furrow, which extends to the posterior margin. Anterior lobe of metathoracic scent gland globose and reniform, posterior lobe sharp, small. Legs: Unarmed; tibiae cylindrical, with sulcus difficult to see and more slender than femora. Scutellum: Triangular, flat, wider than long, with apex subacute. Hemelytra: Staphylinoid, reaching posterior third of abdominal segment III; clavus and corium fused into a coriaceous pad and the wings meeting each other along the midline; hemelytral membrane absent. Abdomen: Connexival segments practically at the same level as abdominal segments; posterior angle of connexiva not extending into short spines; abdominal sternites with medial furrow extending to posterior border of sternite VI. Integument: Body surface rather dull. Pronotum, scutellum and hemelytra scattered punctate. Head, antennae, pronotum, scutellum, legs, hemelytra, thorax and abdomen with short decumbent to suberect golden or silvery bristle- like hairs, intermixed with long erect bristles in antennal segments and legs. Ventral surface with circular grey-white farinose punctures. Female genitalia: Abdominal sternite VII with plica and fissura; plica narrow, elevated and transversely evolved, gonocoxae I squarish, large; paratergite VIII short, square, with spiracle visible; paratergite IX larger than paratergite VIII. Diagnosis The reduction of wings, the prominent pos- 48 MEMOIRS OF THE QUEENSLAND MUSEUM tocular tubercle, the pronotal disk with a deep midline furrow and abdominal stemite VII of the female with plica and fissura might suggest a relationship with Grosshygia Brailovsky. In Grosshygioides the antenniferous tubercles are unarmed; the dorsal head is flat and the man- dibular plates are armed with a large and prom- inent tubercle. In Grosshygia the antenniferous tubercles are armed, the dorsal head is con- spicuously globose and the mandibular plates are unarmed. Etymology Named for its external similarity to Grosshygia. Type Species Grosshygioides mandibularis, sp. nov. Grosshygioides mandibularis sp. nov. (Figs 7, lOB, 16A) Material Examined HolotypE: 9 NEQ: Mt Finnigan, 37 km S Cooktown, 19-22.iv.l982, GBM, Yeates & Cook (850-1 100m) in QMBA(T12712). Paratype: 1 9 : NEQ: 2.5 km SW Mt Hartley, 35 km S Cooktown, 23-24.iv.l982, GBM, Yeates & Cook, in IBUNAM. Description Coloration: Body dark red brown, with bright or dull orange reflections and with following areas red orange yellow: posterior third of con- nexival segments III to VII and pleural margins of abdominal sternites III to VII; antennal seg- ments I to III dark red brown and IV light yellow with base dark red brown; dorsal view of pos- tocular tubercle, spot behind eyes and apex of scutellum yellow ochre; rostral segments I-IV pale hazel brown, coxae bright red brown; troch- anters bicoloured, with external side brown hazel and internal side bright yellow; femora dark red brown with following areas yellow: one subbasal ring and few scattered ventral spots; tibiae pale orange brown with two or three yellow rings: one subbasal, other almost mesal and the third one subdistal; tarsi pale hazel brown with anterior or middle third yellow. Measurements: 9 holotype: Head length: 1.73; intcrocular space: 1.02; width across eyes: 1.58; preocular distance: 1.33; length antennal segments: I, 1.20; II, 1.82; III, 1.14; IV, 1.30. Pronotum: Total length; 2.04; width across fron- tal angles: 1.76; width across humeral angles: 3.00. Scutellar length: 1.05; width: 1.14. Total body length: 12.03. Distribution Rare, known from two adjacent high altitude localities S of Cooktown, NEQ (Fig. 16A). Etymology Named for the appearance of the mandibular plate. Hygia (Australocolpura) subgen. nov. Description Head: Wider than long (across eyes), pen- tagonal, not elongate, dorsally flat, with frons not higher than eyes; tylus unarmed, apically globose, extending anterior to jugae and a little higher than them in lateral view; jugae unarmed, slightly thicker and shorter than tylus; genae un- armed; antenniferous tubercles unarmed with truncated apex; antennal segment I the shortest, robust, thickest, slightly curved outwards and shorter than total length of head; segment II and III cylindrical and IV fusiform; antennal segment II the longest, segment IV longer than III; ocelli conspicuous, raised and situated on an hypotheti- cal line with superior margin just touching the inferior margin of eyes; preocellar pit deep; eyes globose; postocular tubercles prominent; buc- culae angulate, short, not extending beyond an- tenniferous tubercles, with small spiny anterior projection; rostrum reaching the middle third or posterior edge of abdominal sternite III; rostral segment I reaching anterior margin of proster- num. Thorax: Pronotum wider than long, trapez- iform, with disc transversely cleft behind the cal- lar region, leaving anterior and posterior thirds raised; anterior collar wide; frontal angles produced forward as conical teeth; anterolateral border weakly sinuate; humeral angles rounded, not exposed; posterolateral and posterior border straight; callar region transversely convex, separated along the middle line by a slight depres- sion; posterior margin with a transverse ridge, distinctly raised. Anterior lobe of metathoracic scent gland globose and reniform, posterior lobe sharp, small. Legs: Femora unarmed; tibiae terete, sulcate and more slender than femora. Scutellum: Triangular, flat, wider than long, with apex acute. Hemelytra: Macropterous, reaching middle third of posterior margin of the last abdominal segment; claval suture evident; apical margin obliquely straight, with a short apical angle not reaching the middle third of the hemelytral FIG. 7. Dorsal view of Grosshygioides mandibularis sp. nov., 9. membrane; costal margin emarginate; hemelytral membrane with few bifurcate veins. Abdomen: Connexival segments higher than body; posterior angle of connexival segments not produced into spines. Integument: Body surface dull, covered by adpressed small hairs, not distinctly hairy. Pronotum (except callar region), scutellum, clavus, corium, thorax, abdominal stemites and exposed parts of genital segments of both sexes strongly punctate; head, antennae and legs minutely granulate. Male genitalia: Genital capsule: Posteroventral border simple, truncate, with lateral borders weakly elongate (Fig. 4G). Female genitalia: Abdominal stemite VII with plica and fissura; plica transversely broad, but never reaching the middle of the segment; gono- coxae I squarish, medium size, with a convex protruding external margin; paratergite VIII short, square, with visible spiracle; paratergite IX FIG. 8. Dorsal view of Pachycolpuroides monteithi sp. nov.,d. squarish and larger than former paratergite (Fig. 4E,F). Type Species Hygia (Australocolpura) sandaracine sp. nov. Diagnosis Hygia Uhler, contains nine subgenera, Caracolpura Breddin, Colpura Bergroth, Eucol- pura Breddin, Hygia Uhler, Microcolpura Bred- din, Pterocolpura Blote, Sphinctocolpura Breddin, Stenocolpura Breddin and Trichocol- pura Breddin, and approximately .72 species, widely distributed in the Oriental Region throughout Japan, China, Taiwan, India, Burma, Assam, Korea, Cambodia, Malacca, Sarawak (Borneo), West Malaysia, Sumatra, Singapore, Java, Philippines, Sulawesi (Celebes), Mentawei and New Guinea. Hygia (Australocolpura) subgen. nov. is re- lated to Hygia { Caracolpura), the only species of which H. (C). planiceps (Breddin) occurs in A REVISION OF THE TRIBE COLPURINI 50 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 9. Dorsal view of Woodwardhygia bifida sp. nov., 9 . Sulawesi, sharing with it the following charac- ters: 1. Body covered by adpressed small hairs, not distinctly hairy. 2. Genae completely un- armed, 3. Head not elongated, wide, above com- pletely flat and the disc of frons not higher than eyes. 4. Femora unarmed, 5. Antenniferous tub- ercle unarmed. 6. Tylus globose or truncated. 7. Plica of abdominal stemite VII of the female transversely broad, but never surpassing the mid- dle of that segment. Members of this new subgenus are distin- guished by the following combination of charac- ters: anterior angle of the bucculae projecting as a small spine, corium without black spot, antennal segment III shorter than IV, transverse ridge of posterior border of pronotum quite raised; general body coloration is pale orange yellow. In Hygia (Caracolpura) the middle third of the bucculae projects as a large and acute spine, the apex of corium with a dark black spot, antennal segment III larger than IV, posterior border of pronotum not raised and general body coloration dark brown or black, with or without pale areas. Etymology Named for its occurrence in Australia. Hygia (Australocolpura) sandaracine sp. nov. (Figs 4E,G, IID, 16A) Material Examined Holotype S : NSW: New England NP, 1 5.i. 1978, E.I. Schlinger, in QMBA (T12713). Paratype: 1 9 : same as holotype, in CAS. Description Coloration: Pale orange yellow with following areas dark brown or black: apex of rostral seg- ment IV, a large spot located below the spiracle of abdominal stemites III to VI, the genital cap- sule and a few scattered spots into the abdominal stemites III to VII; antennal segments I to III dark orange yellow and IV dark orange yellow with a wide subapical yellow ring; hemelytra membrane amber with veins pale yellow and following areas pale brown: basal margin and few cells into the disc; connexival segments orange brown, with posterior edge yellow; abdominal segments yel- low with bright orange red reflections; anterior and posterior lobe of the metathoracic scent glands creamy yellow, punctures hazel orange. Female: Same colour as male. Transverse Carina of the posterior margin of pronotum, anterior third of claval suture and femora with bright red orange reflections; abdominal seg- ments VIII-IX and abdominal stemites VIII-IX dark orange brown with connexival and pleural margins yellow ochre. Measurements: cJ first, then 9: Head length: 1.33 1.48; interocellar space: 0.41, 0.48; inter- ocular space: 0.89 1.01; width across eyes: 1.60, 1.76; preocular distance: 0.81, 0.89; length anten- nal segments: 1, 1 .22, 1 .20; II, 1 .72, 1 .84, III, 1 .20, 1.32; IV, 1.36, 1.41. Pronotum: Total length: 1 .62, 1 .96; width across frontal angles: 1 .44, 1 .60; width across humeral angles: 2.72, 3.08. Scutellar length: 1.14, 1.20; width: 1.32, 1.32. Total body length: 9.30, 10.65. Distribution Known from a single pair from the high New England Tableland, NENSW (Fig. 64). The two specimens were collected in Nothofagus forest. Etymology From the greek sandaracinos, orange coloured. A REVISION OF THE TRIBE COLPURINI 51 FIG. 10. Dorsal views, Agathyma praecellens Stal 6, (A\Grosshygioides mandibularis sp. nov., 9 (B), Acanthotylafasciataiy^sAV^v) 9(C). Pachycolpuroides gen. nov. Description Head: Wider than long, pentagonal to sub- quadrate, scattered punctate and dorsally slightly convex; tylus unarmed, apically globose, extend- ing anteriorly to the jugae and when viewed laterally practically level with them; jugae un- armed, thick and shorter than tylus; antenniferous tubercles armed with a long robust spine; sides of head in front of eyes unarmed; antennal segment I the shortest and shorter than head; segment II longest of all; segment IV longer than III; anten- nal segment I is the most robust, terete and some- f, what curved outwards; segment II and III terete i and slender; segment IV fusiform; ocelli not raised and based on an hypothetical line, the ( superior margin hardly in contact with eyes; J preocellar pit deep and diagonally excavated; ?! eyes elongate; postocular tubercle prominent; J bucculae rounded, short, not projecting beyond 52 MEMOIRS OF THE QUEENSLAND MUSEUM antenniferous tubercle without teeth and with the external edges thickened; rostrum reaching the anterior margin of metastemum; rostral segment I reaching anterior margin of prostemum; rostral segment II longest; rostral segment I longer than rv and subequal to II and IV longer than III which is the shortest of all. Thorax: Pronotum trapeziform, declivent, wider than long; collar distinct; frontal angles produced forward as conical teeth; anterolateral border obliquely straight; humeral angles rounded, not exposed, feebly convex in lateral view; posterolateral border and posterior border straight; callar region transversely convex, separated along the middle line by a slight depres- sion. Anterior lobe of the metathoracic scent gland elevated, globose and reniform, posterior lobe sharp and small. Legs: Unarmed; tibiae terete, sulcate and more slender than femora. Scutellum: Triangular, flat, wider than long with apex acute. Hemelytra: Macropterous condition (in both sexes) reaching posterior margin of abdominal segment VII; claval suture evident; apical margin obliquely straight, with a short apical angle, not reaching the middle third of the hemelytral membrane; costal margin slightly emarginate; hemelytral membrane composed of hexagonal or quadrate or elongate cells. Submacroptcrous con- dition (in both sexes): Reaching anterior or posterior margin of abdominal segment VI; clavus and corium clearly differentiated; membrane well developed, without cells and with only a few bifurcate veins (Fig. 120,0). Abdomen: Connexival segments higher than body, with posterior angles not produced into spines. Male genitalia: Genital capsule: Posteroventral edge complete, thickened and with a slight median depression. Parameres. Body robust, with anterior lobe slightly convex and posterior lobe long and thick (Fig. 6G,H). Female: Abdomen: Sternite VII complete, without plica or fissura. Genital plates: Gonocoxae I relatively short, oblique, with a con- cave and protruding external margin; paralergite VIII conspicuously short, square, with visible spiracle; paratergite IX elongated, with rounded margin that encompasses sternite X. Diagnosis Externally this new genus resembles the general shape and colour pattern of Pachycolpura Breddin, which is the only genus within the tribe Colpurini that exhibits a long, pointed spine on the side of the head in front of the eyes. In Pachycolpuroides, the frontal angle of the pronotum is produced forward as a conical tooth, the bucculae is uniformly rounded, the tylus is level with the jugum, the antenniferous tubercle is armed with a long spine and the abdominal sternite VII of the female complete without plica or fissura. In Pachycolpura the frontal angle is blunt and not protruding, the anterior angle of the bucculae is sharply protected, the tylus more raised than jugum, the antenniferous tubercle unarmed and truncated and sternite VII is totally cleaved (fissured) dividing the sternite into two parts with plica very short, sometimes difficult to see. Like Acaatholyhas StM this genus has the anten- niferous tubercle armed, the femora unarmed and the abdominal sternite VII of the female complete without plica or fissura. Pachycolpuroides may be lecognizkl by its short length, robust body, bucculae rounded without teeth or spiny projection, rostrum not extending beyond the anterior margin of melaster- num, scutellum wider than long and the hemelytra with macropterous or submacroptcrous condition. In Acantholybas the body is large and a little more slender, the bucculae anned with a clear spine near the middle third, rostmm reaching the posterior bor- der of abdominal sternite IV, scutellum longer than wide and the hemelytra always macropterous. Etymology Named for its external similarity to Pachycol- pura. Type Species Pachycolpuroides monteithi sp. nov. Pachycolpuroides monteithi sp. nov. (Figs 6G,H, 8, 12C,D, 16B) Material Examined Holotype 6: Brisbane, 2. i. 1963, GBM in QMBA (T12714). Paratypes: SEQ: 3 S 6.59 9 : Brisbane, vii.1956, J.C. Donahue, I2.V.I959, B. Wilson, 9.iii.l956, H.J. Lavery, 5.iv.l964, A.E. May, iii.1955, N.J. Thompson, 30.vi.I966, B.K. Cantrell, 21.x. 1956, T.A. Bull, 366 &49 9\n UQIC & 1 9 in IBUNAM. 1 6 : Bunya Mts, 14-15.X.1972, B.K. & J.A. Cantrell in UQIC. 19 : Millaa Millaa, 9.i.I964, GBM in UQIC. 1 6: Laminglon NP, 17-24.V.1965, GBM in UQIC. I d, 29 9 : Mt Glorious, 5- 1 5.vii. 1 964. T. Weir. 13.ix.l%6, GBM, 1 (3 & 19 in UQIC & I 9 in IBUNAM. 3 6 6. 29 9 : NSW; Ebor, Il.iv.l966, GBM, B. Cantrell & T. Weir, 266 & \ 9 in UQIC & \ 6 & ] 9 in IBUNAM. in A REVISION OF THE TRIBE COLPURINI 53 FTG. 1 1. Dorsal views, Grosshygia lobatula sp. nov., 9 (A), G.monticeps sp. nov., 6 (B), Woodwardhygia bifida sp. nov., 6 (C), Hygia (Australocolpura) sandaracine sp. nov., S (D). 54 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 12. Dorsal view Pachycolpura manca Breddin; macropterous, $ (A), brachypterous, 9(B): Pachycol- puroides monteithi sp. nov.; macropterous (5(C), submacropterous <5(D). A REVISION OF THE TRIBE COLPURINI 55 FIG. 13. Dorsal view Sciophyrus spp.; S. diminutus Horvath c5^ (A), 5. sortita (Horvath) 6 (B), 5. australicus sp. nov. c?(C): Grosshygia nigra sp. nov. 6 (D). 56 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 14. Dorsal view of Grosshygia nigra sp. nov., S . Description Dorsal coloration: Pale yellow with following areas pale red orange: two wide bands running laterally along cephalic middle line, antennal seg- ments II and III and the intercallar space; antennal segment IV bright orange yellow, with base bright red orange; postocular tubercle and apex of scutellum creamy white; hemelytral membrane amber brown with pale ochre veins; connexival segments brown red orange with posterior edge pale yellow; dorsal abdominal segments bright pale orange brown with yellow reflections; punc- tures of pronotum, scutellum, clavus and corium hazel orange or pale brown. Ventral coloration: Pale yellow with orange reflections; punctures hazel orange or pale brown or dark red brown; following areas black: discoidal spot in the anterior angle of mesothorax, two lateral spots on mesostemum and a line of discoidal spots running near the pleural margin of abdominal stemites III to VII; abdominal stemites III to VII with several red brown spots, irregularly scattered; coxae bright pale orange; trochanters bright yellow with red orange reflections; anterior and middle femora pale orange; posterior femora yellow with apical third and few spots pale orange; tibiae yellow with two pale orange rings, one subbasal and the other one distal; tarsi yellow with orange reflections; rostral segments I and II pale yellow and III-IV hazel yellow; anterior and posterior lobe of the metathoracic scent gland creamy white. Female: Coloration: Similar to male. Ab- dominal segments VIII and IX and genital plates pale yellow with orange reflections. Variation; The type material exhibits some colour variation on most specimens; 1. Antennal segment I bright orange red or pale yellow ochre. 2. Antennal segment IV bright orange red with apex bright orange yellow. 3. Dorsal pale orange red coloration replaced by pale brown hues. 4. Hemelytral membrane pale yellow or pale amber yellow with cells pale brown. 5. Dorsal ab- dominal segments dark red brown with middle portion bright orange with yellow reflections. 6. Prothorax, mesothorax and metathorax with black irregular spots. 7. Ventral granules of femora dark red brown. 8. Anterior edge of prostemum bright red brown. 9. Coxae bright red brown. Measurements: S macropter, 6 submacropter, 9macropter and $ submacropter: Head length: 0.99, 0.96, 1.09, 0.99; interocellar space: 0.49, 0.43, 0.48, 0.46, interocular space: 0.91, 0.82, 1.01, 0.86; width across eyes: 1.45, 1.30, 1.53, 1.39; preocular distance: 0.66, 0.65, 0.71, 0.63; length antennal segments: 1, 0.68, 0.58, 0.74, 0.65; II, 1 .06, 0.83, 1 .09, 0.86; III, 0.89, 0.7 1 , 0.96, 0.7 1 ; IV, 0.89, 0.80, 0.93, 0.83. Pronotum: Total length: 1 .70, 1 .24, 1.86, 1.33; width across frontal angles: 1.39, 1.33, 1 .42, 1 .36; width across humeral angles: 2.72, 2. 1 7, 3.03, 2.26. Sculellar length 1.30, 0.89, 1.42, 0.96; width: 1.45, 0.99, 1.64, 1.1 1. Total body length: 7.18, 6.18, 8.44,6.70. Distribution From the New England Tableland, NSW to Brisbane and the Bunya Mts, SEQ (Fig. 16B). Etymology Named for Dr G.B. Monteith, Australian A REVISION OF THE TRIBE COLPURINI 57 hemipterist of the Queensland Museum, and friend for many years. Woodwardhygia gen. nov. Description Head: Wider than long, pentagonal and dorsal- ly slightly convex; tylus unarmed, apically bifid, FIG. 15. Distributions, Acantholyhas brunneus Bred- din Acantholybas kirkaldyi Bergroth, Acanthotyla fasciata (Walker), Agathyma praecellens Stal. (A); Pachycolpura manca Breddin, Sciophyrus australicus sp. nov. Sciophyrus diminutus Horvath; Sciophyrus sortita (Horvath) (B). extending anteriorly to the jugae and more raised when viewed laterally; jugae unarmed, slightly thicker and shorter than tylus; antenniferous tubercles unarmed, with apex truncate; sides of head in front of eyes unarmed, straight and shorter than total length of eye; antennal segment I the shortest, robust, thickest, slightly curved out- wards and equal or a little shorter than head; segments II and III terete and slender; segment IV fusiform; antennal segment II the longest, seg- ment III longer or shorter than IV; ocelli con- spicuously evident, raised and based on a hypothetical line with the superior margin just 58 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 16. Distributions, Hygia (Australocolpura) sandaracine sp. nov., Grosshygia lobatula sp. nov., G. monticeps sp. nov., Grosshygioides mandibularis sp. nov., (A): Pachycolpuroides monteithi sp. nov., Wood- wardhygia bifida sp.nov., Grosshygia nigra sp. nov. (B). touching the inferior border of the eyes; preocel- lar pit deep and circular; eyes substylate; pos- tocular tubercles protuberant; bucculae rounded, short, not extending beyond antenniferous tubercle, with sharp spiny anterior projection; rostrum long, reaching the medial third of ab- dominal stemite IV; rostral segment II longest, I longer than IV and IV longer than III, which is the shortest; rostral segment I reaching anterior margin of prosternum. Thorax: Pronotum wider than long, trapeziform, moderately declivent; anterior collar wide; frontal angles produced forward as conical teeth; anterolateral border obliquely straight; humeral angles rounded not exposed; posterolateral and posterior borders straight; cal- lar region transversely convex, separated along the middle line by a slight longitudinal depres- sion. Anterior lobe of metathoracic scent gland globose and reniform, posterior lobe sharp, small. Legs: Femora with two rows of small spines and granules along ventral surface; tibiae terete, sulcate and more slender than femora. Scutellum: Triangular, flat, wider than long, with apex subacute. Hemelytra: Macropterous, reaching posterior margin of the last abdominal segment or extend- ing beyond the apex; claval suture evident; apical margin obliquely straight with a short apical angle not reaching the middle third of the hemelytral membrane; costal margin emarginate; hemelytral membrane with few bifurcate veins. A REVISION OF THE TRIBE COLPURINI 59 Abdomen; Connexivai segments higher than body, with posterior angles not produced into spines: abdominal stemites with medial furrow extending to posterior border of stemite IV or V. Integument: Body surface rather dull. Head, pronotum, scutellum, clavus, corium, thorax, ab- dominal sterna and exposed parts of genital seg- ments of both sexes strongly punctate; antennae and legs minutely granulate. Head, pronotum, scutellum clavus, corium, thorax and abdominal sterna with short decumbent silvery bristle-like hairs, intermixed with a few long erect hairs lo- cated on the abdominal sterna. Male genitalia: Genital capsule; Posteroventral edge elongate and bifurcated, with a short depres- sion between the two lobes (Fig. 4A,B). Parameres: body simple and straight; apical projection short with anterior lobe convex and continuous with the body and posterior lobe short and slender (Fig. 6A,B). Female genitalia: Abdominal stemite VII with plica and fissura; plica narrow, elevated and transversely evolved; gonocoxae I relatively short, with a convex protruding external margin; paratergite VIII short square, with visible spiracle; paratergite IX squarish and larger than former paratergite (Fig. 4C,D). Diagnosis Woodwardhygia gen. nov., most closely resembles Homalocolpura Breddin, including the femora armed with two rows of ventral spines, antenniferous tubercle and tylus unarmed and the abdominal stemite VII of the female with plica and fissura. In Woodwardhygia the anterior angle of the bucculae is sharply projected, the tylus apically bifid, antennal segment II longer than IV, rostrum shorter, reaching the medial third of abdominal stemite VI, the femora and tibiae are longer and more slender and the plica narrow and transver- sely evolved. In Homalocolpura the bucculae are uniformly rounded, the tylus is apically rounded or truncated, antennal segment II is shorter than IV, rostrum longer, reaching or extending beyond the apex of the last abdominal stemite and the plica is wider and triangular. An additional character to separate the two genera is the coloration. In Homalocolpura the total body surface is bright and highly polished with femora unspotted, vs a dull body surface, with femora spotted with dark red brown dis- coidal spots in Woodwardhygia. Etymology Named for the late Dr T.E. Woodward, distin- guished Australian hemipterist. Type Species Woodwardhygia bifida sp. nov. Woodwardhygia bifida sp. nov. (Figs 4A-D, 6A,B, 9, 1 1C, 16B) Material Examined Holotype 6 : NSW: Tooloom Plaieau, via Urben- ville, 31.X.I970, GBM in QMBA (T127I5). Paratypes: I <5,49 9 : same as holotype; 1 d & 2 9 9 in UQIC & 2 9 9 in IBUNAM. SEQ: 3 d d , 2 9 9 : Bunya Mts, 14.xii.l937, F.A. Perkins, 5.vi.l959, LC. Yeo & F. McDonald and 18-I9.X.I972 B. Cantrell 2 dc5 & 29 9 in UQIC & I d in IBUNAM. I d: Lamington NP, I9-22.V.I963, H.A. Rose in UQIC. 1 d: Tomewin Ra., Upper Currumbin, 17.V.I988, GBM in IBUNAM. I d; Brisbane, 6.V.1953, Y. Beri in UQIC. 1 9 : no locality or date in UQIC. Description Dorsal coloration: Pale orange brown with following areas yellow or yellow ochre: jugae, a short longitudinal band running between eye and ocelli up to posterior edge, poslocular tubercle, most of the anterolateral borders of pronotum, a short longitudinal stripe on middle pronotal disc, apex of scutellum and posterior margin of con- nexivai segments III to VII; tylus and vertex diirk brown; ocellar tubercle, internal and external bor- der of callus and irregular spots on corium black; antennal segments, I to III pale orange yellow with red brown spots on I; segment IV bright red brown with a subbasal bright orange yellow ring; hemelytral membrane brown with basal veins yellow; abdominal segments 1 to III and VII black and IV to VI bright orange yellow; punctures dark red brown or pale orange hazel. Ventral colora- tion: yellow to creamy yellow with following areas black; a small spot located on prolhorax, mesothorax and abdominal sterna III to VI; rostral segment I yellow ochre and II to IV orange hazel; coxae and trochanters yellow; femora yel- low and densely covered with dark red brown discoidal spots; tibiae and tarsi pale yellow orange with dark red brown granules; anterior lobe of metathoracic scent gland creamy yellow and posterior lobe brown hazel; punctures orange hazel; abdominal sterna scattered with pale red brown spots. Female: Coloration: Similar to male. Ab- dominal segments VIII and IX black; genital plates yellow, with punctures orange hazel; 60 MEMOIRS OF THE QUEENSLAND MUSEUM paratergite VIII and IX with or without pale red brown discoidal spot. Measurements: 6 first, then 9 : Head length: 1.45. 1.68; interocellar space: 0.49, 0.64; inter- ocular space: 0.84, I.Ol; width across eyes: 1.64, 1.88; preocular distance: 0.89, 1 .01 ; length anten- nal segments:!, 1.44, 1.68; 11,2.04, 2.64; III, 1.64, 2.00; IV, 1.83, 1.90. Pronotum: Total length: 2.08, 2.84; width across frontal angles: 1.48, 1.82; width across humeral angles: 3.04, 3.88. Scutellar length: 1.48, 1,80; width 1.56, 1.92. Total body length: 11.30, 13.40. Distribution From mountains and plateaux in NENSW and SEQ (Fig. 16B). One specimen labelled ‘Brisbane’ needs confirmation. Etymology Named for the bifid apex of its tylus. ACKNOWLEDGEMENTS The following individuals and institutions provided loans and other assistance: Dr Gordon Nishida (BPBM), Mrs Margerison Knight (BMNH) Dr Norman D. Penny (CAS), Dr Andreas Taeger (DEI), Drs Gordon F. Gross, E.G. Matthews and K.L. Gowlett Holmes (SAMA), Mr T. Weir (ANIC), Dr G.B. Monteith (QMBA), Miss Margaret Schneider (UQIC), Dr I.M. Kerzhner (ZIL) and Dr Antti Janson (ZMUH). Biol. Ernesto Barrera, Mrs El via Espar- za, Mr Felipe Villegas, Biol. Cristina Mayorga, Biol. Laura Gonzalez Garcia and Biol. Albino Luna (IBUNAM) prepared the dorsal view il- lustrations and genital drawings. To the Consejo Nacional de Ciencia y Tecnologia, Mexico (C- ONACyT) and Direccion General del Personal Academico de la Universidad Nacional Autonoma de Mexico (DGPA) provided finan- cial assistance. Dr G.B. Monteith commented on the manuscript. The distribution maps were drawn by Mr G.I. Thompson, Queensland Museum. LITERATURE CITED BERGROTH, E. 1909. Hemiptera Nova Orientalia. Annales de la Societe Entomologique de Belgi- que 53: 184-190. BLOIE, H.C. 1936. Catalogue of the Coreidae in the Rijksmuseum van Natuurlijke Histone. Part III, Coreinae, Second Part. Zoologische Mededelin- gen 19:23-66. BRAILOVSKY, H. 1990. Three new species of Indo- Pacific Colpurini (Hemiptera: Heteroptera: Coreidae). Pan-Pacific Entomologist 66(4): 292- 300. BREDDIN, G. 1900. Hemiptera nonnulla regionis Australicae (figuris duodecim illustrata). En- tomologische Nachrichten 26: 17-46. 1909. Hemiptera insulae Lombok in Museo Ham- burgense asseruata adiectis speciebus nonnullis, quas continent collectio auctoris. Mitteilungen aus dem Naturhislorischen Museum in Hamburg 16: 155-194. DOLLING, W.R. 1987. A mimetic coreid bug and its relatives (Hempitera: Coreidae). Journal of Natural History 21: 1259-1271. HORVATH, G. 1900. Hemipera. Denkschriften der Naturforschenden Gesellschaft Jena 8: 629-642. 1919. Hemipteren von den Aru-und Kei Inseln. Abhandlungcn der Senckenbergischen Natur- forschenden Gesellschaft 35 (3): 305-314. KUMAR, R. 1966. Studies on the biology, immature stages and relative growth of some Australian bugs of the Superfamily Coreoidea (Hemiptera- Heteroptera). Australian Journal of Zoology 14: 895-991. SLATER, J.A. 1975. On the biology and zoogeog- raphy of Australian Lygaeidae (Hemip- lera .Heteroptera) with special reference to the southwest fauna. Journal of the Australian En- tomological Society 14:47-64. STAl, C. 1861. Miscellanea hemipterologica. Stet- tiner Entomologische Z^eitung 22: 129-153. 1873. Enumeratio Hemipterorum III. Enumeratiu Coreidarum Africae, Asiae et Australiae. Kngliga Svenska Vetenskaps Akademiens Hand- lingar2(2): 33-163. WALKER, F. 1871. Catalogue of the specimens of Hemiptera Heteroptera in the Collection of the British Museum. Part IV. London: 1-211. CARIDINA ZEBRA, A NEW SPECIES OF FRESHWATER ATYID SHRIMP (CRUSTACEA: DECAPOD A) FROM NORTHEASTERN QUEENSLAND RAINFOREST JOHN W. SHORT Short, J.W. 1993 12 24: Caridina zebra, a new species of freshwater atyid shrimp (Crus- tacea: Decapoda) from northeastern Queensland rainforest. Memoirs of the Queensland Museum 34(l):61-67. Brisbane. ISSN 0079-8835. A new species of freshwater atyid shrimp, Caridina zebra, from high altitude (>400 m) rainforest, northeastern Queensland, is described and illustrated. It is allied to the typus species-group, characterized by a short, dorsally unarmed rostrum and most closely resembles the wide-ranging species, C. typus H. Milne Edwards, 1837. Morphology of the rostrum and lelson, and the large size of developed ova (1.1 mm length), a feature indicative of abbreviated or direct larval development readily distinguish it from C. typus. A distinctive black and white banded pattern distinguishes C. zebra from all known Indo-West Pacific species. □ Crustacea, Atyidae, Caridina, new species, northeastern Australia, freshwater, rainforest, taxonomy, biology, survival status. John W. Short, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia Australian species of Caridina were last revised by Riek (1953) who recorded 7 species: C gracilirostris De Man, 1892; C. indistincta Calnian, 1926; C. rnccullochi J. Roux, 1926; C nilotica (P. Roux, 1833); C. serratirostris De Man, 1892; C thermophila Riek, 1953; and C. typus Milne Edwards, 1837. These are primarily lowland species, although C serratirostris and C. typus occur at moderate elevations in north- eastern Queensland. High altitude species have previously been described from New Caledonia (J. Roux, 1926a; Holthuis, 1970) and Fiji (Choy, 1983, 1984) in the Southwest Pacific Region. Caridina zebra sp.nov. was first collected by Dr S. Bunn and Mr M. Bray, Griffith University during a limnological study for the proposed Tully-Millstream hydroelectric scheme in 1990. Material from the upper Tully and Herbert R. catchments brought to the Queensland Museum for identification showed affinities with C. typus, but the distinctive unarmed rostrum and large eggs immediately suggested that the species was new. A Queensland Museum expedition to the area (November, 1992) yielded further material and habitat information. Distinctive colour in life confirmed its new species status. Further material was examined from the Queensland Fisheries Johnstone Rivers Survey at the invitation of Mr Warren Lee Long, Northern Fisheries Research Centre, Cairns. Two addition- al specimens were later discovered in the Queensland Museum collection, mis-identified as C. typus. MATERIAL AND METHODS All material examined is deposited in the Queensland Museum collection. Measurements were made with the aid of a stereo microscope and vernier callipers or an eyepiece micrometer. A camera lucida was used for the line drawings. Physico-chemical habitat data were obtained using a TPS LC82 dissolved oxygen meter, mer- cury thermometer, Merck Universal indikator pH 0-14 paper, and an Aquasonic water hardness test kit. Altitudes were estimated from the Australia 1:100 000 topographic survey map series R631. The classification of rainforest types follows Webb & Tracey (1981). Cuticle spination and setation terminology fol- lows Felgenhauer (1992). In particular, ‘hamate setae’ (stout, modified setae) have been classified as spines or spinules by previous atyid workers. The term ‘spine’ is restricted here to large cuticular processes without a basal socket formed by folding or extrusion of the epidermis (e.g. the antennal spine). Generally setae are easily distin- guished by the presence of a well-defined basal socket and often the integument is of different appearance to the surrounding cuticle (i.e. more sclerotinous or less pigmented). Developed ova are defined by the presence of eye spots. Ovigerous females are explicitly recorded whereas non-ovigerous specimens are recorded simply as ‘females’. Abbreviations used in text: QM, Queensland Museum; OCL, carapace length measured from orbital margin to dorsolateral invagination of 62 MEMOIRS OF THE QUEENSLAND MUSEUM posterior margin; TBL, total body length from orbital carapace margin to tip of telson; P, pereiopod; PI, pleopod. Caridina zebra sp.nov. (Figs M) Material Examined Holotype: QM W18718, 6 (3.4 mm OCL), O’Leary Ck, 17°50.7’S, 145°37.7’E, fringing simple nolophyll vine forest, altitude c.750 m, water depth 0. 1-0.5 m, scoop-netted, 06/11/1992, J. Short, P. Davie. Allotype: QM18719, ovig. 9 (5.0 mm OCL), same data as holotype. Paratypes: QM W18120, 26 6 (2.9, 3.4 mm OCL), 13 ovig. 9 9 (4.0 - 4.9 mm OCL), 109 9 (3.5 - 4.8 mm OCL), same data as holotype; QM W18 1 3 1 , 3 d d (3.2 - 3.8 mm OCL), ovig. 9 (4.3 mm OCL), 9 (3.3 mm OCL), same locality data as holotype, 0.5- 1.5 m, trapped, 07/1 1/1992, J. Short, P. Davie; QM W18719, 9 (5 .0 mm OCL). same data as holotype; QM W 1 8 1 24, 6 (3.4 mm OCL), 2 ovig, 9 9 (4.3, 4.4 mm OCL), TuUyR. nr Old Culpa, I7‘=55.5’S, 145°37.6’E. fringing simple notophyll vine forest, altitude c.720 m, water depth 1.0 m, scoop-netted, 06/11/1992, J. Short. P. Davie; QM W18127, 5 1 .0 mm length), suggests direct or abbreviated larval development (Choy, 1991 ). Queensland Museum and literature records of Atyidae from the humid tropics are shown in FIG.4. QM and literature records of Atyidae from the humid tropics. Literature records are those of Roux (1926b), Riek (1953), Williams &*Smilh (1979) and Smith & Williams (i982). Stippling indicates major rainforest areas (Australian Heritage Commission, 1986). A NEW FRESHWATER ATYID SHRIMP 67 Fig.4. A large part of the region has not been adequately surveyed for atyid shrimps and pre- vious records are largely the result of incidental collecting. Apart from C. zebra, only Australatya striolata (McCulloch & McNeill, 1923) and Paratya austraiiensis Kemp, 1917, are known from high elevation areas. C. zebra is the only species endemic to rainforest of the humid tropics. ACKNOWLEDGEMENTS Dr Stuart Bunn and Mr John Marshall, Griffith University brought specimens of C. zebra to my attention. Mr Warren Lee Long of the Northern Fisheries Research Centre, Cairns allowed access to the Johnstone Rivers Survey material and donated representative specimens to the QM. Peter Davie encouraged this work and allocated QM resources to the 1992Tully River expedition. Satish Choy and Peter Davie provided construc- tive criticism of the manuscript. The photograph was printed by Gary Cranitch. LITERATURE CITED AUSTRALIAN HERITAGE COMMISSION. 1986. ‘Tropical Rainforests of North Queensland. Their Conservation Significance.’ A Report to the Australian Heritage Commission by the Rain- forest Conservation Society of Queensland. Spe- cial Australian Heritage Publication Series No.3. (Australian Government Publishing Service: Canberra). CALMAN, W.T. 1926. On Freshwater Prawns of the Family Atyidae from Queensland. Annals of the Magazine of Natural History 9: 241-246. CHOY, S. 1983. Caridina fijiana n. sp. (Decapoda: Atyidae) from Nadarivatu, Fiji. New Zealand Journal of Zoology 10: 147-150, fig. 1. 1984. A new atyid shrimp, Caridina nudirostris sp.nov. (Decapoda, Natantia, Atyidae) from nadrau Plateau, Fiji. Crustaceana46(3): 288-294, fig-1. 1991. The atyid shrimps of Fiji with description of a new species. Zoologische Mededelingen, Leiden 65(27): 343-362, figs 14. FELGENHAUER, B.E. 1992. Chapter 2. External Anatomy and Integumentary Structures. Pp. 7- 43. In: Harrison, F.W. & Humes, A.G. (eds). ‘Microscopic Anatomy of Invertebrates. Vol. 10.’ (Wiley-Liss: New York). HOLTHUIS, LB. 1970. Dudes hydrobiologiques en Nouvelle-Caledonie (Mission 1965 du Premier Institut de zoologie de I’Universite de Vienne). IX. The freshwater shrimps (Crustacea Decapoda, Natantia) of New Caledonia. Cahiers ORSTOM, ser. HydrobioL, 3(2): 87-108, figs 14. MAN, J.G. DE. 1892. Decapoden des Indischen Ar- chipels. Pp.295-527. In: Weber, W. (ed.). ‘Zoologische Ergebnisse einer Reise in Niederl^disch Ost-Indien. Vol.2.’ (E.J. Brill: Leiden). MCCULLOCH, A.R. & MCNEILL, F.A, 1923. Notes on Australian Decapoda. Records of the Australian Museum 14; 49-59. MILNE EDWARDS, H. 1837. ‘Histoire Naturelle des Crustaces, comprenant I’anatomie, la physiologie el la classification de ces animaux.’ Vol.2: 1-532, Atlas, pp. 1-532, pis 1-42. RIEK, E.F. 1953. The Australian freshwater prawns of the family Atyidae. Records of the Australian Museum 23: 11 1-121, figs 1-11. ROUX. J. 1926a. An account of the Australian Atyidae. Records of the Australian Museum 15(3): 237-254. 1926b. Crustaces decapodes d’eau douce de la Nouvelle-Caledonie. Nova Caledonia 4(2): 181- 240, figs 1-56. ROUX, P. 1833. Lettre relative a divers coquilles, cmstaces, insectes, reptiles el oiseaux observes en Egypte. Annales des Sciences Naturelles 28: 72-78, pl.7. SHORT, J.W. & DAVIE, P.J.F. 1993. Two new species of freshwater crayfish (Crustacea: Decapoda: Parastacidae) from northeastern Queensland rainforest. Memoirs of the Queensland Museum 34(1): 69-80. SMITH, M.J. & WILLIAMS, W.D. 1982. Taxonomic Revision of Australian Species of Atyoida Ran- dall (Crustacea: Decapoda: Atyidae), with Remarks on the Taxonomy of iUcGener^ Atyoida and Atya Leach. Australian Journal of Marine and Freshwater Research 33(2): 343-361, figs 1-7. WEBB, L.J. & TRACEY, J.G. 1981 . Australian rain- forest: patterns and change. Pp.607-694. In: Keast, A. (ed.). ‘Ecological Biogeography of Australia.’ Monographiae Biologicae Vol.41. (Dr W. Junk bv Publishers: The Hague). WILLIAMS, W.D. & SMITH, M.J. 1979. A Taxonomic Revision of Australian Species of Paratya (Crustacea: Atyidae). Australian Journal of Marine and Freshwater Research 30: 8 15-832, figs 1-4. 68 MEMOIRS OF THE QUEENSLAND MUSEUM TWO NEW SPECffiS OF FRESHWATER CRAYFISH (CRUSTACEA: DECAPODA' PARASTACIDAE) FROM NORTHEASTERN QUEENSLAND RAINFOREST JOHN W. SHORT AND PETER J.F. DAVIE Short, J.W., & Davie, P.J.F. 1993 12 24: Two new species offreshwater crayfish (Crustacea: Decapoda: Parastacidae) from northeastern Queensland rainforest. Memoirs of the Queensland Museum 34(l):69-80. Brisbane. ISSN 0079-8835. Two new species of freshwater crayfish, Cherax parvus y and Euastacus yigara, are described and illustrated from the upper Tully R. catchment in simple notophyll vine forest, on the Cardwell Ra. C. parvus sp.nov. has several morphological features unique to the genus, and does not appear closely related to any extant species, suggesting a long period of geographic isolation. E. yigara sp.nov. most closely resembles E. balanensis Morgan, 1 988, known from the nearby Lamb and Bellenden Ker Ranges, and can be distinguished by the morphology of the first chelipeds and carapace. Keys are provided to Queensland species of Cherax and Euastacus. f^Crustacea, Parasiacklae, Cherax, Euastacus, Queensland, northeastern Australia, freshwater, rainforest, taxonomy, morf?hology, biology, biogeography, sur\>ival status. John W. Short & Peter J.F. Davie, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; 30 August, 1993, Six species of parastacid freshwater crayfish, all belonging to the genus Euastacus, are current- ly considered endemic to wet highland areas (>8(X) m elevation) in north and mideastem Queensland (Morgan, 1991). No Australian species of Cherax has been considered endemic to wet upland or highland areas and most records have been from elevations less than 4(X) m (Riek, 1951, 1969; Short, 1991, 1993a). In the highlands of New Guinea a distinctive Cherax species- group comprising nine species is endemic to the Wissel Lakes area (Holthuis, 1949, 1950). The discovery of a new species of Cherax in an upland/highland, rainforest catchment in north- eastern Queensland, occurring sympatrically with an undescribed Euastacus and two un- described freshwater shrimps, is of considerable interest. Cherax parvus sp.nov. and E. yigara sp.nov. were discovered during a Queensland Museum expedition to the upper Tully River area in November 1992. The primary purpose of the ex- pedition was to collect fully-developed, adult males of an undescribed species of Macro- brachium (presently being studied as part of a revision of Australian species by J.S.) and obtain habitat data for an undescribed Caridina (described as C. zebra Short, 1993), previously known from the area. These new shrimps were first collected by Dr S. Bunn and Mr M. Bray, Griffith University, during a limnological study for the proposed Tully-Millstream hydroelectric scheme in 1990. MATERIAL AND METHODS All material examined is housed in the Queensland Museum collection unless otherwise stated. Measurements were made using vernier callipers and line drawings with the aid of a camera lucida. Photographs were taken with a 35 mm SLR camera and flashes. The terms ‘cheliped’ and ‘chela’ refer to the first pereiopod unless stated otherwise. Euas- tacus terminology largely follows Morgan (1986), particularly spination and sternal keel characters. Colour patterns, habitats and burrow types for Queensland Cherax are derived from the work of Riek (1951, 1969), Horwitz & Richardson (1986) and personal observations by J.S. Rainforest and climatic zone terms follow Webb & Tracey (1981). Physico-chemical habitat data were obtained using a TPS LC82 dissolved oxygen meter, mer- cury thermometer, Merck Universalindikator pH 0-14 paper, and an Aquasonic water hardness test kit. Altitudes were estimated from the Australia 1 : 1(X) (XX) topographic survey map series R63 1 . Abbreviations used in text: QM, Queensland Museum; OCL, carapace length from orbital mar- gin to dorsolateral invagination of posterior mar- gin; T, thoracic stemite, T6, thoracic stemite six etc., imm., immature; P, pereiopod; Pr, sternal keel processes, Prl, first sternal keel processes, etc; Qld, Queensland; N.T., the Northern Ter- ritory. 70 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS Cherax parvus sp.nov. (Figs M) Material Examined Holotype: QM W1812L 6 {22 J mm OCL), O’Leary Ck, tributary of the Tuliy R. above Koom- booloomba Dam, 17°50.7’S. 145°37.7’E, in short phreatic burrows under rocks, rocks/sand substratum, leaf litter, fallen timber, low to moderate flow, water clarity high, fringing simple notophyll vine forest, pH 5.5, hardness <10 ppm, water temperature 20“C, dis- solved oxygen 6 ppm, altitude c. 750 m, water depth 0. 1-0.3 m, 06/1 1/1 992, J. Short, P. Davie. Paratypes: QMW18I23, 9 (14.0 mm OCL), upper TuUy R. near Old Culpa above Koombooloomba Dam, 17°55.5’ S, I45°37.6’E, amongst leaf litter, rocks/sand substratum, fallen timber, low to moderate flow, water clarity high, fringing simple notophyll vine forest, pH 5.5, hardness <10 ppm, water temperature 18°C, dis- solved oxygen 5.8 ppm, altitude c. 720 m, water depth 1 m, netted, 06/11/1992, J. Short. P. Davie; QM W18I33,6(J<3(lZ4-21.9mmOCL),29 9 (12.6,16.0 mm OCL), imm. (9.9 mm OCL), same collection data as holotype except collected from short phreatic bur- rows under rocks in small, clay bottomed pool above main creek; QMW17494, 10c? 6 (9.0-16.6 mm OCL), 49 9 (8.3-14.4 mm OCL), same locality and habitat data as holotype, water depth 0. 1-0.5 m, 06/1 1/1992, J, Short, P. Davie. Description of Male Holotype Cephalothorax. Carapace punctate; slender, breadth c. 0.5 X CL (0.5-0.6 in paratypes), depth c. 0.6 X CL (0.5-0.6 in paratypes); cephalon with few tubercles ventrally; branchiostegites uninflated, with few indistinct tubercles along ventral cervical groove. Rostrum slender, triangular, tapering strongly in distal third, length c. 1.4 X breadth (L0-L6 in paratypes), reaching distal penultimate segment of antennular peduncle, dorsally flattened, not recurved dislally, punctate along inner margins of lateral carinae, smooth medially, setose distally near acumen; lateral carinae moderately developed, commencing at base of rostrum, ter- minating at acumen, without tubercles or spines; ventro-lateral margins setose; acumen blunt, up- turned (strongly in paratypes^ Post-orbital carinae almost obsolete, unarmed anteriorly, excavated with well separated puncla- tions, commencing close to orbital margin of carapace, medially curved anteriorly, diverging posteriorly; cervical groove setose (setation less developed in smaller paratypes); branchiocardiac grooves obsolete. Eyes with cornea large, globular, well pig- mented; eyestalks largely concealed by rostrum, Scaphocerite length slightly exceeding length of rostrum, broadest at mid-length; lamina broadly rounded mesially, lateral margin terminating in well developed spine. Antennal peduncle setose ventrally, coxocerite acute anteriorly, basicerite without lateral spine (acute in smaller paratypes). Epistomc strongly concave medi^ly, setose anteriorly (densely setose in paratypes), with dis- tinct tubercles laterally, lateral margins entire. Mouthparts without unique features for genus. Branchial formula typical for genus (cf. Holthuis, 1949) with posterior arthrobranch above P4 reduced. Pleurocoxal lappets well developed; operculiform; fringed with long plumose setae; lappet between P4-5 unusually thin, circular, plumose setae very long. Sternal keel sharp posteriorly, more rounded anteriorly, lateral processes setose, without ob- vious pores, Pr4 juxtaposed, Pr5 similar, Chelipeds. First chelipeds large, chela length exceeding carapace length (shorter in juvenile paratypes), distal merus reaching end of scaphocerite (carpus reaching end of scaphocerite in juvenile paratypes), isomorphic. Outer margin of chela moderately convex; fingers slightly gaping (without gape in un- developed paratypes); opposing edges densely setose ventrally with poilex setation continuing onto manus, bearing rounded teeth, one tooth large and prominent on dactylus at mid-length, similar slightly smaller tooth on poilex (teeth isomorphic in paratypes); poilex broad basally, evenly tapering; dactylus broadest at mid-length, equal to manus in length; manus moderately broad, breadth equal to len^h (greater than length in juveniles), dorsum slightly convex long- itudinally, strongly convex laterally, mesial mar- gin serrations continuing onto distal half. Carpus bearing large broad uncinate mesial spine (missing on left cheliped), angle of disto- ventral condyle not produced into spine or tubercle, ventro-mesial angle similar, followed by few tubercles postero-mesially, disto-mesial angle glabrous, disto-dorsal condyle strongly developed, enlarged laterally to form broad sclerotinous plate. Merus of typical shape, dorsal carina without obvious tubercles or spines (small distal tubercle in paratypes), ventral surface sparsely setose, with numerous strong cuticular processes, one or two larger than remainder. Colour Body dark brown; first chelipeds dark brown TWO NEW FRESHWATER CRAYFISH 71 FIG.l. Cherax parvus sp.nov., QM W18121, c? holotype (22.7 mm OCL). 72 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Cherax parvus sp.nov., QM W18121, S holotype, anterior carapace and rostrum. Scale bar divisions in millimetres. FIG. 3. Cherax parvus sp.nov., QM W18121, 6 holotype, A. ventral first right cheliped. B. dorsal first left cheliped. Scale bar divisions in millimetres. disto-dorsally with reticulated pattern on manus, fingertips orange, proximal merus and ischium orange, ventral manus and fingers cream; second chelipeds and ambulatory legs greenish-cream dorsally, light cream ventrally. Biology Freshwater, short phreatic burrows under rocks or amongst leaf litter in shallow open water ( 1 m), rocks/sand or clay substrates, fringing simple notophyll vine forest, 720-750 m altitude, zero to moderate flow, water clarity high. Recorded physico-chemical tolerances; pH 5.5, hardness <10 ppm, water temperature 18-20°C, dissolved oxygen 5. 8-6.0 ppm. Close association of this species with rainforest is suggested by its absence in Nitchaga Ck, another tributary of the Tully R., above Tully Falls. This creek drains open sclerophyll forest and dries to a series of pools during extended dry periods (as in November 1992, when sampled). Here it is replaced by a species of the wide-rang- ing "depressus* complex (sensu Riek, 1951). This is the smallest species so far described in the genus. The holotype, which has a post-orbital body length of only 55 mm and a post-orbital carapace length of 23 mm, shows the robust, broad chelae with an obvious gape, and the well developed dentition on the opposing margins of the fingers, characteristic of a well developed male. Of the 24 paratypes only two exceed 1 7 mm in post-orbital carapace length. At the type locality C. parvus is sympatric with E. yigara sp.nov., Caridina zebra Short. 1993, and an undescribed Macrobrachium. A F1G.4. Cherax parvus sp.nov., QM W18133, d paratype (17.4 mm OCL), pleurocoxal lappets. A. P4-5. B. P3-4. C. P2-3. D. PI-2. Scale bar I mil- limetre. TWO NEW FRESHWATER CRAYHSH 73 Distribution Recorded from the upper Tully R. and its tributary, O’Leary Ck, above Koombooloomba Dam, at 720-750 m altitude, Cardwell Ra., NEQ. Survival Status Probably secure. Although abundant at the type locality, more data is required on the distribution of the species and its relative abundance at other sites. Only one specimen was collected in the main Tully R. at Old Culpa where the substrate was clean sand. Two higher elevation sites (c. 900 m), approx. 12 km past O’Leary Ck, travelling east on the main forestry road, failed to produce crayfish, although C zebra Short, 1993, was very abundant. Both sites had a silty sand substratum and the fringing vegetation showed indications of regrowth, possibly as a result of timber logging. Etymology Latin (adjective), meaning small. Systematic Position The distribution of Cherax in the humid tropics of north Queensland is shown in Fig.5. Although records of the new species lie within the broad geographical range of ‘C. depressus' (sensu Riek, 1951), C. parvus does not appear closely related to that complex. The following unique features suggest a long period of geographic isolation. 1. The almost obsolete post-orbital carinae commencing very close to the orbital carapace margin. 2. The disto-dorsal condyle on the carpus of the first chelipeds very strongly developed and en- larged laterally to form a broad sclerotinous plate. 3. The pleurocoxal lappet between P4-5 un- usually thin and fringed with very long plumose setae. 4. The branchiostegites uninflated in developed specimens. With regard to the last character, inflated branchiostegites and narrow areolae are generally correlated with the enlargement of the branchial chambers and increased surface area of gills. This is most pronounced in fully grown adults of species living in poorly oxygenated habitats. The relatively small branchial chambers of C. parvus may reflect its preference for cool, well- oxygenated, rainforest streams. In contrast to the unarmed rostrum of C. parvus ^ all highland New Guinean species, except C. monticola, have two or more pairs of lateral processes (tubercles or spines) on the rostrum. In C. monticola there is generally one or two pairs, FIG.5. QM locality records of freshwater crayfish in the humid tropics. Hatched areas are cloudy wet mountains inferred as major refugia (Webb & Tracey, 1981). Question marks indicate poorly-sampled refugial areas on which future investigations should focus. but occasionally none or three. This species, how- ever, has two well-defined uncalcified patches on the first cheliped of mature males and is clearly allied to the quadricarinatus species-group (Short, 1991). KEY TO QUEENSLAND CHERAX 1 Adult rostrum with obvious, well-developed spines on lateral carinae; cervical groove of adults bor- dered by number of w'ell developed spines 2 — Adult rostrum without well-developed lateral spines, lateral carinae terminating in blunt process or unarmed; cervical groove bordered by blunt tubercles or small spines 4 2(1) Rostrum with 1 pairoflateral spines; lateral pollex of first chelipeds of mature males without uncal- cified region C. disparKiek, 1951 [Body bluish or green-grey, chelipeds dark blue in ma- ture specimens, lighter blue ventrally, chelae orange in juveniles. Between Elliot R. & Brisbane R.,SEQ, includ- ing large sand islands. Permanent freshwater, including 74 MEMOIRS OF THE QUEENSLAND MUSEUM FIG.6. Euastacus yigara sp.nov., QM 18134, 9 holotype (22.9 mm OCL). !l TWO NEW FRESHWATER CRAYHSH 75 acidic perched lakes and large coastal streams. Short burrows in, or connected to open water.) — Rostrum with 2 or more pairs of lateral spines; lateral pollex of first chelipeds of mature males with distinct uncalcified region (most developed in fully grown males) 3 3(2) Post-rostral carinae short, terminating slightly be- hind line of post-orbital spines; dorso-medial rostrum slightly concave, lateral carinae moderate- ly elevated ....C. rhynchotus Riek ,951 [Body green-brown dorsally, chelipeds similar except lateral chela orange, and uncalcified portion on lateral pollex of mature males peach cream. Northern Cape York Pen. and Badu Is., Torres Strait. Acidic coastal swamps and perched lakes. Burrows connected to open water or water table.) — Post-rostral carinae long, almost extending to end of post-orbital carinae. Dorso-medial rostrum strongly concave, lateral carinae well elevated C. quadricarinatus (von Martens, 1 868) [Body colour from light green-blue to almost black, light flecks on lighter specimens, abdomen with purple bands dorso-laterally. chelipeds slightly darker than body, manus patterned dorsally, base of dactylus white, uncal- cified portion on pollex of mature males red. Exorheic drainage systems from Daly R. N.T., to Normanby R., NEQ. Permanent freshwater, absent from acidic swamps and perched lakes. Short burrows in, or connected toopen water.] 4(1) Antero-dorsal carpus of first chelipeds with con- dyle developed into long sclerotinous plate; post- orbital carinae commencing very close to carapace margin; adults with branchiostegites not obviously inflated C parvus sp.nov. [Dorsum dark brown, chelipeds dark brown disto-dorsal- ly, patterned on dorsal manus, finger tips orange, proximal segments orange. Upper Tully R. catchment, NEQ. Rainforest streams. Short burrows in open water.] — Antero-dorsal carpus of chelipeds with condyle typically developed into rounded lobe; post-orbital carinae widely separated from carapace margin; adults with brachiostegites obviously inflated ... 5 5(4) Branchiocardiac grooves well defined on adults, very narrowly separated or confluent anteriorly; mesial margin of manus of cheliped generally very short, much shorter than dactylus C. punctatusCXdsk, 1936 [Mary R. catchment, SEQ. Burrows independent of open water or the water table, located on clay hillsides.) — Branchiocardiac grooves variably defined on adults, when pre.sent well separated throughout length; mesial margin of manus of cheliped of moderate length, about equal in length to, or slight- ly shorter than, length of dactylus 6 6(5) Serrations on manus of chelipeds clearly terminat- ing within proximal half of mesial margin C. cartalacoolah Short, 1993 [Dorsum blue-grey to rusty brown, venter vivid purple or blue, meral/carpal joint red.Cape Flattery, NEQ. Coas- tal sand dune wetlands, acid water. Burrows connected to open water or the water table.) — Serrations on manus of chela terminating at least halfway along mesial margin 7 7(6) Setation on ventral pollex of chelipeds very broad, clearly continuing onto manus C robustus Kick, 1951 [Body bluish or purple-grey to black, chelae of similar colour dorsally, ventrally vivid purple or blue, Joints red. Fraser Is. and Cooloola to N. Slradbroke Is. SEQ.Coastal sand dune wetlands, acid water. Burrows connected to open water or water table. ) — Setation on ventral pollex of chelipeds restricted to opposing edge, not continuing onto manus 8 8(7) Mesial surface of cheliped carpus usually with well developed setal pubescence; post-orbital processes obsolete ‘C. destructor' complex' [Body from pale cream to brown or deep blue, chelipeds slightly darker with dorsal manus patterned, joints red. Wide-ranging inland and southern Australia; endorheic catchments, Murray-Darling system, and the Dawson R., MEQ. Permanent and ephemeral water bodies. Bur- rows in, Of connected to open water.) — Mesial surface of cheliped carpus without distinct setal pubescence, generally asetose, sometimes with scattered short setae; post-orbital processes moderately developed, spinate in juveniles, blunt in adults ‘C. depressus' complex^ [Body green-brown to blue-black or dark brown, often mottled in females, dorsal manus of chelae generally patterned, finger tips orange, joints red or orange. The Gold Coast, SEQ to Jardine Swamps, Cape York Penin- sula. Ephemeral waters including acidic swamps. Bur- rows connected to open water or the water table.] ' Includes the nominal species, C. davisi Clark, 1941 and C. destructor Oark, 1936, in Queensland ^ Includes the poorly-defined nominal species, C. depressus Riek, 1 95 1 and C cam\sensis, C. gladsfonensis, C. was- selli and C. urospinosus, all of Riek, 1969. Euastacus yigara sp.nov. (Figs 6-8) Material Examined Holotype: QM W18134, 9 (22.9 mm OCL), O’Leary Ck, 17°50.7’S, I45°37.7'E, freshwater, rocks/sand, leaf litter, fallen timber, fringing simple nolophyll vine forest, under rock in pool along edge of creek, pH 5.5, hardness <10 ppm, water temperature 20°C, dissolved oxygen 6 ppm. altitude c. 750 m, water depth 0.1-0.5 m, 07/I1/T992, J. Short, P. Davie. Paratype: QM W 1 8 1 25. imm. (11.1 mm OCL), same 76 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. Euastacus yigara sp.nov., QM 18134, 9 holotype (22.9 mm OCL), anterior cephalon and rostrum. Scale bar divisions in millimetres. locality and habitat data as holotype, 06/11/1992, J. Short, P. Davie. Description of Female Holotype Cephalothorax, Carapace slender, breadth c. 0.5 X CL; cephalon with low, blunt, anteriorly- directed tubercles anterolaterally, extending in a band ventrally; thorax punctate, punctations bear- ing short setae, armed with 1-2 low cervical tubercles, otherwise smooth. Rostrum moderately long, acute, basally broad, length c. 1.2 X breadth, reaching base of anten- nular flagella; acumen spinous, upturned, sclerotinous at tip; lateral carinae well developed, slightly convex, armed with 3-4 pairs of blunt processes (more prominent and acute in juvenile paratype), processes diminishing in size f>osteriorly (3 on right side, but with no trace of fourth; on left side second spine missing but fourth present at about level of posterior edge of cornea), extending onto proximal half of rostrum; rostral carinae extending posteriorly to about half distance between posterior margin of orbit and post-orbital spine. Post-orbital carinae long, sulcate, sulci setose, margins raised, armed with single blunt spine anteriorly. Branchiocardiac grooves distinct, relatively widely separated, areola divergent anteriorly. Antennal scaphocerite with lateral margin straight, unarmed, apex acute, greatest breadth at mid-length; basicerite spine (suborbital spine) moderately developed; coxocerite with 2 small acute processes (on right), 1 small acute process and 2 more indistinct processes (on left); basal exopod spine (basipodite spine) absent. Epistome (interantennal spine) moderately elongate with weakly scalloped margins, reaching distal end of first segment of antennal endopod. Sternal keel lacking spines; Prl posterior mar- gin sloped, processes apart and parallel; Pr2 apart and open; Pr3 with posterior edges straight, or very slightly rounded; Pr4 with anterior edges angular, posterior borders sharp and convex. Chelipeds. Moderately stout, dimorphic, left larger and more robust than right; left cheliped FIG.8. Euastacus yigara sp.nov., QM 18134, 9 holotype (22.9 mm OCL), dorsal left chela. Scale bar divisions in millimetres. TWO NEW FRESHWATER CRAYHSH 77 dactyl length c. 0.6 X propodus length; propodus length c. 2.1 X width. Dactylus without spines above tuberculate op- posing edge, bearing small apical mesial spine, mesial basal spines absent, dactylar groove deep. Propodus with single lateral spine row, spines blunt, becoming more prominent and closer together distally, not formed into distinct ridge; ventral spine row vestigial, single blunt mid- lateral spine; 5 marginal mesial spines; dorsal apical spines absent; without spines above oppos- ing edge, opposing edge with single prominent rounded tubercle at about proximal quarter; 1 small distinct tubercle and 2 smaller indistinct tubercles on base of dorsal pollex lateral to dac- tylar base; uneven row of tubercles dorso-medial- ly on manus, ventral pollex with 1-2 tubercles lateral to dactylar base. Carpus with deep dorso-Iongitudinal groove; 3-4 mesial spines, distal-most the largest and offset from remainder; minute lateral spine; ar- ticulation spine absent; small but prominent dorso-mesial tubercle inside longitudinal groove; minute dislo-ventral process medially, 2 ventro- mesial processes, 1 slightly proximal to disto- ventral process the other placed below large distal carpal spine (processes much smaller on right cheliped). Merus with 1-3 (1 on left, 3 on right) small dorsal spines; outer spine absent. Abdomen. Bearing sparse short tufts of setae, smooth, punctate, second pleurae with 2 indis- tinct tubercles vcntrally. Dorsal telsonic spines absent; surface anterior to small disto-lateral spines coarsely punctate, punctalions bearing tufts of setae; posterior surface with fine long- itudinal ridging. Colour General colour greenish-grey. Biology Freshwater, under rocks in shallow water (<0.5m), rocks/sand substratum, fringing simple notophyll vine forest, c. 750m altitude, water clarity high. Recorded physico-chemical toleran- ces : pH 5.5, hardness < 1 0 ppm, water temperature 18-20°C, dissolved oxygen 5. 8-6.0 ppm. At the type locality E. yigara is sympatric with C. parvus sp.nov., Caridina zebra Short, 1993 and an undescribed Macrobrachium. Distribution Presently known from O’Leary Ck, a tributary of the upper Tully R. above Koombooloomba Dam, Cardwell Ra., NEQ. Survival Status Unknown. More data is required on the abun- dance and distribution of the species. The two type specimens were located during 2 days col- lecting in the upper TuIIy area and c. 4 man-hours at the type locality. The species may be more common at higher elevations as other north Queensland species are restricted to altitudes above 800 m (Morgan, 1991). Etymology The aboriginal word for ‘crayfish’ in the Djirbal language group of the upper Tully R. area (Dixon, 1972). The specific epithet is to be treated as a noun in apposition. Systematic Position E. yigara sp.nov. most closely resembles E. balanensis Morgan, 1988, which is known from the nearby Lamb and Belienden Ker Ranges. It can be separated from that species by the follow- ing features: 1 . An obvious tubercle on the disto-mesial car- pus inside the longitudinal groove. 2. The presence of 3-5 distinct dorso-medial tubercles in an irregular longitudinal row on the manus, with small setose punctations anteriorly. In E. balanensis the dorsal manus is ornamented with broad punctations but is without elevated tubercles. 3. The rostrum armed with blunt marginal tubercles which extend onto the proximal half. On E. balanensis they are restricted to the distal half. 4. A relatively wider areola which diverges anteriorly rather than having subparallel margins. Fifteen species of Euastacus arc now known from Queensland. The distribution of Euastacus species in the humid tropics and their close as- sociation with the cloudy, wet, mountain areas, inferred as rcfugial by Webb & Tracey (1981), is illustrated in Fig. 5. Question marks indicate poorly-sampled rcfugial areas on which future investigations should focus. Of particular interest are the Walter Hill, Herberton and Hugh Nelson Ranges, between the present distribution of E. yigara and E. balanensis. KEY TO QUEENSLAND EUASTACUS (Modified from Morgan, 1991) 1 Well-defined, longitudinal groove on dorsal carpus of cheliped 2 78 MEMOIRS OF THE QUEENSLAND MUSEUM — Without well-defined, longitudinal groove on dor- sal carpus of cheliped, at most with broad, shallow depression 14 2(1) Large species with well-developed abdominal spines; large adults usually with distinct spines on thorax and/or telson and 2 spines on mesial carpus of cheliped 3 — Small species with reduced or absent abdominal spines; large adults with spines on thorax and telson absent or minute; 3 or more spines on mesial carpus of cheliped 6 3(2) No mesial basal spines on cheliped dactylus. E. vfl/enm/w^Riek, 1951 [Body deep green or green-brown. From Cumimbin Ck SEQ to Clarence R., NSW. Wet scleropbyll forest and rainforest, 0-600m.] — One or more mesial basal spines on cheliped dac- tylus 4 4(3) Large adults with dorsal spines absent on thorax and absent or very small on telson E. suIcatusKlek, 1951 (Body bright blue, red or brown. East flowing streams from Logan R.,SEQ, to Richmond R., NSW;also the Condamine R. flowing west. Wet sclerophyll forest and sublropical/rainforest, >300 m.] — Large adults with distinct spines on thorax; spines on telson usually distinct and often large 5 5(4) Large adults with distinct dorsal spine(s) on ab- dominal somites 2-4 E. sunoniC\ark, 1941 [Body very dark, spines on thorax and abdomen bright red. Severn R. and Dumaresque R., SEQ, flowing west, and Clarence R., NENS W, flowing east. Dry sclerophyll forest and heath, >680 m.] — Large adults with dorsal spines absent on ab- dominal somites 2-4 E, hystricosus Riek, 195 1 [Body, including spines, deep green. Conondale and Blackall Ranges, SEQ. Wet sclerophyll forest and rain- forest, >550 m.j 6(2) Spines above opposing edges of chelae in rows reaching almost full length of fingers; ventral spine row on lateral chelae well developed 7 — Spines above opposing edges of chelae not reach- ing beyond mid-length of fingers and sometimes absent; ventral spine row on lateral chelae absent or poorly developed (fewer than 4 spines) 9 7(6) Post-orbital spine absent; Isl processes of sternal keel well separated and converging anteriorly E. monteithorum Morgan, 1989 [Body and chelipeds dark brown dorsally except for orange merus, ventral body orange. From Kroombil Ck, Kroombit Tops, Calliope Ra., SEQ. Rainforest. >800 m elevation.] — Post-orbital spine present; 1st processes of sternal keel close and parallel 8 8(7) Numerous tubercles on dorsal and ventral surfaces of chelae near base of pollex, extending some distance along finger; without dorso-mesial tubercle on carpus of cheliped E. Morgan, 1989 [From North Ck, Mt. Elliot, NEQ. Rainforest, 1000 m .] — Only a few tubercles near base of fingers, these not extending along pollex; dorso-mesial tubercle/s present on carpus of cheliped E. eungella Morgan, 1 988 [Body green-blue with purple tinges, especially on chelae. Tributaries of Pioneer R, Clark Ra., MEQ. Rain- forest, >740 m.] 9(6) First processes of sternal keel apart 10 — First processes of sternal keel close together 11 10(9) Carpus of cheliped with dorsal disto-mesial tub- ercle, manus with irregular longitudinal row of dorso-medial tubercles; rostrum with marginal processes extending onto proximal half E. yigara sp.nov. [Body green-grey. O’Leary Ck, tributary of upper Tully R, Cardwell Ra.. NEQ. Rainforest, 760 m.) — Carpus of cheliped without dorsal disto-mesial tubercle, manus without dorso-medial row of tubercles; rostrum with marginal processes restricted to distal half E. balanensis Morgan, 1988 [Body green-brown or olive, abdominal spines paler brown or green. Lamb and Bellenden Ker Ranges, NEQ. Rainforest, >800 m.] 1 1(9) Suborbital spine large E. setosus Riek, 1 956 [Body deep red-brown with green on chelae. Mt. Glorious, D’Aguillar Ra., SEQ. Rainforest, >500 m.] — Suborbital spine medium-sized 12 1 2( 1 1 ) Third sternal keel processes parallel E. urospinosus Rick, 1956 [Body red-brown with green tinges. Conondale and Blackall Ranges, SEQ. Rainforest, >240 m.] — Third Sternal keel processes not parallel 13 13(12) Adult epistome of moderate length, clearly shorter than first segment of antennal endopod. E. maidaeRick, 1956 [Body orange-brown, chelipeds similar except for finger tips blue. Upper Currumbin and TallebudgeraCks, SEQ. Rainforest, >150 m.] — Adult epistome elongate, reaching to distal end of first segment of antennal endopod or beyond E. yagara Morgan, 1988 [Flaggy Ck, Mistake Mts, SEQ. Rainforest, 900 m.] 14(1) Lateral ventral spine row on chelae poorly developed, usually 4-5 spines on mesial margin of manus; rostrum U-shaped on large specimens E.fleckeri (Watson, 1935) TWO NEW FRESHWATER CRAYHSH 79 [Body very dark green, thoracic and abdominal spines red. Mt. Carbine Tableland, NEQ. Rainforest, >1000m.] — Lateral ventral spine row on chelae well developed and reaching apex of pollex; 6-9 spines on mesial margin of manus; rostrum usually triangular. E. Monroe, 1977 [Body dull blue-purple. Mt Finnigan and Thornton Peak, NEQ. Rainforest, >1000 m.] DISCUSSION The presence of four undescribed freshwater decapods in the upper Tully R. catchment, three of which appear endemic (the two crayfish species described above and an undescribed Mac- robrachium), suggests something unique about the physiographic/climatic history of the area. Features which may have promoted speciation include: 1. A well-developed, physical, dispersal barrier in the form of Tully Falls, which is a sheer drop of 200 m. Potential crustacean competitors and predatory fish such as Hephaestus fuUginosus (Sooty Grunter), Kuhlia rupestris (Jungle Perch) and Tandanus tandanus (Eel-tail Catfish) are also isolated by this barrier. 2. The high, consistent, annual rainfall (2689 mm average at Koombooloomba Dam) and plateau topography, resulting in an unusually large, upland/highland catchment of permanent streams of high water quality and cool water temperature. 3. A cloudy, wet, rainforest environment which may have acted as a refugium during periods of climatic warming and decreasing rainfall. The Cardwell Ra. was listed by Webb & Tracey (1981) as one of their inferr^ refugial areas for the humid tropics. 4. The long, stable geological history of the Cardwell Range, which is largely granitic in com- position (De Keyser, 1964). Neighbouring granite massifs such as the Lamb and Bellenden Ker Ranges are of Permo-Triassic origin (Wil- mott et al, 1988). Unlike the nearby Atherton Tableland, there is presently no evidence of larva flows on the Cardwell Range during the Cainozoic (De Keyser, 1964). ACKNOWLEDGEMENTS We are grateful to the Queensland Forest Ser- vice for permits to transverse and collect within State Forest 605 in the Atherton Forestry District. Gary Cranitch printed the photographs. LITERATURE CITED CLARK, E. 1936. The freshwater and land crayfishes of Australia. Memoirs of the National Museum of Victoria. 10:5-58, pis 1-11. 1941. New species of Australian freshwater and land crayfishes (family Parastacidae). Memoirs of the National Museum of Victoria 12:31-40, pi. 10. DDCON, R.M.W. 1972. ‘The Dyirbal Language of North Queensland.’ Cambridge Studies in Lin- guistics. 9. (Cambridge University Press: Cambridge). HOLTHUIS, L.B. 1949. Decapoda Macrura with a revision of the New Guinea Parastacidae. Zoological Results of the Dutch New Guinea Expedition 1939. No 3. Nova Guinea 5:289-328, pis 2-9. 1950. Results of the Archbold Expeditions. No.63. The Crustacea Decapoda Macrura collected by the Archbold New Guinea expeditions. American Museum Novitates 1461:1-17, figs 1-4. HORWITZ, P.H.J. & RICHARDSON, A.M.M. 1986. An Ecological Classification of the Burrows of Australian Freshwater Crayfish. Australian Jour- nal of Marine and Freshwater Research 37:237- 42. KEYSER, F. DE. 1964. Innisfail, Queensland - 1:250 (XX) Geological Series. Bureau of Mineral Resources, Australia, Explanatory Notes SE/55- 6 . MARTENS, E. VON. 1866. On a new species of Astacus. Annals of the Magazine of Natural His- tory (3) 17:359-60. MONROE, R. 1977. A new species of Euastacus (Decapoda: Parastacidae) from North Queensland. Memoirs of the Queensland Museum 18:65-67. MORGAN, G.J. 1986. Freshwater crayfish of the genus Euastacus Clark (Decapoda, Parastacidae) from Victoria. Memoirs of the Museum of Vic- toria 47(l):l-57, figs 1-23. 1988. Freshwater crayfish of the genus Euastacus Clark (Decapoda: Parastacidae) from Queensland. Memoirs of the Museum of Victoria 49(l);l-49, figs 1-25. 1989. Two new species of the freshwater crayfish Euastacus Clark (Decapoda: Parastacidae) from isolated high country of Queensland. Memoirs of the Queensland Museum 27(2):555-562, figs 1-3. 1991. The spiny freshwater crayfish of Queensland . The Queensland Naturalist 3](l-2):29-36, figs 1-5. RIEK, E.F. 1951. The freshwater crayfish (family 80 MEMOIRS OF THE QUEENSLAND MUSEUM Parastacidae) of Queensland. Records of the Australian Museum 22:368-388, figs 1-13. 1956. Additions to the Australian freshwater crayfish. Records of the Australian Museum 24:1-6. 1969. The Australian freshwater crayfish (Crus- tacea: Decapoda: Parastacidae), with descrip- tions of new species. Australian Journal of Zoology 17:855-918, figs 1-20. ROUX, J. 1933. Note sur quelques Crustaces decapodes d’eau douce provenant de I’Australie seplentrionale. Revue suisse Zoology 40:343- 348. SHORT, J.W. 1991. Cherax nucifraga^ a new species of freshwater crayfish (Crustacea: Decapoda: Parastacidae) from the Northern Territory, Australia. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 8(1 ): 1 1 5- 120, figs 1-4. 1993a. Cherax cartalacoolah, a new species of freshwater crayfish (Decapoda; Parastacidae) from northeast Australia. Memoirs of the Queensland Museum 33(l):55-59, figs M. 1993b. Caridina zebra, a new species of freshwater atyid shrimp (Crustacea: Decapoda) from north- eastern Queensland rainforest. Memoirs of the Queensland Museum 34(l):61-67. SOKOL, A. 1987. Yabbies at Dalhousie Springs, Northern South Australia; Morphological Evidence for Long Term Isolation. Transactions ofthe Royal Society of South Australia 1 1 1 (3-4): 207-209, figs 2. WATSON, K. 1935. A new Astacopsis from north Queensland. Memoirs of the Queensland Museum 10:232-235. WEBB, L.J. & TRACEY. J.G. 1981. Australian rain- forest: patterns and change. Pp.607-694. In Keast, A. (ed). ‘Ecological Biogeography of Australia’. Monographiae Biologicae Volume 41. (Junk: The Hague) WILMOTT, W.F., TREZISE, D.L., O’FLYNN, M.L., HOLMES, P.R. & HOFMANN, G.W. 1988. Cairns Region, Sheets 8064 & 8063 (part), Queensland - 1:100 000 Geological Map Com- mentary. Queensland Department of Mines. THE BIODIVERSITY OF AUSTRALIAN MYGALOMORPH SPIDERS. 1. TWO NEW SPECIES 0¥NAMIREA (ARANEAE: DIPLURIDAE). ROBERT! RAVEN Raven, RJ. 1993 12 24: The biodiversity of Australian mygalomorph spiders. I. Two new species oiNamirea (Araneae:Dipluridae). Queensland Museum 34(l):81-88. Brisbane. ISSN 0079-8835. Two new species of Namirea^ N. dougwallacei and N. johnlyonsi, are described from northern Australia. This is the first record of males from the region. Males of N. dougwallacei appear to pose a phylogenetic dilemma because of homoplasies with Australothele. □ Mygalomorphae, Dipluridae, taxonomy, phylogeny, Namirea, biogeography, biodiver- sity, disturbance, Australia. Robert J. Raven, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia: 4 July 1993. The genus Namirea was described by Raven (1984) along with other ischnotheline diplurids from Australia. Namirea included five new species. The genus is found in areas from Mt Lewis, NEQ, through southeastern Queensland, to the Blue Mountains, NSW. Males of the genus were known only in the southern portion of its range. The most northerly known males of Namirea were of N. planipes which is widespread in open forest areas and found commonly in road- side embankments in southern Queensland. Here two new species including males from the north- ern part of the range are described. Their mor- phology, especially that of males, allows a better understanding of the relationships of the group. Methods are explained in Raven (1984). SYSTEMATICS Namirea Raven, 1984 Namirea dougwallacei sp. nov. (Figs 1-4. Tables 1, 2) Material Examined Holotype d. Allotype 9, Mt Archer, summit, 23°2rS 150°35’E, MEQ, D. Wallace, May 1984, QMS19706, 19707. Diagnosis Males differ from those of all other Namirea with known males in having a mounded spur on tibia I and from Australothele in lacking the secondary spine on the spur and the metatarsal thorn on leg II. Females differ from those of N. eungella in the sp^rmathecae having three, rather than only two, spiralled loops, a larger basal lobe and longer apical lobe. Medium-sized; carapace length ca. 5. Preening combs present on metatarsi I, II, IV. Sper- mathecae two, each an inner long sclerolised spiralled duct with short straight unsclerotised outer lobe, males. Tibia I subdistally incrassate with group of strong retrolateral spines; metatar- sus I basally incrassate, distally arched; retrolaterally a low but distinct mound, set on retroventral comer, with strong spines almost forming a scoop; a long broad depression defined below by distinct keel from proventral comer proximally rising to upper junction with metatar- si. Leg II, viewed prolaterally, distally a long conical spur capped with short conical megaspine; two long upeurved spines on edge above spur. FIG. 1. Records of Namirea in north eastern Queensland; inset shows known range. 82 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Natnirea dougwallacei sp. nov., holotype 6 (QMS 19706). a, carapace & chelicerae, dorsal view; b, spinnerets, ventral view; c, eyes, dorsal view; d, chelicerae, sternum, maxillae & labium, ventral view; e, f, abdomen, dorsal (e), ventral view (0- Scale line =lmm (d); 2mm (a, b, e, f); 0.5mm (c). Etymology The specific epithet is a patronym in honour of Mr Doug Wallace, renowned arachnologist of Rockhampton, who prepared detailed and thorough arachnid displays for the Queensland Ambulance Service and Rockhampton Botanical Gardens, from which the types were ‘rescued’. Description Holotype male QMS 19706 Carapace 4.88 long, 4.00 wide. Abdomen 4.60 long, 3.20 wide. Total length, 10. Colour in alcohol. Carapace yellow brown with darker lines posteriorly on caput and beside anteromedial line, chelicerae, and legs yellow brown, none banded. Abdomen dorsally and ventrally dark brown without pattern. Carapace. Caput low. Pilosity: light cover of fine brown and silver hairs; striae broad, shallow. Bristles: one foveal pair; one very long and few finer on clypeal edge; all other setae slender. similar. Fovea a short, straight transverse pit. Clypeus ca. 0.16 from ALE. Leg 1 2 3 4 Palp Femur 3.63 3.44 3.44 4.19 2.19 Patella 2.19 2.19 1.88 2.25 1.50 Tibia 2.31 2.38 2.38 3.25 1.63 Metatarsus 2.56 2.81 3.56 4.50 Tarsus 1.19 1.56 1.88 2.06 0.88 Total 11.88 12.38 13.14 16.25 6.20 Table 1. Leg lengths of N. dougwallacei, holotype S . Leg 1 2 3 4 Palp Femur 3.28 3.32 3.24 4.00 2.40 Patella 2.12 2.12 2.00 2.28 1.44 Tibia 2.16 2.20 2.24 2.96 1.60 Metatarsus 2.32 2.48 3.08 4.00 — Tarsus 1.20 1.32 1.56 1.68 1.76 Total 11.08 11.44 12.12 14.92 7.20 Table 2. Leg lengths of N. dougwallacei, allotype $ . TWO NEW NAMIREA FROM NORTH QUEENSLAND 83 FIG. 3, Namirea dougwallacei sp. nov., holotype S (QMS19706). a-c, left tibia I, prolateral (a), ventral (b), retrolateral (c) views; d, tibia & metatarsus II, prolateral view; e, tibia, cymbium & palpal bulb, ventral view. Scale line =lmm. Eyes. Tubercle distinct. Group occupies 0.50 of head-width; front width, length, 38: 1 8. Front row procurved; back row recurved. MOQ front width, back width, length, 20:30:17. AME:ALE: PME:PLE, 10:11:9:10. Eye interspaces: AME- AME, 0.2; AME-ALE, 0.1; ALE-ALE, 2.0; PME-PLE, 0. 1 ; PME-PME, 1 .6; ALE-PLE 0. 1 . Chelicerae. Small; wide band of long erect brown bristles prodorsally. Intercheiiceral tumes- cence small, pallid, distinct. Furrow promargin with 5 large mixed with 8 smaller teeth, basomesally no teeth evident. Labium. 0.50 long, 0.88 wide. Anterior edge pallid, not indented; setae cover uniform, no pat- tern. Labiostemal suture a narrow continuous groove. Maxillae. 1.25 long in front, 1.18 long behind, 0.75 wide; anterior edge pallid. Heel truncated; anterior lobe well defined by long groove, lobe small; semila distinct. No modifications or strong setae on anterior face. Sternum. 2.50 long, 2.18 wide. Cordate, with strong ‘shoulders’ lateral of labium; setae on mar- gin not enlarged; covered with long erect brown bristles. All sigilla small, oval with long axis transverse, within one length of margin. Legs. I: viewed prolaterally, tibia subdistally incrassate with group of strong retrolateral spines evident; metatarsus basally incrassate, distally arched; retro laterally a low but distinct mound, set on retroventral comer, with few small and 1 1 long strong spines almost forming a scoop, as in Australothele; a long broad depression defined below by distinct keel from proventral corner proximally rising to upper junction with metatar- si. II: viewed prolaterally, distally a long conical spur capped with short conical megaspine; two long upeurved spines on edge above spur; metatarsus basally excavate, then a short incras- sate mound quickly reducing to normal leg diameter. Preening combs: I, III, none; II, 1 of 2 weak setae proventrally; 1 of 5 strong setae on retrolateral IV. No scopula. Spines. Distinction between ‘sj^ines’ and thick- ened bristles subtle. Leg 1, fe d7, pa pi, ti p2, v 5 short distal, 3 long slender plus 10 long megaspines in retroventral group and 1 proventrally, me v2; ta, 0. Leg 2, fe d6, pa p4, ti p3, v2 + 3 megaspines, me p2, v5; ta, v4. Leg 3, fe d6, rl, pa p5, d4, r2, ti p3, d3, r3, v6, me p4, r3, v5; ta, pi, v4; leg 4, fe d4, rl , pa p4, r2, ti p3, d2, r3, v6, me p5, r6, v8; ta, p4, rl, v8; palp, fe d5, pa pi, d2, ti pi, r2, la, 0. Claws. 8 long teeth in S-shaped line on paired claws of I, IV, unpaired claws bare. Trichobothria. Two rows, each of ca. 1 0 for 3/4 of tibiae; ca. 10-15 on metatarsi in line; ca. 10 filiform on tarsi. Palp. Bulb reniform, quickly tapering to long slender embolus extending past basal tibia; cym- bium short, indented, aspinose; tibia basally in- crassate with cluster of many long bristles, distally excavate. Spinnerets. PMS 1.25 long, 0.25 wide, 1.00 apart, and ca. 0.45 of basal PLS in diameter. Basal, middle, apical, total segments of PLS, 1.80, 1.88, 2.00, 5.68 long, respectively. Allotype female QMS 19707 Carapace 4.92 long, 4.08 wide. Abdomen 5.00 long, 3.60 wide. Total length, 11. 84 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Namirea dougwallacei sp. nov., allotype 9 (QMS19707). a, spinnerets, ventral view; b, carapace & chelicerae, dorsal view; c, chelicerae, sternum, maxillae & labium, ventral view; d, eyes, dorsal view; e, f, abdomen, dorsal (e), ventral view (0; g, spermathecae. Scale line =2mm (a, b, e, f); lnim (c); 0.5mm (d, g). Colour in alcohol. As in S . Carapace. Pilosity: as in S but 2 pairs of foveal bristles. Fovea short, transverse; set 45% of carapace length from back edge. Caput low but sloping down to fovea. Clypeus width, 0.12 to base of tubercle. Eyes. Tubercle distinct. Front row procurved; back row recurved. Group occupies 0.44 of head- width; front width, back width, length, 38, 38, 1 8, respectively. MOQ front width, back width, length, 20, 29, 15, respectively. AME:ALE: PME:PLE, 9:12: 7:10. Eye interspaces: AME- AME, 0.3; AME-ALE, 0.2; ALE-PLE, 0.2; PME- PME, 1.9; PME-PLE, 0.1; ALE-ALE, 2.0. Chelicerae. Geniculate, stout; setation like S . Furrow promargin with 4 large mixed evenly with 1 1 small teeth, basomesally with 4 small teeth. Labium. 0.55 long, 1.03 wide. Anterior edge indented. Labiostemal suture a narrow con- tinuous groove. Maxillae. 1 .28 long in front, 1.10 long behind, 0,83 wide. Heel truncated; anterior lobe short, with long defining groove. Sternum. 2.50 long, 2.20 wide. Cordate, widest TWO NEW NAMIREA FROM NORTH QUEENSLAND 85 at coxae I; all sigilla small, distinct, subcircular, marginal. Marginal setae like central. Legs. Preening combs: 1 of 3 strong setae on proventral I, II; 1 of 5 strong setae on retrolateral IV. Spines. Leg 1, fe d5w, pa p4w, ti p3. v5, me pi, v6, ta vl ; leg 2, fe d5w, pa p4w, ti p3, v5, me p3, v7, ta v3; leg 3, fe d3w, r2w, pa p3w, d2w, r3w, ti p3, d2, r3, v5, me p5, r3, v8, ta v3; leg 4, fe d5w, pa p3w, r2, ti p3, r3, v8, me p4, r3, v9; ta v3; fe d3, pa p2, dl, ti p2, v6, ta v7. Claws, ca.9 teeth in S-shaped row on paired claws of I, IV; unpaired claw with 1 small tooth; palpal claw with 10 teeth in straight line on medial keel. Trichobothria. As in <5 . Spermathecae. Two, each an inner long sclerotised spiralled duct with short straight un- sclerotised outer lobe. Epigastric furrow ex- tended in curve from outer edge of anterior booklung cover extending posteriorly to line at about half length of posterior booklung covers. Spinnerets. PMS 1.42 long, 0.32 wide, 1.20 apart, and ca. 0.50 of basal PLS in diameter. Basal, middle, apical, total segments of PLS, 2.25, 2.15, 2.27, 6.67 long, respectively. Distribution and Habitat Known only from vine thicket on the summit of Mt Archer, MEQ. Phytogeny The phylogenetic position of this species poses a dilemma. A phylogeny of some Australian Evagrini was implicit in Raven (1984). All Australothele males have a secondary spine on the spur of tibia I and distinct thorn on lower metatarsi 1. Males of all species, except A. rnagna, which lack spurs on tibia I, have clustered spines on tibia I. Hence, A. magna is the sister group of all other species of Australothele. Neither the secondary spine nor the metatarsal thorn are found in Namirea. In Namirea, the first leg lacks a spur but in N. insularis, a cluster of spines is present. Also, metatarsus and sometimes also tibia I have a dorsal flat area demarcated by a distinct ridge (Raven, 1984, figs. 201, 202). In N. dougwallacei, the form of the spur and spine cluster on tibia 1 resembles that in some Australothele. However, other aulapomoqjhies in Namirea override any notion that the spur similarity derives from recent common descent. Equally, the encumbent increase in homoplasies in both Australothele and Namirea imposed by interposing N. dougwallacei between A. magna and other Australothele species is too severe to be acceptable. Namirea johnlyonsi sp. nov. (Figs 1, 5, 6. Tables 3, 4) Material Examined Holotype d, Mt Spec National Park, 19°00’S 146°09’E, NEQ, on embankment at ecotone of rain- forest, Casuarina, and eucalypt forests, 3 Sep 1988, R. Raven, J. Gallon, T. Churchill, QMS 1 1225; Allotype 9 , 2 paralype 9 9 , same data as holotype, QMS 1 9704- Diagnosis Females differ from those of N. montislewisi in the narrow form of the spermathecae and males differ from those of N. dougwallacei in having a lower spur on tibia I. Small to medium-sized; preening combs present on legs I, II, IV. Two spermathecae, each a bipartite lobe broadest basally. Tibia I with a ventral mound bearing one long and several smaller megaspines; metatarsus I proximally in- crassate, slightly flattened dorsally. Tibia II dis- toventrally with long single spur tipped with small black megaspine. Palpal bulb long, pyriform with short embolus; cymbium with sclerotised distal edge. Etymology The specific epithet is a patronym in honour of Mr John Lyons, Chairperson, Management Com- mittee, Museum of Tropical Queenland, a branch of the Queensland Museum. Description Holotype male QMS 1 1225 Carapace 3.20 long, 2.92 wide. Abdomen 3.44 long, 2.08 wide. Total length, 7. Colour in alcohol. Carapace, chelicerae and legs yellow brown, ocular area black, carapace with light mottled brown margins and two long triangles laterally on caput; tibia I orange brown. Abdomen entirely dark brown. Table 3. Leg lengths of N. johnlyonsi, holotype 6 . Leg 2 3 4 Palp Femur 3.20 3.24 3.52 3.72 2.28 Patella 2.04 2.12 1.80 2.20 1.52 Tibia 2.36 2.28 2.08 3.12 1.68 Metatarsus 2.36 2.44 3.08 3.84 — Tarsus 1.16 1.28 1.52 1.56 1.68 Total 11.12 11.36 12.00 14.44 7.16 86 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Namirea johnlyonsi sp. nov,^ holotype 6 (QMSl 1225). a, eyes, dorsal view; b, carapace & chelicerae, dorsal view; c, spinnerets, ventral view; d, abdomen, ventral view; e. chelicerae, sternum, maxillae & labium, ventral view; f, tibia I, retrolateral view; g, tibia & metatarsus IT, prolateral view; h, palp showing tibia, cymbium & palpal bulb, prolateral view. Scale line =2mm (b-d); 1mm (a. e-h). Carapace. Pilosity: margins with line of fine brown setae; one pair of foveal bristles; light cover of fine gray hair on caput and interstrial ridges; striae distinct, glabrous. Caput low, flat. Fovea a small open pit. Clypeus absent. Eyes. Tubercle steep, distinct. Front row slight- ly procurved; back row recurved. Group occupies 0.49 of head-width; front width, back width, length, 34, 37, 20, respectively. MOQ front width, back width, length, 18, 26, 17, respective- ly. AME:ALE:PME:PLE, 10:13: 9:10. Eye inter- spaces: AME-AME, 0.3; AME-ALE, 0.2; ALE-PLE, 0.4; PME-PME, 1.6; PME-PLE, 0; ALE- ALE, 1.5. Chelicerae. Small, rounded, knee-like; with few long and many short curved bristles prodor- sally. Intercheliceral tumescence a small pallid area. Furrow promargin with 4 long mixed with 13 smaller teeth, basomesally with 5 teeth. Labium. 0.40 long, 0.76 wide. Labiostemal su- ture a shallow continuous groove. Maxillae. 0.98 long in front, 1 .00 long behind, 0.60 wide. Heel rounded; anterior lobe rounded, distinct, serrulate. Sternum. 1 .68 long, 1 .54 wide. Cordate with uniform cover of long erect bristles and short hairs. Sigilla six; all small, rounded. Legs. Tibia I distally incrassate; apically a ventral mound bears one long and several smaller megaspines; metatarsus I proximally incrassate, TWO NEW NAMIREA FROM NORTH QUEENSLAND 87 FIG. 6. Namirea johnlyonsi sp. nov., allotype 9 (QMS 19704). a, eyes, dorsal view; b, carapace & chelicerae, dorsal view; c, sternum, maxillae & labium, ventral view; d, abdomen, ventral view; e, spermathecae, dorsal view; f, spinnerets, ventral view. Scale line =0.5mm (a, f); 2mm (b, c); 4mm (d); 0.25nim(e). slightly flattened dorsally. Tibia II distoventrally with long single spur tipped with small black megaspine, passing prolateral when metatarsus II is contracted; metatarsus II proximally excavate distal of which is incrassate area. Preening comb of 4 setae on retroventral IV. Scopula absent but sparse band of erect hairs ventrally on all tarsi. Spines. Leg l,fed5,pa 0, tiplw, vll.me 0, ta, 0; leg 2, fe d5w, pa p3w, ti p3w, v2 + megaspine, me p2, v5 ta, 0; leg 3, fe d5 w, pa p4w, r2w, ti p3w, d2w, r3w, v6, me p3, r3, v5, ta, v2; leg 4, fe d6w, pa p2w, r2w, ti p3, r^ v5, me p3, r3, v5; palp, fe d5w, rest, 0. Claws. One S-shaped row of 9 (I) to 7(IV) teeth on paired claws; unpaired claw bare. Trichobothria. Two rows, each of 10 for half length of tibiae; ca. 10 on metatarsi in straight line; ca 8 filiform in line on tarsi. Palp. Bulb long pyriform with short embolus; cymbium with sclerotised distal edge. Spinnerets. Australotheline crescent distinct. PMS 0.72 long, 0.16 wide, 0.76 apart, and about 0.50 of basal PLS in diameter. Basal, middle, apical, total segments of PLS, 1.64, 1.60, 1.80, 5.04 long, respectively. Allotype female QMS 19704 Carapace 4.92 long, 4.00 wide. Abdomen 6.48 long, 4.80 wide. Total length, 13. Colour in alcohol. Carapace and legs light orange brown, caput, lateral margins and inter- strial ridges of carapace slightly darker. Abdomen brown. Carapace. Oval; with uniform cover of fine gray hairs; lateral margins with few longer hairs; one pair of foveal bristles arising deep in fovea. Fovea a small pit with steep front edge forming recurved arc. Striae distinct, glabrous. A line of 4 erect setae on clypeal edge; clypeus narrow but distinct; few hairs between ALE and PME. Eyes. Front row procurved; back row recurved. Group occupies 0.45 of head-width; front width, back width, length, 48, 52, 28, respectively. MOQ 88 MEMOIRS OF THE QUEENSLAND MUSEUM front width, back width, length, 22, 35, 20, respectively. AME:ALE:PME:PLE, 9; 15: 11: 12. Eye interspaces: AME-AME, 0.6; AME-ALE, 0.4; ALE-PLE, 0.6; PME-PME, 2.6; PME-PLE, 0.1; ALE-ALE, 2.7. Chelicerae. Stout, porrect with ca 20 long black and many shorter setae dorsally. Fangs short. Furrow promargin with 3 large mixed with 13 smaller teeth, basomesally with 7 teeth and 20-30 granules. Labium. Like cJ, 0.48 long, 1.12 wide. Maxillae. Rectangular; anterior lobe distinct but with delimiting groove; 1.48 long in front, 1.48 long behind, 0.92 wide; heel indistinct. Sternum. Like cJ; 2.40 long, 2.24 wide. Sigilla all oval, marginal but anterior pair farthest from margin. Legs. Covered with long curved setae and fine hair. Scopula absent. Preening combs: 1 of 3 setae on prolateral I, II, none of III; 1 of 4 setae retroventrally on IV. Spines. Thicker setae scored as weak spines on femora. Leg 1, fe p6w, pa p3w, ti p3, v5, me pi, v5; la, 0; leg 2, fe d5w, pa plw, li p3, v5, me p3, v5, ta, v2; leg 3, fe d5w, pa p3w, r3, ti p3, r3, v5, me p3, r3, v7, ta, v2; leg 4, fe d5w, pa p3, r2, ti p3, d4w, r3, v6, me p3, r3, v7, ta, dl, v3; palp, fe d5w, pa 2w, ti p2,w, v7, ta v9. Claws. S-shaped line of 1 5 (I) to 9(IV) teeth on paired claws; 3 on unpaired claws; palpal claw with 12 teeth in one medial line. Trichobothria. Two rows, each of ca. 10 for half length of tibiae; ca. 18 on metatarsi; ca. 8 on tarsi. Spermathecae. Two, each a bipartite lobe broadest basally. Spinnerets. PMS 1.48 long, 0.32 wide, 2.08 apart, and about 0.53 of basal PLS in diameter. Basal, middle, apical, total segments of PLS, 2.56, 2.08, 2.08, 6.72 long, respectively. Distribution and Habitat Known only from Mt Spec NP, NEQ, in a mixed forest ecotone including Casuarina, eucalypts, and rainforest. The species was not found at higher altitudes in an evidently highly disturbed rainforest where ground fauna was generally depauperate. Nor was it evident in less Table 4. Leg lengths of M johnlyonsi, allotype 9 Leg 2 3 4 Palp Femur 2.64 2.40 2.60 3.08 1.68 Patella 1.44 1.60 1.36 1.36 0.96 Tibia 1.92 1.92 1.96 2.56 1.36 Metatarsus 1.92 2.12 2.72 3.60 Tarsus 1.04 1.32 1.40 1.80 0.72 disturbed adjacent areas of rainforest. Clearly, the preference of this group for embankments indi- cates that it will take advantage of the microhabitats provided by road cuttings. (A similar phenomenom was noted by Raven, 1991 in New Caledonian diplurids.) Like many mygalomorphs in Australia, this group is more diverse in open forest habitats. Web The web had bluish white curtains of silk ex- tending 8-10 cms from the overhang of an em- bankment beside a road. Several corridors lead to a branching main tube extended only 3-4cm into the soil. ACKNOWLEDGEMENTS Illustrations were skillfully draw by Clare Bremner (2-4), and Bronwyn Mitchell (5, 6). This paper was produced with the support of Australian Research Council grants (A 1 87 1 5 1 05, AD9031859). LITERATURE CITED RAVEN, R.J. 1 984. The Australian curtain-web spiders (Ischnothelinae: Dipluridae: Chelicerata). Australian Journal of Zoology. Supplementary Series 93: 1-102. 1985. The spider infraorder Mygalomorphae (Araneae): cladistics and syslematics. Bulletin of the American Museum of Natural History 182: 1-180. 1991. A revision of the mygalomorph family Dipluridae (Araneae) in New Caledonia. In: Chazeau, J. & Tillier, S. (eds), Zoologica. Neocaledonica 2. Memoires de la Museum na- tionale d’Histoire naturelle, A, 149: 87-117. FUNCTIONAL SIGNinCANCE OF NEST CONSTRUCTION BY AN AUSTRALIAN RAINFOREST FROG: A PRELIMINARY ANALYSIS STEPHEN J. RICHARDS Richards, SJ. 1993 12 24: Functional significance of nest construction by an Australian rainforest frog: a preliminary analysis. Memoirs of the Queensland Museum 34(l):89-93 Brisbane. ISSN 0079-8835. A frog of the Litoria lesueuri complex from northeastern Queensland rainforests sometimes deposits eggs in small basins constructed on sandy banks adjacent to streams. Observations and experiments on nests at Elphinstone Ck showed that basin construction commenced by early September, and that the mortality of embryos from desiccation was extremely high. Embryos removed from basins and placed in the stream suffered no delectable predation, ^d developmental rates of embiyos placed in streams were similar to those left undisturbed in basins. Basin construction may have evolved as a mechanism to secure egg masses to a homogeneous and mobile substrate. Hylidae, aquatic nest, embryo, development, predation, Litoria lesueuri complex. S.J.Richards, Zoology Department, James Cook University, Townsville, Queensland 481 1, Australia; 20 August, 1993. A few species of frogs that breed along rain- forest streams construct small water-filled basins (‘nests’) for egg deposition (Duellman & Trueb, 1986). Two hypotheses have been proposed to explain the evolution of this behaviour. It evolved either to exclude potential predators of eggs and embryos, or to t^e advantage of higher water temperatures to ensure more rapid embryonic development (Crump, 1974; Lammote & Les- cure, 1977). There have been few attempts to test these hypotheses, and two recent studies (Caldwell, 1992; Kluge, 1981) reached different conclusions about the functional significance of nests. Richards & Alford (1992) reported the first example of this reproductive strategy in Australia and speculated on the benefits of nest construc- tion. The si>ecies involved here is undescribed but is referred to as Litoria lesueuri (Richards & Alford, 1992). I report the results of preliminary experiments designed to lest the proposed hypotheses, and propose an additional one to explain the evolution of nest construction in this species. MATERIALS AND METHODS I made a series of observations on 12 nests over a 15 day period in September, 1991 at El- phinstone Ck, a lowland stream west of Ingham, NEQ (18° 29’S, 146° Ol’E). Each nest was marked with a flag for identification, and infor- mation on mortality of eggs and tadpoles was recorded. Each nest was examined for the presence of potential predators. Nests were clas- sified as ‘separate’ if there was no connection between the stream and the nest, and ‘connected’ if there was even a small channel between the nest and the stream. Separate nesLs therefore included those separated from the water only by the nest wall, and some located more than one metre from the stream edge. Only nests separate from the stream were used in the experiments reported below. To test the hypothesis that water temperatures are higher in nests than in the adjacent stream, and confer an advantage in terms of more rapid embryonic development, I placed a max-min thermometer in each of two nests, and another in a small basin artificially constructed in the stream adjacent to them. Nests and artificial basins were about 5 cm deep, and in each case the ther- mometer was about 3 cm below the water surface. Approximately 50 eggs were excised from the egg mass in each nest and placed in the artificial basins in the adjacent stream. Maximum, mini- mum and ambient temperatures were recorded after three days when hatching had commenced. These temperatures were recorded, and the embryos preserved, within a 15 minute period. Between 1400 - 1500 on the afternoon of 6.9.91 ambient temperatures in the 12 nests and adjacent stream were recorded. Temperatures in each nest/stream combination were recorded within 30 seconds. Dry nests often contained no desiccated tad- poles so, to determine whether tadpoles could escape drying nests, I constructed a moat (about 90 MEMOIRS OF THE QUEENSLAND MUSEUM 10 cm deep) around each of four nests that con- tained embryos and were separate from the stream. As the nests dried the moats (which retained water) were checked for the presence of tadpoles leaving the nests. Water levels in nests were regulated by seepage from the stream so moat construction did not contribute to drying of the nests. The staging tables of Gosner (1960) were employed. RESULTS Male L. lesueuri (Fig. 1) constructed small circular basins (Fig. 2) in sand banks along El- phinstone Ck (Fig. 3), between September and December, 1991. At the time of the study the area was extremely dry and water levels were falling. No nests were observed after late summer storms flooded the creek in March 1992. Egg masses were never observed away from nests, and in nearby rocky creeks frogs attempted to construct nests amongst coarse pebbles (Fig. 4). Half of the nests observed initially contained a single egg mass; the remainder contained recently hatched, free-swimming tadpoles. Eight of the 12 nests were separate from the stream, two were sub- merged completely, and two (8 & 12) were con- nected to the stream by shallow channels. Mortality Observations on survivorship of embryos in 12 nests over a 15 day period (Table 1) show mor- tality was extremely high due to desiccation as water levels dropped. Free-swimming tadpoles escaped from only two of the 12 nests (17%) (Table 1). Both of these nests were submerged, and the tadpoles escaped before the nests were separated from the creek by falling water levels. Tadpoles in another nest that was not studied intensively were observed to swim from the nest FIG. 1. Male nest-building frog L. lesueuri . via a shallow channel, and subsequently return to the nest. As water levels fell these tadpoles died from desiccation. Moats dug around four nests contained water but no tadpoles when the nests dried up. Three of these nests contained desiccated tadpoles, but one was empty. Ants were observed in dried nests, FIG. 2. Aquatic nest with a single egg mass at El- phinstone Creek. FIG. 3. The main study site, Elphinstone Ck. Nests were located in the sand banks at the water's edge. RG. 4. Nest constructed in a rocky creek near the study site. FUNCTIONAL SIGNIHCANCE OF FROG NESTS 91 TABLE 1. Mortality of embryos and tadpoles in 12 nests at Elphinstone Ck, NEQ, S = nest separate from stream, C = nest connected to stream; T=tad- poles; St = tadpole stages according to Gosner ( 1 960); E = unhatched embryos, tadpoles at stage 24 and above are free-swimming; dry = 100% mor- tality. Where a stage is not given, the tadpoles have hatched, but data on stage are not available. Date Nest 6.9.91 9.9.91 12.9.91 20.9.91 I S.T dry 2 S,E S,St 24 S,St25 dry 3 S T 95% dead S,St 22 dry — 4 S,T dry — 5 S,T S,T, almost dry dry — 6 C.E C,T escaping — — 7 C.T C, St 25 escaping — — 8 C,E S,T dry 9 S,E hatching S,St 24 S,St 25 dry 10 S,T S.Sl 24 S,St25 dry 11 S,E S,St 20 S, St 24 dry 12 C,E S,T S,T dry and egrets and herons were observed along the stream bank. They may have removed tadpoles from some nests. Development and Predation Table 2 summarises information on the temperature regime and tadpole development in two nests and the adjacent stream. Nest 2 became much warmer than nest 1 , and the adjacent stream at this site was also warmer than the stream ad- jacent to nest I . Both nests were warmer than the adjacent stream during the day, but were slightly colder at night. On 6.9.91 four shaded nests were 1 -5°C cooler than the adjacent stream, and four unshaded nests were 4.2 - 7.7°C warmer than the adjacent stream. Four connected nests were 4 - 5.7°C warmer than the adjacent stream. There was no difference between developmen- tal rates of embryos in nests and in the adjacent artificial basins in the stream over three days at two sites (Table 2). Differences in developmental stages of embryos between sites partly reflected different initial stages of embryos at the two sites, but embryos at the warmer site (nest 2) also appeared to develop faster than those at the cooler site (Table 2). There were large numbers of fish and con- specific tadpoles in the creek, and large (stage 36-7) conspecific tadpoles were seen in two sub- merged nests (6 & 7, Table 1) that contained smaller embryos. If egg predation occurred it was minimal. All egg masses, including those in sub- merged nests, remained intact throughout the study. However it was impossible to determine whether small numbers of eggs were consumed from the surface of the egg masses. Embryos translocated from two nests into the stream suc- cessfully hatched at both sites. DISCUSSION Mortality of embryos deposited in nests during this study was extremely high. It was attributed primarily to desiccation, although predation may occur as water levels in nests drop. As water levels in the nests are maintained by seepage (Richards & Alford, 1992) falling water levels result in rapidly drying nests. Most nests were separate from the creek, and as water levels dropped their isolation from the stream increased. Isolated nests are unlikely to release tadpoles to TABLE 2. Temperature regimes in two nests and ad- jacent stream, with stages (Gosner, I%0) of tadpoles transferred to .stream and left in nests, 9.9.91 and 12.9.91. Differences between nests reflect different initial stages of embryos in each nest (nest 1 = stage 19, nest 2 = stage 9). All ambient temperatures taken within fifteen minutes at time of data collection. NEST 1 NEST 2 Nest Stream Nest Stream Temp °C Temp °C Max: 30.0 27.5 34.0 31.0 Min: 14.5 17.5 15.5 17.0 Amb: 30.0 25.0 23.5 26.0 Tadpole stages (Gosner, 1960) (n = 10) 22 23 20 20 23 23 20 20 23 23 20 20 23 23 20 20 23 23 19 20 23 23 20 20 23 23 20 20 23 23 19 20 23 23 20 20 23 23 20 20 92 MEMOIRS OF THE QUEENSLAND MUSEUM the stream, as tadpoles rely on erosion of the nest wall (presumably from fluctuating water levels) to escape. Tadpoles of the Hispaniolan Frog, Hyla vasta were reported by Noble (1927) to wriggle over wet rocks from rocky basins into an adjacent stream, but tadpoles of L, lesueuri were unable to escape drying nests. All nests were constructed at the waters edge, (or were submerged) minimising the probability of nests becoming isolated from the stream. Nests reached much higher temperatures than the adjacent stream during the day, but were cooler at night and appeared to cool more rapidly than the stream when shaded. Nests constructed along rainforest streams may be in shade for most of the day, and thus may be cooler than the adjacent stream much of the time, so explaining the similarity between developmental rates of embryos in and out of nests. Given the apparent spatial heterogeneity of temperature regimes ob- served at the two nests manipulated during this study, further work is required to determine whether frogs select sunny positions for nest sites rather than shady positions, and whether in* creased temperature may enhance development. Although there were large numbers of fish and conspecific tadpoles in the stream, tadpoles from both submerged nests observed escaped into the stream. Embryos translocated from nests into the stream also hatched with no apparent mortality from predation. My observations differ from those of the most comprehensive study to date on the biology of a nest-building frog: Hyla rosenbergi from Panama (Kluge, 1981), where there was little difference between nest and stream temperatures. Nest temperatures were, however, more variable. H. rosenbergi embryos were consumed rapidly by a number of predators if they were removed from nests and placed in the adjacent stream. Kluge (1981) argued that predation of embryos by small fish, and hetero-and conspecific tadpoles played a key role in the evolution of nest construction in that species. Although mortality from desiccation was considered insignificant, he suggested that it may be higher in the drier, early wet season. Caldwell ( 1 992) found that nests of Hyla boans in Brazil were warmer than those in the stream, and showed that embryos developed more rapidly in nests. The temperature regime was variable among nests and she found that nests with seepage from the stream were cooler and had embryos with slower development than isolated nests. Despite this apparent advantage, Caldwell (1992) also found that embryos in nests of H. boans suffered extremely high levels of mortality from predation and desiccation. Only two of seven nests observed by Caldwell (1992) produced any surviving tadpoles. In contrast, embryos from all clutches laid by //. boans out- side nests escaped into the stream (Caldwell, 1992). That study was also conducted during the dry season when water levels were falling. The functional significance of nest construc- tion appears to vary between localities and species. Both of the hypotheses proposed to ex- plain the advantages of nest construction gain some support from the studies of Kluge (1981) and Caldwell (1992). However the selective ad- vantage of nest construction by L. lesueuri is less clear, and additional studies are required to fur- ther test the two hypotheses. Further experiments should be conducted to determine predation rates on egg masses by fish and conspecifics, and variability in temperature regimes in nests at many sites also needs to be established. My preliminary results suggest that predation of embryos by aquatic organisms may not be a major cause of mortality. Increased temperatures may not provide benefits in terms of enhanced development because nests may be cooler than the stream for most of the day. Mortality in nests later in the wet season needs to be assessed but observations during 1991-2 suggested that nest construction during monsoon rains may be an ineffective strategy as creeks flood and nests and tadpoles are washed away. What then, is the functional significance of this strategy? I have observed many submerged nests in streams in northern Queensland. The sandy substrates at many of these sites are uniform and mobile. Egg masses removed from nests and placed in the current are frequently rapidly washed downstream. The evolution of nest construction by this species initially may have been a response to a substrate on which egg deposition was virtually impossible without a depression in which to protect the clutch from the force of the current. The observed benefits of nest construction by several species may in fact be a consequence, rather than the cause of, the evolu- tion of nest construction. A large proportion of nests constructed by H. boans in central Amaz- onia are connected to the stream (Hero, pers. comm.), and the nests oi'Rana blythi from Borneo are completely submerged in the stream (Emer- son, 1992). FUNCTIONAL SIGNIRCANCE OF FROG NESTS 93 ACKNOWLEDGMENTS Field work was supported by ARC grants A 187 15248 and 18931875 to R.A. Alford. L. Schwarzkopf. M. Crossland, and Amanda, Cate and Nicolas Richards assisted in the field. R. Alford encouraged the study, and J-M. Hero, MJ. Tyler and M. Mahony kindly commented on the manuscript. LITERATURE CITED CALDWELL, J.P. 1992. Diversity of reproductive modes in anurans: facultative nest construction in gladiator frogs. Pp. 85-97 In Hamlett, W.C. (ed). ‘Reproductive diversity of South American vertebrates*. (Springer- Verlag: New York). CRUMP, M.L. 1974. Reproductive strategies in a tropical anuran community. University of Kansas Museum of Natural Histoiy Miscellaneous Pub- lications 61: 1-68. DUELLMAN, W.E. & TRUER, L. 1986. ‘Biology of amphibians’. (McGraw Hill: New York). EMERSON, S.B. 1992. Courtship and nest-building behavior of a Bornean frog, Rana blythi. Copeia (4):1 123-1 127. GOSNER. K.L. 1 960. A simplified table for staging anuran embryos and larvae with notes on iden- tification. Herpetologica 16: 183-190. KLUGE, A.G. 1981. The life history, social organiza- tion, and parental behavior of Hyla rosenbergi Boulenger, a nest-building gladiator frog. Mis- cellaneous Publications of the Museum of Zool- ogy. University of Michigan. 160: 1-170 LAMMOTTE, M. & LESCURE, J. 1977. Tendences adaptives a I’affranchissement du milieu aquati- que chez les amphibiens anoures. Terre Vie 31: 225-311. NOBLE, G.K. 1927. Ihe value of life history data in the study of the evolution of the Amphibia. An- nals of the New York Academy of Science 30: 31-128. RICHARDS, S.J. & ALFORD, R.A. 1992. Nest con- struction by an Australian rainforest frog of the Litoria lesueuri complex (Anura: Hylidae). Copeia (1992): 1120-1123. 94 MEMOIRS OF THE QUEENSLAND MUSEUM RECOGNITION OF LITORIA EVCNEMIS (LONNBERG) in AUSTRALIA. Memoirs of the Queensland Museum 34(1): 94. 1993:- In 1979, in the Mc- Ilwraith Ra., Cape York Peninsula (CYP), one of us (KMcD) recorded the call of a frog in the Litoria eucnemis species- group (Tyler & Watson, 1985). The call, a series of short growls, has been confirmed recently (September, 1993, KMcD). It clarifies nomenclature of Australian taxa in the L eucnemis group. Species of this group occur in New Guinea as well as Australia and three names have been proposed for them — Hyla eucnemis Lonnberg, 1900 (described from Sattelberg, Huon Peninsula, PNG); H. genimaculata Horst, 1883 (‘Gebeh* Is, west of Waigeo Is., West Irian); and H. serrata Andersson, 1916 (Malanda, Atherton and Carrington, NEQ). Australian 'eucnemis' occur in three discrete populations: southern (Paluma-Big Tableland, 19°0rS 146°12’E-I5®42’ 145^16’, NEQ); middle (Mcllwraith Ra.-Iron Ra.. 13°50’S 143°17’E-12®46* 143‘^16’, CYP); and northern (Wenlock R.- Ducie R.. 12°16’S I4r59’E-12“07’ 142"2r, CYP). These populations have always been treated as a single taxon, but the names Litoria serratOy L. genimaculata and L eucnemis have variously been applied (c.g. Ingram & Covacevich, 1981; Cogger et al, 1983; Tyler & Watson, 1985). There are, however, at least two taxa in Australia with distinctly different mating calls. Males of the southern popula- tion produce a series of soft ‘ticks’ (SIR, KMcD, GJI, pers. obs.; Fig. 1). Those of the middle population have a series of short growls (Tyler & Watson. 1985). Calls of the northern population have not been recorded. In New Guinea there are also two taxa with distinct calls: one resembling a short, repeated ‘growl’ (described as ‘waa waa' by Menzies (1976)andas ‘groups of low, soft chuckling notes’ followed by a ‘series with several louder, shorter, pulsed calls’ by Zweifel (1980)); the other a series of soft ‘ticks’. Both authors also concluded there were two mor- phologically similar species distinguished primarily by mating call structure, and by slight differences in degree of finger webbing and size. They assigned the name SECONDS Fig. 1 . Sound spectrograph of call of L. genimaculata recorded 23 January, 1977, Tully Falls, NEQ, by G.J. Ingram. Fig 2. L. eucnemis, Mcllwraith Ra., September, 1993. genimaculata to the taxon with the ‘soft tick’; and eucnemis | to the other. j Calls of the two Australian taxa closely resemble those of ' the two New Guinean species. We conclude that the ap- propriate name for the population between Paluma-Big Tableland is Litoria genimaculata; and that for the Iron Ra. -Mcllwraith Ra. taxon is Litoria eucnemis (Fig. 2). Thus H. serrata Andersson, 1916, becomes a junior subjective j synonym of L. genimaculata (for photograph, see Cogger, | 1992: 139). Further, pending investigation of the call of the Ducie R.-Wenlock R. population, it seems reasonable to as- sign that poiulation to L eucnemis. Acknowledgements I R.G. Atherton, J.W. Winter and D. L. Storch assisted with field work in the Mcllwraith Ra. in 1979 and 1993. Research on rainforest frogs was funded by the Wet Tropics Manage- ment Authority, Queensland Department of Environment and Heritage, Australian Nature Conservation Agency, Queensland Museum and ARC grants A18715284 and A 1893 1875 to R.A. Alford. Literature Cited Cogger, H.G. 1992. ‘Reptiles and amphibians of Australia.’ 5th ed. (Reed Books: Frenchs Forest). Cogger, H.G., Cameron, E.E. & Cogger, H.M. 1 983. Amphibia and Reptilia. Zoological Catalogue of Australia 1:1-31 3. Ingram, G.J. & Covacevich. J. 1981. Frog and reptile type specimens in the Queeasland Museum, with a checklist of frogs and reptiles for Queensland. Memoirs of the Queensland Museum 20: 291-306. Menzies, J.I. 1976. ‘Handbookof common New Guinea frogs’ (Wau Ecology Institute: Wau). Tyler, M.J. & Watson, G.F. 1985. On the nomenclature of a hylid tree frog from Queensland. Transactions of the Royal Society of South Australia 1 10: 193-194. Zweifel, R.G. 1980. Results of the Archbold Expeditions. No. 103. Frogs and lizards from the Huon Peninsula, Papua New Guinea. Bulletin of the American Museum Natural History 165; 387-434. SJ. Richards, Zoology Department, James Cook University 4811: K.R. McDonald, Queensland Department of Environ- ment and Heritage, PO Box 834, Atherton 4883; G.J. Ingram, Queensland Museum, PO Box 3300, South Brisbane 4101. A REVIEW OF THE LEAF-TAILED GECKOS ENDEMIC TO EASTERN AUSTRALIA- A NEW GENUS, FOUR NEW SPECIES, AND OTHER NEW DATA P.J. COUPER, J. A. COVACEVICH AND CRAIG MORITZ Couper, P.J., Covacevich, J.A. & Moritz, C. 1993 12 24: A review of the leaf-tailed geckos endemic to eastern Australia: a new genus, four new species, and other new data. Memoirs of the Queensland Museum 34(1):95-124. Brisbane. ISSN 0079-8835. Saltuarius gen. nov. is separated from Phyllurus by external and internal characters, and by karyotype. To it are assigned S. comutus (Ogilby, 1892); S. salebrosus (Covacevich, 1975); 5. swaini (Wells & Wellington, 1985) and S. occultus sp.nov. To Phyllurus sensu stricto are assigned P. caudiannulatus Covacevich, 1975; P. platurus (Shaw, 1 790); P. isis ; P. nepthys and P. ossa spp. nov. The leaf-tailed geckos (Saltuarius spp. and Phyllurus spp.) are confined to narrow, isolated patches of rainforest or heathlands in coastal Australia between the Mcllwraith Range, far northeastern Queensland (13''45’S, 143°19’E) and the Hawkesbury Sandstone area, near Sydney (33°55’S,15]°13*E), mideastem New South Wales. Endemic species lists for several rainforests and lists of rare, endangered or vulnerable species have been changed by this revision. P.occultus is recognised as the first reptile species endemic to rainforests of the Mcllwraith Ra., FNEQ; S. comutus is no longer a wide-ranging species, but is confined to the Wet Tropics, NEQ; and P. isis, P. nepthys, P. ossa spp. nov. and P. caudiannulatus join lists of very narrowly endemic rainforest species. □ Saltuarius, Phyl- lurus , rainforest, endemism, Reptilia, Squamata,Gekkonidae, Australia. P. J. Couper &J. A. Covacevich, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101; C. Moritz, Zoology Department, University of Queensland, Queensland 4072, AustraHa,4 September 1993. Leaf-tailed geckos are well known as being strongly associated with tropical and subtropical rainforests and, to a lesser extent, heaths on sandstones and granites in eastern Australia. Covacevich, 1975 completed a revision of Phyl- lurus, recognising Phyllurus platurus (Shaw, 1790), P. caudiannulatus Covacevich, 1975 (both small species); P. comutus Ogilby, 1892, and P. salebrosus Covacevich, 1975 (both large species). Since then, additional work has been undertaken on Phyllurus species, notably by Wells & Wellington (1985) and by Bauer (1990). Further, many new specimens of Phyllurus have been added to the reference holdings of the Queensland and Australian Museums. Wells & Wellington (1985) separated P. swaini from P. comutus, recognising the specific status of southern and northern forms, previously treated as one species Covacevich (1975). Bauer (1990) confirmed the monophyly of Phyllurus, clarified relationships with other carphodactyline geckos, and provided new data on two distinct subgroups wilhan Phyllurus, a new key, and new species diagnoses. He did not recognise P. swaini. Rainforest vertebrates in Australia are now generally regarded as being fairly well known, at least in taxonomic terms. However, rainforests of mideastem Queensland are not as well known as those further north. In an attempt to redress that situation, the National Rainforest Conservation Programme funded research in unsurveyed rain- forests in this area. Two of the new species of Phyllurus from isolated rainforests of mideastem Queensland were collected during this field work. T^eir discovery prompted this review. Some 6 1 specimens of Phyllurus have been added to the Queensland Museum’s holdings since Covacevich (1975). Most significant are specimens of a new species from far northern Queensland, three new species from isolated rainforest blocks in mideastern Queensland, and specimens from southern Queensland and New South Wales. The last nientioned confirm recognition of P. swaini Wells & Wellington (1985) and provide new dis- tributional records and other data. Bauer’s (1990) definition of two subgroups within Phyllurus prompted investigation of pos- sible separate generic status for the large species of Phyllurus. In particular, we examined the dis- tribution of the diagnostic characters defined by Bauer in our new specimens and added new in- formation from karyology. As a result, we define a new genus, Saltuarius, including four species of large leaf-tails-^, comutus (Ogilby 1892), 5. occultus sp. nov., S. salebrosus (Covacevich, 1975) and 5. swaini (Wells & Wellington, 1985). To Phyllurus sensu stricto we assign two pre- viously recognised species - P. caudiannulatus 96 MEMOIRS OF THE QUEENSLAND MUSEUM Covacevich, 1975 and (Shaw, 1790) and three new species - P. isis, P. nepthys and P. ossa from the rainforests of mideastem Queensland. Body measurements follow Covacevich (1975) for snout to vent length (SVL); tail length (T), from posterior margin of cloaca to tip of tail; attenuated tip of original tail (TT); head length (HL); head width (HW); snout length (S), Addi- tional measurements include - length of front leg (LI) axilla to tip of longest digit; length of hind leg (L2) groin to tip of longest digit; neck length (NL) axilla to posterior margin of ear. Morphological characters follow Covacevich (1975) also, with the following modifications - labial scale counts are for both sides of each specimen of newly described species; subdigital lamellae of the 4th toe are for both sides of each specimen (in a sample of 20, ‘n’ for labial and toe lamellae counts could be a maximum of 40). Skeletal definitions follow Bauer, (1990). Colours are defined subjectively, e.g. cream, rather than according to Ridgway (1912), e.g. cartridge buff. Axillary pits (acarodomatia) are described for both new and previously known species, wherever they are present. As it is not universally agreed that they are useful taxonomic tools (Loveridge, 1951; Arnold, 1986), we use them with caution, only as secondary features, in support of our species definitions. GENERIC HISTORY The genus Phyllurus has had a fairly stable taxonomic history, save for the successive in- clusion in, and exclusion from it of Under- woodisaurus spp. Vnderwoodisaurus has been treated as a synonym of Phyllurus by Kluge (1967) and Russell (1980), and as distinct from Phyllurus by Covacevich (1975) and Bauer (1990). Bauer (1990) provided a 107 character matrix for all carphodactyline taxa and con- structed a consensus cladogram of the Australian padless carphodactyline genera from it. His node 5 defined a monophyletic group including Car- phodactylus laevis with all the then known Phyl- lurus spp. Further, he identified five characters, three of them unique synapomorphies, that con- firmed the monophyly of Phyllurus. In PhylluruSy he recognised two diagnosable subgroups (nodes 7,8). In the former, he placed P. caudiannulatus and P. platurus. To the latter he referred P. cor- nutus and P. salebrosus. This separation of what are essentially the small from the large Phyllurus sensu lato, foreshadows the recognition here of a new genus, Saltuarius, for the large geckos formerly assigned to Phyllurus. We have examined Bauer’s characters for the two phylogenetic divisions in leaf tail geckos (Phyllurus sensu lato) in our expanded collection, added new characters and reassessed the evidence using parsimony analysis. One important charac- ter, the expansion of the epipubic cartilage, was found to exhibit more variation than recognised by Bauer (1990), with one species of each puta- tive group (S. occultus gen, et sp. nov. and P. nepthys sp. nov.) having an intermediate condi- tion (Fig, 1). Another significant character used by Bauer was the presence or absence of an anterior process of the interclavicle which, if present, was a narrow spike vs a broad process. Again, we recognize an intermediate state, in P. nepthys (Fig. 2). Important evidence derives from preliminary analysis of karyotypes. Previous studies of Phyl- lurus and related genera of carphodactyline geckos (Nephurus and Carphodactylus) have reported variation in the number of chromosome arms, but not in the number of chromosomes, all species having the presumed ancestral chromosome number of 2n = 38 (King, 1 987). We have obtained new data confirming this result for S. cornutus and showing that other species of Saltaurius (S. swaini and S. salebrosus) also have the ancestral condition. Species within Phyllurus are unique among carphodactylines in having reduced chromosome numbers. The mideastern Queensland species (P. caudiannulatus, P. nep- thys and P. ossa) have 2n = 28 or 2n = 30 chromo- A B C D E F G H FIG. 1 . Variation in si7x and shape of epipubic cartilage in Saltuarius spp. and Phyllurus spp. (A) S. cornutus - J48178; (B) S. occultus - J37037; (C) S. salebrosus J33700; (D) S. swaini - J8075; (E) P. caudiannulatus - J3363 1 ; (F) P. tsis - J535 1 2 (G) P. nepthys - J34024; (H) P. ossa - J53391. See Bauer (1990) for condition in P. platurus. REVIEW OF LEAF-TAILED GECKOS 97 T T T ▼ ▼ T ▼ FIG. 2. Condition of the anterior inlerclavicular exten- sion in Saltuarius spp. and Phyllurus spp, (A) S. comutus - J48178; (B) S, occultus - J37037: (C) S. salebrosus - J33700; (D) S. swaini - J8075; (E) P. caudiannulatus - J33631; (F^ P. isis - J53512; (G) P. nepthys - J34024; (^) P. ossa - J53391. See Bauer (1990) for the condition in P. platurus. somes (Fig. 3) with predominantly metacentric or submetacentric chromosomes, suggesting reduc- tion from the 2n = 38 condition via centric fusion. P . platurus has an even more reduced chromosome number, 2n = 22. Such dramatic reductions in chromosome number have been reported previously within genera (even ‘species’ of geckos, e.g. Nactus, Moritz, 1987; Diplodac- tylus. King, 1987). Our analysis is limited by small sample size (1-2 specimens per species) and the absence of data for two species, S. occultus and P. isis. Nonetheless, it provides strong sup- port for the grouping of the species Phyllurus. The chromosome evidence, along with a series of morphological characters (Table 1) were used in a parsimony analysis (using PAUP 3.0s with ordered stales and C. laevis as an outgroup) to re-evaluate the phylogenetic division proposed within Phyllurus sensu lato (Bauer, 1990). Three shortest trees were obtained with the branch and bound search option, and each supported the monophyly of Saltaurius and Phyllurus (Fig. 4). These major clades were each strongly supported in a bootstrap analysis present in 98% of pseudo- replicalcs, the former being defined by a mini- mum of four characters, the latter by six. There was also evidence for grouping of S. swaini, S. cornutus and S. salebrosus to the exclusion of S. occultus, although this rests in part on the inter- mediate state of the epipubic cartilage in S. occul- tus. The presence of two clearly defined monophyletic lineages within leaf-tailed geckos supports the conclusions of Bauer (1990) and our proposal to recognise Saltaurius as a genus. Saltuarius gen.nov. See Cogger, et al., 1983; Bauer, 1990. Referred Species Saltuarius comutus (Ogilby, 1892); S. occultus sp. nov.; S. salebrosus (Covacevich, 1975); S. swaini (Wells & Wellington, 1985). Diagnosis Nostril in contact with rostral (vs not contacting the rostral in Phyllurus); anterior margin of inter- clavicle flat, or forward projecting without dis- tinct process. Fig. 2a-d (vs bearing a distinct process); axilla not, or only very rarely and shal- lowly, invaginated (vs always, sometimes deeply invaginated); epipubic cartilage moderate to large, wedge-shaped, Fig. la-d (vs small - moderate, wedge-shaped); male preanal pores present, Fig. 5a-c, save for one species, Fig. 5d 98 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1: Character matrix on which the phylogeny (Fig. 4) for species of Phyllums and Saltuarius gen. nov. is based. This analysis includes only the char- acters that vary within the leaf-tailed geckos. Monophyly of Saltuarius and Phyllurus with respect to other carphodactyline geckos was X!! AH Kl U li •i xi gg 1 9 fi demonstrated by Bauer (1990). 1 ft « Characters ,1-15 • ► mm salebrosus swaini occultus corutus platurus isis nepthys ossa caudiannulatus laevis 002110111111010 002170101111010 0011 ?? 01111 ? 01 ? 002110111111010 210001000000111 21007100000010 ? 1110?1000000101 210071000000101 210001000010101 010110010100000 Characters 1. Broadened process on anterior interclavicle exten- sion: 0, narrow splint; 1, intermediate; 2, broad 2. Anterior interclavicle extension: 0, absent; 1 , present 3. Epipubic cartilage: 0, not expanded; 1, moderate; 2, greatly expanded 4.SuprdOcular portion of frontal: 0, flat; 1, furrowed S.Inscriptional ribs: 0, 0-1 ; L 2-4; 2, 5-7 ribs. ^ b.Poslmental scales: 0, enlarged anteriorly; 1, subequal 7. Scales of rosettes: 0, not spinose; 1, spinose S.Preanal pores: 0, absent; 1, present 9. Rostral contacting nostril: 0, no contact; 1, contact 10. Body size: 0, max SVL <103mm; 1, niax SVL >103mm 11. Rostral: 0. not divided or only partially divided; 1, completely divided 12. TaiI: 0, simple; 1, elaborate edge flanges 13. Attenuated tip on regrown tail: 0, not elongate; 1, elongate 14. First autotomy septum: 0, 5; 1,6 15. Chromosome number: 0, 2n=38; I,2n<38 FIG. 3. Karyotype (2n=30) of Phyllums nepthys from Finch Hatton Gorge, MEQ. salebrosus cornutus swaini occultus platurus isis ossa caudiannulatus nephthys laevis (vs male preanal pores absent); original tail^ strongly flared, with an elaborate outer flange, Fig. 6a (vs cylindrical to simply flared, lacking an elaborate outer flange. Fig. 6b); regenerated tail strongly flared, with only a small attenuated lip (vs cylindrical to moderately flared, with a pronounced attenuated tip). Max SVL 108- 144mm, medium to large (vs max SVL 76- 103mm^ small - medium); karyotype 2n=38 (vs 2n=30 or 22); rostral completely divided. Fig. FIG. 4. Phylogenetic hypothesis for species of Phyl- lurus and Saltuarius derived from parsimony analysis of morphological characters with ordered character states and 100 bootstrap replicates. The numbers above the branches are the number of replicates in which the group to the right was monophylelic. The analysis provides strong support for the monophyly of species within Saltuarius and within Phyllums. The monophyly of the two genera with respect to Carphylodactylus laevis is assumed on the basis of Bauer’s (1990) analysis. ^The original tail of 5". occultus is not known, because the species is based on only 4 type specimens, none with original tails. Bauer (1990) gives SVL 112 mm for one P. caudiannulatus. TTie max SVL for P. caudiannulatus here is 91.24 mm. Since Bauer’s work, P. nepthys has been separated from P. caudianannulatus. The max. SVL for P. nepthys is 103 mm. REVIEW OF LEAF-TAILED GECKOS 99 FIG. 5. Presence or absence of preanal pores in Saltuarius spp. Preanal pores present (A) S. comutus - J5 1097, (B) S. occultus - J37038, (C) S. salebrosus - J33730. Preanal pores absent (D) 5. swaini - J5 1639. Habitat S. occultus and S. comutus, are obligatory rain- forest species. S. salebrosus occurs in complex notophyll vine forest in Bulburin SF, SEQ, and in araucarian vine thicket in the Goodnight Scrub, SEQ. Throughout the rest of its range, it is found in dry rocky (sandstone) situations. S. swaini is found in both rainforest, and in granite-based heathlands. Reproduction Oviparous, producing two soft-shelled eggs. Etymology From the Latin - ‘keeper of the forest’. A key to Saltuarius 1 .Throat smooth 2 Throat strongly tuberculate S. salebrosus 2. Lower flank tubercles hooked and surrounded by spinose basal scales (Fig. 9a) S. comutus Lower flank tubercles not as above 3 3. Neck extremely elongate and slender (Fig. 10a) S. occultus Neck not as above (Fig. 1 Ob) S . swaini 7a-d (vs partially divided. Fig. 7e-j, save for one species). Distribution Eastern Australia: from Mcllwraith Ra., Cape York Peninsula, FNEQ, to central coastal NSW (Fig. 8). Individual species accounts contain further details. FIG. 6. Original tails of (A) S. salebrosus (J33730) - strongly flared with an elaborate outer flange. (B) P. platurus (R106497) - simply flared, lacking an elaborate outer flange. 100 MEMOIRS OF THE QUEENSLAND MUSEUM I J FIG. 7. Variations in rostral grooving (A-D) S. swaini (Fig. 7A is the typical condition for all Saltuarius spp. and also for P. caudiamulatus); (E) P. isis (also the condition for P. nepthys); (F-H) P. ossa; (I-J) P. platurus. Saltuarius cornutus (Ogilby 1892) (Figs la; 5a,d;7a,b; 9a, b; 11; 12a, b; 13) Gymnodactylus cornutus Ogilby, 1 892. Phyllurus lichenosus Ganthcty 1897. Gymnodactylus spyrurus Barrett, 1950. Material Examined Queensland Museum: J25394, Home Rule (15°44’, 145°17’) NEQ; J27265, Mt Finlay (15°49’, 145°2r) NEQ; J51631'2, Bell Pk, Malbon Thompson Ra., (\r05\ 145° 52’) NEQ; J48279, Danbulla SF, Kauri Ck, Ml Haig Rd (17°07’. 145°38’); J25373, Atherton Tableland (17°16’, 145°29’) NEQ; J9532, Atherton, 11km SW Walsh Camp (17°20M45°25’) NEQ; J3099I, Boonjee, via Malanda (17°22’, I45°45’) NEQ; J52775, Millaa Millaa, Palmerston Hwy (17°3r, 145°37’) NEQ; J48254, Ravenshoe SF, near Vine Ck (]7°4r, 145°3r) NEQ; J48178 (alizarin stained), J48179, Koolmoon Ck, S of Ravenshoe (17°44’, 145°34’) NEQ; J28355, Koombooloomba Dam, viaTully Falls (I7°50\ 145°36’)NEQ; J510%- 7, Paluma SF, 2km from turnoff to Paluma Dam (19°01, 146°12’)NEQ; J2126,?Northem Rivers; J149, Queensland. Australian Museum; R26118-9, Evelyn, approx 14.5km from Ravenshoe (17°30’, 145°27’) NEQ; R128869, El Arish (17°49’, 146°00’) NEQ; R12935, (Queensland. The original description of S, cornutus is ac- curate. It has been refined by Covacevich (1975) and Bauer (1990). The former recognised three variable, geographically-isolated forms within S. cornutus - from rainforests of NEQ; SEQ- MENSW, and the granite-based heaths of the Stanthorpe area, SEQ. TTie latter recognised four forms of S.comutus - disjunct populations from near Coen, FNEQ; between Cooktown and Townsville, NEQ; from extreme SEQ to MENSW, and the Stanthorpe area, SEQ. Specimens from far northeastern Queensland were collected in 1 979, and referred to S. cornutus at registration into the Queensland Museum ref- erence collection. Cogger (1992) included them in his distribution map for Phyllurus (now SaU tuarius) cornutus and Covacevich & McDonald (1991a) note their collection locality, Mcllwraith Ra. FNEQ (13°50’, 143°180, as the northern limit of the range of Phyllurus {=Saltuarius) cor- nutus. The far northeastern Queensland specimens of T. cornutus' (which were not included in any of the descriptions of that species) are here REVIEW OF LEAF-TAILED GECKOS 101 B FIG. 9. Flank tubercles of (A) S. comutus (J48254), (B) S. swaini (J8183). described as S. occultus . Specimens of "P. cornutus' from SEQ- MENSW have been described as S. swaini (Wells & Wellington, 1 985). The type description is incomplete. It does not include any reference to the features that most readily separate S. swaini from S. cornutus and does not provide a diagnosis. These, and other deficiencies in the description are redressed under S. swaini. The population of T. cornutus' iso- lated on the Stanthorpe area granites is here referred to S. swaini. With these recent descriptions of new taxafrom the wide-ranging former 'P. cornutus', there are now three species of Saltuarius in addition to the nominate species. 5. cornutus sensu stricto is now confined to the rainforests of the area between Big Tableland and Mt. Spec, NEQ, in the Wet Tropics, one of Australia’s World Heritage sites. Thus, in the light of these changes, data for S. cornutus should be modified. Diagnosis 5. cornutus is a moderately spinose, large Sal- FIG. 10. X-rays showing the cervical vertebrae of (A) S. occultus and (B) 5. comutus. Elongation of the cervical vertebrae in S. occultus results in this species having a proportionately longer neck than other Saltuarius spp. Condition (B) is shared with S. salebrosus and S. swaini. 102 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 11.5. comutus, Paluma, NEQ (S. Richards). tuarius (maximum SVL 144mm). It is distin- guished from S. salebrosus by its chin scalation (smooth, with no raised tubercules vs rough, covered with raised tubercules); from S. occultus by flank tubercules (long and hooked - Fig. 9a, surrounded by smaller spines vs not long and hooked and not surrounded by smaller spines), by the number of lumbar vertebrae (2 - Fig. 1 2a vs 3 - Fig. 12b) and by the size of neck vertebrae (not elongate vs elongate); and from S, swaini by flank tubercules (long and hooked, surrounded by smaller spines - Fig. 9a vs not hooked and not surrounded by smaller spines - Fig. 9b). It is further distinguished from S. swaini by prcanal pores (males with preanal pores - Fig. 5a vs males without preanal pores - Fig. 5d). Description With the separation of S. occultus and S. swaini from S. cornutus, the descriptions provided by Covacevich (1975) and Bauer (1990) for this species require minor modification. Bauer(1990) refers to the flank tubercules as being ‘enlarged and hooked’ (Fig, 9a). We found this character to be confined to NEQ (Big Tableland-Mt Spec) specimens. This character is absent in far NECJ specimens (S. occultus) and SEQ through MENSW specimens {S. swaini). Bauer’s sample (in litt.) was based predominantly on NSW and SEQ specimens from the Australian Museum’s collection. This explains why his diagnosis makes no reference to preanal pores, which are present in male S. cornutus (Fig. 5a) and absent in male 5. swaini (Fig. 5d). Covacevich (1975) recognised the difference in the flank tubercules between the NEQ (long hooked spines sur- rounded by smaller spines - Fig. 9a) and the SEQ-NSW (shorter, unhooked spines - Fig. 9b) populations. At the time she treated this as accept- able variation within a single species. The far NEQ specimens (5. occultus) were unknown at the time of her revision. We now regard this feature (long hooked flank tubercules surrounded by smaller spines - Fig. 9a) as key to the diagnosis of S. cornutus. Covacevich (1975) and Bauer (1990) provide size ranges for ‘F. cornutus'. Of the three species now recognised from the former P. cornutus sensu lato, S. cornutus is the largest (maximum SVL 144mm). S. swaini has a maxi- mum SVL of 131mm and S. occultus (based on the 4 known specimens) has a maximum SVL of 108mm. The following measurements and characters have not been included in previous descriptions of S. cornutus: Proportions as % SVL: LI 43-50 (nl8, mean 46.4); L2 53-62 (nl7, mean 56.1); S REVIEW OF LEAF-TAILED GECKOS 103 FIG. 12. X-rays showing the number of lumbar vertebrae in (A) S. cornutus (J28355), two lumbar vertebrae; (B) S. occultus (J37040), firee lumbar vertebrae. Condition (A) is shared with S. salebrosus and S. swaini. 12-15 (n20, mean 13.0); NL 13-21 (n20, mean 17.3). Rostral completely divided by a single vertical groove, Fig. 7a (nl8), or by two grooves forming an additional scale and dividing the rostral into three (n2), seen also in 5. swaini, (Fig. 7b). Axilla not invaginated. Preanal pores present in males (Fig. 5a). Skeletal Features Material examined: (X rays) J28355, J30991, J48254; (alizarin stained) J48178. Supraocular portion of frontal grooved (n4); anterior process of interclavicle projecting slight- ly forward, Fig. 2a (n 1 ); epipubic cartilage greatly expanded, Fig. la (nl); presacral vertebrae 25-26 (n3, mean 25.3); sacral vertebrae 2 (n3); lumbar vertebrae, 2-Fig. 12a (n3); 1st autotomy septum 6 (nl); abdominal vertebrae bearing reduced ribs 4 (nl); rib free cervicals 3 (n3); cervical vertebrae not enlongate; sternal ribs 3 (n 1 ); mesostemal ri bs 2(nl). Distribution Big Tableland (15°43\ 145°16\ sight record, K.McDonald pers. comm.) to Mt Spec (19°0r, 146°12’) NEQ (Fig. 13). The locality given for QM J212 is ‘?Northern Rivers’, which usually refers to an area in northern NSW. The previous entry in the register (QM J2125) is a specimen of Nephrurus asper from the same locality. As neither S. cornutus nor A. asper occurs in NSW, nor are they sympatric elsewhere, the locality ‘?Northem Rivers’ should probably be regarded as an error. Habitat Tree trunks and rocks, tropical rainforest. Reproduction A female from the Malbon Thompson Ra., NEQ laid two oval-shaped eggs on I December, 1990 (±3 days). These measured 26.62 x 16.05 mm and 27.24 x 16.44 mm. The eggs took 100 ±3 days to hatch at a temperature of approximately 24°C. Hatchlings measured (SVL) 50.63mm and 51.41mm. 104 MEMOIRS OF THE QUEENSLAND MUSEUM Diet A faecal sample from QM J48179 contained fragments of cockroaches, a cricket, and a spider (Blattellidae, Gryllidae and Sparassidae). Saltuarius occultus sp. nov. (Figs lb, 2b, 5a-d, 7a, 10a, 12b, 14, 15) Material Examined Holotype: Queensland Museum: J37040 9 (with regenerated tail) Peach Ck, Mcllwraith Ra., 1 9km ENE of Mt Croll (13°45’, I43°19’) FNEQ, J.W. Winter, R.G. Atherton, P.A. Matthew, 7 June 1979. Paratypes: Queensland Museum: J37037 (alizarin preparation), J37038-9 sununit of Mcllwraith Ra., I7km ENE of Mt Croll (13°46’, 143°19’), FNEQ. Diagnosis S. occultus is a moderately spinose, medium- sized Saltuarius (maximum SVL 108mm). An extremely long, thin neck, a function of elongate neck vertebrae (Fig. 10a), mean % NL/SVL 24.2; a series of enlarged, spinose tubercles fringing the regenerated tail; and three lumbar vertebrae (Fig. 12b) separate it from all its congeners. S. occultus resembles S. cornutus and S. salebrosus in having male pre-anal pores (Fig. 5a-c). These are not present in S. swaini (Fig. 5d). In S. occultus they are very pronounced, and form a raised cluster (vs not pronounced, only just visible without magnification, and flat). Distribution Mcllwraith Ra. (13^45’, 143°19*), Cape York Peninsula, FNEQ (Fig. 15). Habitat Microphyll / notophyll vine forests, on a variety of substrates, including granite. Habits Known from only four specimens, all of which were found at night, on granite boulders in rain- forest. Reproduction The only females known (J37037, J37040) were found with enlarged vitellogenic follicles between late May and early June, 1974. Diet ^ The gut of QM J37038 contained spider frag- ments. Etymology From the Latin - ‘hidden’ and so, difficult to find. Description SVL (mm): 99-108 (n4, mean 102.7). Propor- tions as % SVL; LI 45^9 (n4, mean 47.1); L2 56-61 (n4, mean 59.0); HL 25-27 (n4, mean ; 25.8); HW 16-19 (n4, mean 17.2); S 12-14 (n4, mean 13.0); NL 23-27 (n4, mean 24.2). Head: large, depressed, elongate, shovel- shaped, distinct from neck; covered in small granules which are intermixed with larger conical tubercles (top of snout without larger tubercles); skin of head and skull not co-ossified; rostral divided, bisected by a deep vertical groove (Fig. 7a); rostral contacting nostril; ear opening ex- tremely narrow, elliptical, vertical, much less than half as large as eye; supralabials 19-21 (n8, mean 19.75), infralabials 18-22 (n8, mean 19.5). Neck: long and slender. Body: moderate, depressed, covered in small granules; dorsal granules intermixed with larger conical tubercles; REVIEW OF LEAF-TAILED GECKOS 105 1 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 cz> •car FIG. 14. 5. occultus (hololype J37040), Peach Ck, McIIwraith Ra., NEQ (B. Cowell). tubercles moderate on back, small on flanks, no more prominent on sides of neck than back; basal scales surrounding flank tubercles not enlarged; no enlarged tubercles or granules on ventral sur- face of body. Preanal Pores: present in males, extremely prominent (Fig. 5b). Limbs: long, covered in pointed tubercles dorsally (much more prominent on hind limb); proximal portion of forelimb bearing slightly enlarged tubercules on ventral surface; digits strong, compre.ssed distal- ly; subdigital lamellae (fourth toe) 23-25 (n4, mean 24). Original tail: unknown, because all of the type series have regenerated tails. Reg- enerated tail: (n4) depressSl, broad and leaf-like, contracted at base, not attenuated at tip; lacking spinose tubercles on dorsal surface; edges sur- rounded by broad spinose tubercles. Skeletal Features Material examined: (X rays) J37038-40; (alizarin stained) J37037. Supraocuhir portion of frontal grooved (n4); anterior process of inter- clavicle projecting slightly forward (Fig. 2b) n 1 ; epipubic cartilage slightly expanded (Fig. lb) n 1; presacral vertebrae 25 (n3); sacral vertebrae 2 (n3); lumbar vertebrae 3 (Fig. 12b) n 3; 1st autotomy septum 6 (n3); abdominal vertebrae bearing reduced ribs 3 (nl); rib free cervicals 3 (n3); cervical vertebrae greatly enlarged (Fig. lOa); sternal ribs 3 (nl); mesostemal ribs 2 (nl). Pattern In spirit, dorsal base colour tan; heavily marked 106 MEMOIRS OF THE QUEENSLAND MUSEUM with large distinct, dark brown blotches which form a series of four irregular crossbands between the axilla and groin. Limbs bearing irregular dark brown crossbands. Head marked with less dis- tinct brown blotchings. Ventral surface cream with pale brown mottling on belly, throat and limbs. Labials, mainly dark brown broken by patches of white. Regenerated tail tan to cream, mottled brown dorsally and ventrally. Saltuarius salebrosus (Covacevich, 1975) (Figs 7a, b; 16; 17) J2874I, Goodnight Scrub, via Wallaville (25°12’, 151 ®55’) SEQ; J6198, Burnett R., Goodnight Scrub (25°12\ 15U550 SEQ; J28802, J29778, J36115, Robinson Gorge, Taroom (25°17\ 149°09’); SEQ; J6382, Cracow, Cracow Ck. (25°I8\ 150°18’) SEQ. Little that is new can be added to the data on S. salebrosus. The type description is accurate and no change to the species definition is warranted following re-examination of the specimens of Covacevich (1975), and examination of those added to museum collections recently. The following measurements and characters have not been included in previous descriptions of 5. salebrosus: Material Examined Queensland Museum; J51091-2, Blackdown Tableland, 500m from top (23°46\ 149°06’) MEQ; J36114, Cania Kroombil goldfield , near Dawes Ra., via Monto (24^38’, I50°58’) SEQ; J33730-2, Granite Ck. tributary crossing to forestry camp, Bulburin (24°31\ I5I°29’) SEQ; J33700 (alizarin stained), J51090, Bulburin SF (24’=3I’. 15r29*) SEQ; J9770, Lowmead (24°32\ 15U45’) SEQ; J22288, Bulburin SF, Granite Ck, via Many Peaks (24^35^ I5r29*) SEQ; J8377, Injune (25^51’, 148°34’) SEQ; J8142, Monto (24°52\ I5r07’) SEQ; J56919, Bania SF, via Monto (24^57’, 150^30’) SEQ; J5390, J25360, Proportions as % SVL: LI 45-50 (nl9, mean 47.1); L2 53-61 (nl7, mean 57.1); S 11-13 (n20, mean 12.2); NL 15-21 (n20, mean 18.5). Rostral completely divided by a single vertical groove (Fig. 7a) n 17, or by two grooves forming an additional scale and dividing the rostral into three (n2) seen also in S. swaini (Fig. 7b). The axilla is normally not invaginated (nl6), but is sometimes slightly invaginated (n4). For skeletal description see Bauer, (1990). Some modification to distribution of S. saleb- rosus is required following re-examination of FIG. 16. S . salebrosus, Bulburin SF, SEQ (S. Wilson). REVIEW OF LEAF-TAILED GECKOS 107 FIG. 17. Distribution of 5. salebrosus. data for one specimen originally believed to be from south-central Queensland, remote from the near-coastal range of the species. The record was based on QM J4897 ‘Coongoola, SCQ’. The specimen in fact comes from Coombooloolaroo Station, a locality well within the near-coastal range of S. salebrosus, and quite close to the Blackdown Tableland, an area in which S. salebrosus is well known now. In addition to restricting the western range of S. salebrosus, new data extends it a little to the north and slightly to the south west of its former range, Fig. 17 (minus the ‘Coongoola, SCQ’ record). New records since those of Covacevich (1975) include Blackdown Tableland, MEQ, J35400, J35448, J5 1091-2 (23°46\ 149°06’), where specimens of S. salebrosus are common in heaths on both sandstones and granites, and Bania State Forest, via Monto, SEQ, J56919, (24°57’, 150°30’) in rainforest. Reproduction New data on breeding by S. salebrosus are also available. A young specimen (J51090, SVL 52.8mm) was collected at Bulburin State Forest on 1 8 May, 1991 . This individual is similar in size to newly emerged S. cornutus hatchlings (50.63- 51.41mm). It seems reasonable to regard it as a newly emerged hatchling. The incubation period recorded for S. cornutus is approximately 103 days. Because of close taxonomic affinity, it seems reasonable to suggest that the eggs of S. cornutus and S. salebrosus would have a similar incubation period. If this is so, the oviposition date for hatchling J51091 would have been early February, 1990. Diet The gut of QM J33732 from rainforest, con- tained one large cockroach and a pygmy gras- shopper (Tetrigidae). QM J51092 from open forest, contained fragments of a spider. Saltuarius swaini (Wells & Wellington,1985) (Figs 2d; 5a, d; 7a-d; 9a,b; 18a,b; 19) Phyllurus swaini Wells & Wellington, 1985. The specimen chosen as the holotype of S. swaini was unfortunate considering the many specimens available to the authors. Specimen AM RI 16978 (formerly AM Field Series no. 16799), the holotype, is extremely faded and all but patternless; has been totally eviscerated; has a badly damaged left lower jaw; and has a separate, regenerated tail. The type description of S, swaini, which is based on this specimen and one published photograph, is neither complete nor accurate. Characters key to separating S. swaini from S. cornutus are not described (i.e. smaller size, lateral tubercles that are neither hooked nor surrounded by at least some spinose basal scales (Fig. 9b), and the absence of preanal pores (Fig. 5d). Further, part of the discussion of this species is erroneous. ... ‘Another species occasionally confused with this taxon is Phyl- lurus caudiannulatus of southeast Queensland; Covacevich, 1975, included northeast New South Wales specimens of P. swaini in her analysis of P. caudiannulatus'.... P. caudiannulatus has not been (and could hardly be) confused with S. swaini. The former is very distinct in being con- siderably smaller than 5. swaini, and in having a distinctly white-banded original tail. It is restricted to the Dawes/Many Peaks Ranges, SEQ. No mention of specimens from north- eastern New South Wales is made in the descrip- tion of P. caudiannulatus by Covacevich, (1975) although she does refer to P. platurus of ...‘ more open forest in rocky, especially sandstone areas of mideastem New South Wales’... This re-description of S. swaini is based on a sample of specimens selected from the collec- tions of the Queensland and Australian Museums, to give maximum range in terms of distribution and morphological diversity. Material Examined Austral ian Museum R 1 1 6978, holotype, (formerly AM 108 MEMOIRS OF THE QUEENSLAND MUSEUM Field Series no. 16799) Richmond Ra. SF, (28°3r, 152°44’) NSW. Other SpecimensiQueensland Museum: J398, J2409. J2933-34, J3254, J4439, J8 1 83, J8359, J886 1 , J 1 0440. J 1 2257, J5 1 095 Mt Tamborine, (27°55\ 153'’10’) SEQ; J4819, Mt Tamborine, Eagle Heights (27‘^55’, 153°120 SEQ; J148 Canungra Ck (27°58^ 153W) SEQ; J3215 Canungra (28“0r, 153°ir) SEQ; J4I98, J5690 Mudgeeraba (28*^05’. 153°22’) SEQ; J5649 Rying Fox Valley, Beechmont (28°08’. 153^12’) SEQ; J33I3 Tallebudgera (28“08\ 153°26') SEQ; J5382 Lamington NF (28°12\ 153°05’) SEQ; J8646 Lamington NP Binna Burra (28°12\ 153'’! r) SEQ J23937 near Mt. Ballow, (28°16\ 152^37') SEQ; J51094 Mt. Superbus SF, via Warwick, (28°13*, 152'^28’) SEQ; J5 1639 151637^0 O’Reilly’s, Lamington NP (28°!4\ 153^08’) SEQ; J8074, J8075 (alizarin stained), J8099 Mt Clunie, via Boonah (28°18\ 152^32’) SEQ; JI143 Tweed R. (28°18’, 153^^27’) NSW; J5757 Chillingham, Murwil- lumbah (28^19’, 153*^17’) NSW; JI0565 Mt Lindesay (28°23M 52^=43’) SEQ; J35401 BoonooBoonoo Falls, via Tenterfield (28°48’, 152°10’) NSW; J24250, J27349 near Girraween (28°50\ 151^55’) SEQ; J25374 Girraween NP, via Slanlhorpe (28°50’, 15 1°55*) SEQ; J28648-9 Girraween area, near Wyber- ba (28"50\ 15U55’) SEQ; J29115-7 Stanthorpe area, ?Girraween (28^50’, I5r55’) SEQ; J30677 Stan- Ihorpe, Aztec Temples, near underground river (28°50’, 152°05’)SEQ;J51093 edge of Girraween NP (28°50’. I51°56’) SEQ; J51633-6 Girraween NP, Natural Arch track (28°50\ I51°55’) SEQ; J54847 Bookookoorara, Boonoo SF (28°5r. 152'^] U) NSW J50345 Girraween NP (28°5 ! M 5 I ”55 ’ ) SEQ; J30420 near Wyberba (28^52’, 15I^S2') SEQ; J53984 Teapot Ck, Narrow Pass Fire Trail, Mann R. Nature Reserve (29°45\ 152*^02’) NSW; J54846 Bray’s Ck. Border Ranges NP (28^24’, I53TO’) NSW; J9054 Bulah- delah, 96km NE Newcastle (32^25’. 152^12’) NSW; J56894 found on warehouse wall, Brisbane. Australian Museum: R97823 Mt Superbus,(28°13’, 152°26’) SEQ; R2409 Murwillumbah, Tweed R., (28°20’, I53°24’) NSW; RI10510 Mt. Warning, (28°24’, 153°16’)NSW; R1 1860Huonbrook(28°32M53°2r) NSW; R92121-3 6km NW of Amiens, (28°34’, 15U46’) SEQ; R98332-3 ca. L5km NW of Amiens, (28°34’, I5U46’)SEQ;R1309I1 YabbraSF,{28°37’, 152°30’ ) NSW; AM Field No. 4 1 650 B lack Ck, 6. 1 km SE along Black Hole Trail, Curramore SF, (29°30’, 152°ir) NSW; AM Field No. 41770 Willy’s Ck. 300m downstream from Oakwood Fire Trail, (29°53’, 152°02’) NSW; R43870 33km E of Guyra, (30^15’, 152°00’) NSW; R69866-7, R8192I Coff’s Harbour, (30°18\ I53°08’) NSW; R43875 Dorrigo, (30"20’. 152°43’)NSW; R4387I-3, R 1 6905, R 1 6989, R17008 Pt Lookout, via Armidale, (30^^29’, 152^25’) NSW; R54071 Brinerville, via Thora, (30°29’, 152^33’) NSW; R97670-2, R 106749 Cogger’s property, near Brinerville, (30^29’, 152''33’) NSW; AM Field No. 41683 first falls below Blue Hole. Gara R. Gorge, (30°36’, 151 °48’) NSW; R6284 Gurravembi, near Macksvilleon Nambucca R., (30°44’, 152°59’)NSW; R 103031 4.5km N. of Wonders Hill Homestead, (31°16’, 152°19’) NSW; R71372-3 Comboyne Pk, (3r35’, 152^32’) NSW; R6247 Bulga Tableland, near Bobin, Manning R., (3F37’, 152°15’) NSW; R6915 Bulong, via Wingham (31°37’, 152°18’) NSW; R59313 vicinity of Bird Tree, Middle Brother SF, (31°4r, 152M2’) NSW; R593I4 5km W. of highway on Middle Brother Mt (3P4r, 152M2’) NSW; R43874 Lansdowne, Manning R., (3r47’, 152^32’) NSW; R8253 Wallis Lake, Tuncurry, (32°17’. I52"29’) NSW; R101338 O’Sullivan’s Gap Forestry Reserve, Buladelah, (32°20’, 152°16’) NSW; RI5412 Girvan, via Stroud, (32°28’, 152 W’) NSW; R 123490 Tullawudjah Ck. (?) NSW. Diagnosis S. swaini is a moderately-spinose, large Sal- tuarius (maximum SVL 131mm). It is distin- guished from S. cornutus by its flank tubercules (tubercules not hooked and not surrounded by smaller spines, Fig. 9b vs tubercules hooked and surrounded by smaller spines. Fig. 9a); from S. salehrosus by its chin scalation (chin lacking tubercles vs chin with tubercles); from S. occultus by the colour of its labial scales (labials pale, blotched with grey or brown vs labials dark brown with a few patches of white). It is further distinguished from S. cornutus by male preanai pores (males without preanai pores, Fig. 5d vs males with preanai pores. Fig. 5a). Diet The gut of QM J5 1 640 contained a large cricket (Gryllacrididae). REVIEW OF LEAF-TAILED GECKOS 109 DESCRIPTION SVL (mm); 41-131 (nlOl, mean 92.0). Adult SVL (mm): 82-131 (n81, mean 102.6). Propor- tions as % SVL: LI 37-62 (n83, mean 45.1); L2 50-65 (n81, mean 56.3); T 58-81 (n43, mean 68.1); TT 20-33 (n43, mean 27.8); HL 26-32 (nl01,mean 28.1); HW 19-25 (nlOl, mean 21.5); S 1 1-14 (nlOl, mean 12.4); NL 15-23 (n95,mean 19.7). Head: large, depressed, triangular, distinct from neck; covered in small granules which are inter- mixed with large rounded to conical tubercles; skin of head co-ossified with skull; rostral com- pletely divided, usually by a single deep groove (nlOl, for variations sec Fig. 7a-d); rostral con- tacting nostril; ear opening elliptical, vertical, much less than half as large as eye; supralabials ll-20(nl98, mean 14.5); infralabials 9-1 6 (n 192, mean 12.4); Neck: broad; Body; moderate, depressed, covered in small granules; dorsal granules intermixed with larger conical tubercles; tubercles moderate to large on back, flanks and neck; lower flank tubercles small to large, often associated with a lateral flange running from axil- la to groin; basal scales surrounding flank tubercles slightly enlarged (Fig. 9b); granules on chest often noticeably larger than adjacent granules. Preanal Pores: Absent (Fig. 5d). Limbs: long, covered in pointed tubercles dorsally; digits strong, compressed distally; subdigital lamellae (fourth toe) 18-26 (nl64, mean 21.7). Original tail: (n43) - depressed, broad and contracted at base and attenuated at tip; anterior flared portion surrounded by an undulating flange which bears slender, sharply pointed tubercles around its mar- gin; dorsal surface of tail (except along midline of flared portion) covered in large conical tubercles which are particularly pronounced on the attenuated tip; tubercles extend to tail tip; number of rows of enlarged spinose tubercles across attenuated tip 2-7 (n43, mean 5.3); atten- uated tip accounts for 34-45% of total tail length; ventral surface smooth, some specimens with a shallow groove along the midlinc. Regenerated tail: (n37) depressed, broad and leaf-like, con- tracted at base and only just attenuated at tip; tail margin is a broad, thin flange which bears minute spinose tubercles around the edges; tail free from spinose tubercles on both dorsal and ventral sur- faces; ventral surface without any indication of a shallow groove along the midline. Skeletal Features (X rays)QM J2409, J3313, J4439, J5649, J8359, J28648-9, J51633, J5 1635-6, J53984; (alizarin stained) J8075, J291 15. Supraocular portion of frontal grooved (nl3); anterior process of interclavicle not present, anterior edge flat. Fig. 2d(n2); epipubic cartilage expanded. Fig. Id (n2); presacral vertebrae 24-25 (n8, mean 24.8); sacral vertebrae 2 (n9); lumbar vertebrae 2 (n7); 1st autotomy septum 6 (n6); abdominal vertebrae bearing reduced ribs 3-4 (n2, mean 3); rib free cervicals 3 (nIO); cervical ver- tebrae not enlongate; sternal ribs 3 (n2); meso- stemal ribs 2 (nl). Pattern In spirit, three colour forms are readily distin- guished by dorsal colour and pattern - gray to medium brown with both paler and darker blotches in the base colour; these blotches are edged with brown or black lines to give a ‘lichen- like’ effect (Fig. 19a); often with a pale vertebral streak: gray or mid-dark brown with a pale ver- tebral streak: heavily blotched with pale gray and black, often with a pale fawn vertebral streak (Fig. 19b). These three forms have cream ventral sur- faces, faintly to heavily marked by small clusters of brown scales. Original tails are marked with ‘lichen-like’ blotches on dorsal surface. Some specimens of the three forms have indistinct light bands across the attenuated tip, sometimes ex- tending to the ventral surface. Ventrally the tails in all three forms are grey to brown, bearing pale blotches and small dark specks. Regenerated tails are pale with cream and brown mottling. Fonns ‘lichen-like’ (Fig. 19a) and ‘gray- mid- dark brown’ are the dominant colour morphs in rainforests of southeastern Queensland and New South Wales. Form ‘heavily blotched’ (Fig. 19b) is dominant in dry eucalypt/granite habitats centred on the Stanthorpe region, SEQ, and the New England Tableland, NSW. However, ‘lichen-iike’ (R98333) and ‘gray- mid-dark brown’ (R92122) individuals also occur in dry forests near Stanthorpe, and one ‘heavily- blotched’ individual (R 11 05 10) has been col- lected from rainforest on Mt Warning, NENSW. Phyllurus Goldfuss, 1820 See Bauer, 1990. Referred Species Phyllurus caudiannulatus Covacevich, 1975; P. isis sp. nov.; P. nepthys sp, nov.; P. ossa sp. nov.; P. platurus (Shaw, 1790). 110 MEMOIRS OF THE QUEENSLAND MUSEUM mmm FIG. 19. (A) S. swaini (J51094) Ml. Superbus, SEQ (S. Wilson). (B) S. swaini Girraween NP, SEQ (S. Wilson.) REVIEW OF LEAF-TAILED GECKOS 111 FIG. 20. Distribution of Phyllurus spp. Diagnosis Nostril not in contact with rostral shield (vs nostril contacts the rostral in Saltuarius); anterior margin of the interclavicle with a distinct process, Fig. 2e-h (vs flat or bearing a slight projection, Fig. 2a“d); axilla always, sometimes deeply, in- vaginated (vs very rarely, invariably shallowly invaginated); epipubic cartilage small - moderate, wedge-shaped, Fig. le-h (vs moderate - large, wedge shaped. Fig. la-d); male preanal pores absent; original tail cylindrical, without flaring to simply flared, carrot-like (vs strongly flared, with an elaborate spinose outer flange); max SVL 76- 103mm, small to medium (vs max SVL 108- 144mm, medium to large); karyotype 2n=30 or 22 (vs 2n=38); rostral shield usually partially divided^ Fig. 7e-j (vs rostral always completely divided. Fig. 7a-d). Distribution Eastern Australia from the Mackay/ Proseipine district MEQ - to the Sydney district, NSW (Fig. 20 ). Habitat P. caudiannulatus, P. isis, P. nepthys and P. ossa occur in complex notophyll vine forests on a variety of substrates. F. platurus is confined to heaths on sandstones. Reproduction Oviparous, producing two soft-shelled eggs. A KEY TO PHYLLURUS 1 .Rostral completely divided (Fig. 7a) P. caudiannulatus Rostral not as above (Fig. 7e-j) 2 2.Venter distinctly ‘peppered* with brown... P. nepthys Venter not as above 3 3.0riginal tail plain, without light bands P. platurus Original tail not as above (with light bands) 4 4. Rostral with 2-3 partial divisions (Fig. 7f,g) ..P. ossa Rostral not as above, 1 partial division (Fig. 7e,h). 5 5. Anterior-most white tail marking a broken band P. ossa Anterior-most white tail marking two narrowly- spaced, midline blotches P. isis Phyllurus caudiannulatus Covacevich, 1975 (Figs 7a, e; 21; 22) Phyllurus nepthys Couper et aL 1993. In part. Material Examined Queensland Museum: J 1 56 1 9, Holotype, Bulburin SF, 24km SW Miriam Vale (24°3r, 15r29’) SEQ; J28356, J3363 1 , J33649, J3365 1 , J33684-5, J33692-5, J33703-4, J33709, J51 1034, as for holotype. Australian Museum: R47641, R47654, R47657, R47888,R90205, Bulburin SF(24^3r,15r29’)SEQ. The type description of P. caudiannulatus is accurate, but requires modification in light of our separation of P. nepthys from it. P. caudian- nulatus was described from Bulburin SF, SEQ. As the description went to press, a specimen (J2541 1) apparently similar to P caudiannulatus, came to hand from isolated rainforest of the Eun- gella NP (Clarke Ra.) MEQ, some 480km north of Bulburin. This was referred to P caudian- nulatus, despite recognition of certain differen- ces, notably in the tail, between material from Bulburin SF and Eungella NP. Re-examination of specimens of P caudiannulatus, of the Eun- gella specimen referred to it, and examination of ^P. caudiarmualtus is the exception. It has a completely divided rostral, a character oi Saltuarius gen.nov. 112 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 21. P. caudiannulatuSy Bulburin SF, SEQ (B. Cowell). additional specimens recently collected from both localities, shows that the differences originally observed are consistent and that there are other distinct characters. Thus, we recognise as distinct P. nepthys sp. nov. from the Eungella NP and Crediton SF on the Clarke Ra., MEQ. Description Now removed from the series of specimens on which the description of P. caudiannulatus was based is QM J2541 1. P. caudiannulatus and P. nepthys resemble each other in size, body propor- tions, colour pattern and spinosity. They differ in the condition of the rostral scale (completely divided in P. caudiannulatus. Fig. 7a vs piirtially divided in P, nepthys. Fig. 7e). Further differen- ces are given in the description of P. nepthys. The following measurements and characters have not been included in previous descriptions of P. caudiannulatus. Proportions as % SVL: LI 37-45 (n20, mean 42.3); L2 46-54 (n20, mean 50.6); S 11-12 (n20, mean 1 1.6); NL 17-23 (nl9, mean 19.5). Rostral completely divided by a single vertical groove - Fig. 7a (n20). Axilla shallowly to deeply invaginated. Distribution The range of P. caudiannulatus is also restricted with the removal from P. caudian- nulatus of specimens from the Clarke Ra. P. caudiannulatus is now narrowly restricted to rainforests of Bulburin SF, on the Dawes Ra. and Many Peaks Ra., SEQ (Fig. 22). Bauer (1990) notes its occurrence ‘to the south and west of Brisbane*. This locality is based on an AMNH specimen from Nerang. Given the SVL (112 mm) of this specimen, it seems reasonable to suggest that it is referable to S. swaini, which is known to occur in the Nerang area. Diet The gut of QM J33709 contained numerous small beetle fragments, and pieces of a large beetle larva (Coleoptera). Skeletal Features Material examined: (X rays) J28356, J33651, J33684-5, J33693, J33695, J33703-4, J33709; (alizarin stained) J33631. Supraocular portion of frontal flat (n9); anterior process of interclavicle distinct(nl); epipubic cartilage not expanded (nl ); presacral vertebrae 26 (n9); sacral vertebrae REVIEW OF LEAF-TAILED GECKOS 113 FIG. 22. Distribution of P. caudiannulatus. 2 (n9); lumbar vertebrae 2 (n9); 1st autotomy septum 5 (n5); abdominal vertebrae bearing reduced ribs 4 (nl); rib free cervicals 3 (n9); sternal ribs 2 (nl); mesostemal ribs 3 (nl). Phyllurus isis sp. nov. (Figs lf;2f;7a,e-h;23;24) Material Examined Holotype: Queensland Museum J 53511 gravid 9 (with original tail) Mt Blackwood NP (21°02’, 148°56’), MEQ, P.J. Couper, J.A. Covacevich, K.R. McDonald, 10 October 1991 . Paratypes: Queensland Museum: J 53485-6; J53518 Coffee Ck, Mt Jukes (2r00’ 148°57’), MEQ; J 53480, J53512 (alizarin preparation), J 53591, J53602-3 Mt Blackwood NP (2r02\ 148°56’),MEQ. Diagnosis P, isis is the least spinose and smallest Phyl- lurus (maximum SVL 76mm). It most resembles P. ossa. From P. ossa it is distinguished by rostral grooves (a single groove partially dividing the rostral, Fig. 7e vs 2-3 grooves, usually 3, rarely 1 , partially dividing the rostral, Fig. 7f-h). It is fur- ther distinguished from P. ossa by the size of its flank tubercules (small vs moderate). P. isis is readily distinguished from P. caudiannulatus by the shape of both its original and regrown tail (flared vs cylindrical) and by the nature of the rostral groove (rostral partially divided, Fig. 7e vs rostral fully divided, Fig. 7a); from P. nepthys by ventral colour pattern (immaculate vs ‘peppered’ brown); from P. platurus by original tail colour pattern (distinctly banded white vs lacking white bands). Distribution Mt Blackwood (2r02’, 148°56’) and Mt Jukes (2r02’, 148°57’), Mackay district, MEQ (Fig. 24). Habitat Complex notophyll vine forest, on quartz- syenite and granite. Habits All specimens examined, except QM J53512, were collected during the early evening on rock outcrops. J53512, also active during the early evening, was found sitting on a stick on the forest floor, after light rain. Reproduction Gravid females (J5351 1, J53512, J53602) were present in the population in early October, 1992. Diet The gut of QM J53512 contained moth, cock- roach and cricket fragments (Lepidoptera, Blat- todea; Gryllidae). Etymology Selected from Egyptian mythology (Macken- zie, 1978). The epithet is to be treated as a noun in apposition. Description SVL (mm): 53-76 (n9, mean 69.2). Adult SVL (mm): 68-72 (n8, mean 71.2). Proportions as % SVL: LI 41-45 (n9, mean 42.6); L2 48-56 (n9, mean 52. 1 ); T 73 (n 1 ); TT 40 (n 1 ); HL 28-32 (n9, mean 29.4); HW 22-26 (n9, mean 23.1); S 12-13 (n9, mean 12.2); NL 15-19 (n9, mean 16.8). Head: large, depressed, triangular, distinct from neck; covered in small granules which are inter- mixed with larger rounded to conical tubercles; skin of head co-ossified with skull; deep vertical groove partially dividing rostral scale (Fig. 7e); rostral excluded from nostril; ear opening ellipti- cal, vertical, much less than half as large as eye; supralabials 14-18 (nl8, mean 15.7); infralabials 13-15 (nl8, mean 13.9). Neck: broad. Body: 114 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 23. P. isis, Mt Blackwood, MEQ (B. Cowell). moderate, depressed, covered in small granules; dorsal granules intermixed with larger conical tubercles; tubercles indistinct on back, small on flanks, most prominent on sides of neck; basal scales surrounding flank tubercles not enlarged; no enlarged tubercles or granules on ventral sur- face of body. Preanal pores: absent. Axilla: In- vaginated. Limbsilong, covered in small pointed tubercles dorsally; proximal portion of forelimb bearing enlarged tubercles on ventral surface; digits strong, compressed distally; subdigital lamellae (fourth toe) 18-20 (n9, mean 18.6). Original tail: (nl) depressed, strongly flared, con- tracted at base and attenuated at tip; without en- larged, spinose edge tubercles; covered dorsally with numerous minute, spinose tubercles; tubercles terminate approximately half-way along the attenuated tip which is long, fine, and terminates with a minute rounded ‘knob’ distally; dorsal tubercles fail to form clearly defined rows anteriorly across the attenuated tip; attenuated tip accounts for 52% of total tail length; ventral surface smooth, deeply grooved along midline (excluding attenuated tip). Regenerated tail: (n6) depressed, flared, contracted at base and at- tenuated at tip; lacking spinose tubercles on dor- FIG. 24. Distribution of P. isis. REVIEW OF LEAF-TAILED GECKOS 115 sal surface and edges; ventral surface without deep groove along midline. Skeletal Features Material examined: (X rays) J53480, J53485, J53511, J53518, J53591, J53602, J53603; (alizarin stained) J53512. Supraocular portion of frontal flat (n7); anterior process of interclavicle distinct - Fig. 2f(nl); epipubic cartilage not expanded - Fig. If (nl); presacral vertebrae 26 (n7); sacral vertebrae 2 (n7); lumbar vertebrae 2 (n7); 1st autotomy sep- tum 5(n6); abdominal vertebrae bearing reduced ribs 4 (nl); rib free cervicals 3 (n7); sternal ribs 2 (nl); mesostemal ribs 3 (nl). Pattern In spirit, dorsal base colour grey to pale brown; heavily marked with distinct, dark grey to black blotches on head, body and limbs. Body ventrally cream, slightly translucent. Labials pale grey to brown, mottled with white. Original tail dorsally black, indistinctly patterned with irregular grey markings; three distinct creamish bands on at- tenuated tip, extending to the ventral surface; two broken bands which appear as one irregular blotch on either side of the vertebral line on anterior flared portion; ventrally grey, mottled with white. Regenerated tail totally lacking cream bands; dorsally, grey to brown with obscure creamish blotching; ventrally mottled but paler than dorsal surface. Phyllurus nepthys sp. nov. (Figs Ig; 2g; 7a,e,f-h; 25; 26) Phyllurus caudiannulatus Covacevich, 1975. In part. Material Examined Holotype: Queensland Museum J34058 9 (with original tail) Finch Hatton NP(21 °06’, 148°38’) MEQ, R. Monroe, J.A. Covacevich, P. Filcwood, 7-14 April 1975.PARATYPES: Queensland Museum: J50993 Eun- gella NP, Dalrymple Rd (2r03\ 148^34’) MEQ; J35128 Thurgood farm, 18km N Dalrymple Hts (21^04’, 148''36’)MEQ;J34024 (alizarin preparation), J34025, Finch Hatton NP (2P06’, 148^38’) MEQ; J51098-10] Finch Hatton Gorge (2^06’, 148'^38’) MEQ; J34076-9 Finch Hatton NP (2r06’, I48°38’) MEQ; J2541 1 (also a paralype of P. caudiannulatus) Broken R. headwaters, EungellaNP(21°08\ 148°30’) MEQ; J50992 Eungclla NP, near Vlasak property (2ri0’, 148^24’) MEQ; J32634-35, J32695 Crediton (2ri3\ 148°34') MEQ; J32669, J32674-76, J32696, J32733, J32740 Crediton (2L13\ 148'^34’) MEQ; J53330-2 Rocky Dam Ck, via Crediton (2n8’, 148^32’ ) MEQ; J53359-62 Rocky Dam Ck, via Crediton (21°]9’, 148°34’)MEQ. Australian Museum: R47901 -14, R47959Mt William, via Eungella (21°0r, 148°36’) MEQ; R47512 R47836-49 Eungella (2r08\ 148°30’) MEQ; R47738-62 Eungella, Dalrymple Hts (21°08’, 148°30’) MEQ; R47551-6, R47957 Eungella, near Vlasak property (2nOM48°24’)MEQ; R61473 AM building? - no original locality data. Diagnosis P. nepthys is an extremely spinose, large Phyl- lurus (maximum SVL 103 mm). A ‘peppered’ brown venter separates it from all its congeners (with immaculate or mottled venters). P. nepthys is further distinguished from P. caudiannulatus by its rostral groove (partially divided. Fig. 7e vs completely divided. Fig. 7a); from P. isis and P. platurus by the depth of its acarodomatia = axil- lary pits (moderately to deeply invagi nated vs shallowly invaginated); and from P. ossa by rostral grooves (1 groove partially dividing the rostral, Fig. 7e vs 1-3 grooves, usually 3, only rarely 1 or 2, partially dividing the rostral. Fig. 7f-h). Distribution Confined to the Clarke Ra., west of Mackay, MEQ. Much of this range is included in the Eungella NP (Fig. 26). Habitat Complex notophyll and mesophyll vine forests on a variety of substrates, including granite. Habits Commonly found in association with both rocks and trees, and also with buildings. Reproduction Gravid females (J53332, J53362) were present in the population during late October-late November, 1991. Diet The gut of QM J5 1 098 contained a beetle larva (Coleoptera). QM J5 1099 contained fragments of a fulgoroid leafhopper. QM J51100 contained cockroach and moth fragments (Blattodea and Lepidoptera). Etymology Selected from Egyptian mythology (Macken- zie, 1978). The epithet is to be treated as a noun in apposition. 116 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 25. P. nepthys, Eungella, MEQ (T. Helden). Description SVL- (mm); 39-103 (n98, mean 91.0). Propor- tions as % SVL: LI 41-49 (n90, mean 43.7); L2 49-60 (n88, mean - 54.3); T 61-89 (n49, mean 75.5); TT 28-50 (n49, mean 40.9); HL 26-31 (n97, mean 27.9); HW 20-24 (n97, mean 21 .5); S 9-13 (n97, mean 11.8); NL 14-24 (n96, mean 18.9). Head: large, depressed, triangular, distinct from neck; covered in small granules which are inter- mixed with larger conical tubercles (tubercules extremely prominent on snout); skin of head co- ossified with skull; deep vertical groove partially dividing rostral scale, Fig. 7e; rostral excluded from nostril; ear opening elliptical, vertical, much less than half as large as eye; supralabials 12-20 (nl96, mean 15.4); infralabials 10-17 (nl95, mean 13.9). Neck: broad. Body: moderate, depressed, covered in small granules; dorsal granules intermixed with larger conical tubercles; tubercles moderate on back, large on flanks and extremely prominent on sides of neck; basal scales surrounding flank tubercles only slightly enlarged; no enlarged tubercles or granules on ventral surface of body. Preanal pores: absent. Axilla: moderately to deeply invaginated. Limbs: long, covered in large pointed tubercles dorsally; without enlarged tubercles on ventral surface; digits strong, compressed distally; subdigital lamellae (fourth toe) 18-24 (nl73, mean 20.5). Original tail: (n49) - depressed, flared to carrot- shaped, contracted at base and attenuated at tip; covered with prominent enlarged spinose tubercles over entire dorsal surface; 6-8 rows of enlarged spines across the attenuated portion of the tail; attenuated tip accounts for 45-62% of total tail length; ventral surface smooth, grooved or ungrooved along midline (excluding at- tenuated tip). Regenerated tail: (n22) depre.ssed, broad and strongly flared, contracted at base and attenuated at tip; covered with spinose tubercles which are most prominent around the edges and on the attenuated tip; ventral surface without groove along midline. Skeletal Features (X rays) QM J32669, J32674-5, J34057, J5333I, J53359, J53362; (alizarin stained) J34024. Supraocular portion of frontal flat (n7); anterior process of interclavicle distinct - Fig. 2g; REVIEW OF LEAF-TAILED GECKOS II7 FIG. 26. Distribution of P. nepthys. epipubic cartilage slightly expanded - Fig. Ig; presacral vertebrae 26 (n7); sacral vertebrae 2 (n7); lumbar vertebrae 2 (n7); 1st autotomy sep- tum 5 (n3); abdominal vertebrae bearing reduced ribs 4; rib free cervicals 3 (n7); sternal ribs 2; mesostemal ribs 3. Pattern In spirit, dorsal base colour tan to dark brown; mark^ with irregular dark brown blotches on head, body and limbs. Body and limbs ventrally cream, faintly to heavily p)eppered with brown specks; labials cream, mottled with dark brown. Original tail dorsally tan to dark brown, marked with irregular dark brown blotches; four or five bold cream bands on tail, only those on the attenuated portion extend- ing to the ventral surface; some specimens exhibit faint, obscure banding ^tween the bold cream bands; ventrally cream, peppered with brown specks. Regenerated tail totally lacking cream bands; dorsally tan to brown, mottled with cream and dark brown blotches; ventrally mottled but paler than dorsal surface. Phyllurus ossa sp. nov. (Figs Ih; 2h; 7a,e,f-h,i,j; 27; 28; 29) Material Examined Holotype: Queensland Museum: J53444 6 (with original tail) Mt Ossa/Ossa Ck, via Mirani (20°56’, 148°49’) MEQ, PJ. Couper, D. Limpus, M. Cunnin- gham, 20 October 1991. Paratypes: Queensland Museum: J56775, J5679I-2, Vine Ck, Mt Dryander (20°15’, 148°33’) MEQ; J56766-71, Brandy Ck, Con- way Ra. (20^21’, 148°4r).MEQ; J56772-4, near Lit- tle Conway Mtn, Conway SF (20°27’, 148°44’), J53443, J53445-7, J53507 Mt Ossa/Ossa Ck, via Mirani (20^56’, 148°49’) MEQ: J53426-8 St Helen’s Gap, via Mt Charlton (2 ! °00’ , 148°43’ ) MEQ; J53389- 90, J53391 (alizarin preparation), J53392-3, J53414, J56311, Mt Charlton foothills, via Mt Charlton (21°0r, 148°44’)MEQ. Diagnosis P. ossa is a moderately-spinose, medium-sized Phyllurus (maximum SVL 89mm). P. ossa may be distinguished from P. caudiannulatus by its rostral shield (rostral partially divided. Fig. 7f-h vs rostral totally divided, Fig. 7a); from P. isis, P. nepthys, and P. platurus by its rostral grooves (usually 3 grooves, sometimes 2, rarely 1 , partial- ly dividing the rostral, Fig. 7f-h vs a single groove partially dividing the rostral. Fig. 7e,i,j). It can be distinguished further from P. nepthys by its venter (immaculate vs heavily ‘peppered’ brown); from P. platurus by its original tail (with distinct white bands vs lacking white bands); and from P. isis by the size of its flank tubercules (moderate vs small). Distribution Disjunct populations occur in the Mt Ossa/Mt Charlton area, (2! W, 148°43’) north of Mack- ay, MEQ, in the Conway Ra., (20°27’, 148M4’) and Mt. Dryander, (20°15’, 148° 33’E) via Proserpine, MEQ (Fig. 28). Habitat Complex notophyll vine forest on a variety of substrates. Habits Usually found on rocks in rainforest or on the trunks of trees that are near rock. At Mt Dryander, P. ossa was found on large boulders in a dry creek bed. Activity starts soon after dark. This species appears to be common. Reproduction Gravid females were present in the population in October, 1992. Diet The gut of QM J53391 contained spider frag- ments. QM J56768 also contained spider frag- ments, and pieces of a cricket (Gryllidae). 118 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 27. F. ossa. Ossa Ck, MEQ (P. Couper). Etymology Named for Mt Ossa, the type locality. The epithet is to be treated as a noun in apposition. Description S VL (mm): 36-89 (n28, mean 72.9). Adult S VL (mm): 63-89 (n22, mean 78.4). Proportions as % SVL: LI 41-46 (n2I, mean 43.2); L2 48-59 (n22, mean 53.6); T 79-93 (n8, mean 86.1); TT 39-53 (n7, mean 48.3); HL 28-32 (n28, mean 29.7); HW 23-26 (n28, mean 24.3); S 11-13 (n28, mean 12.2); NL 14-23 (n28, mean 18.6). Head: large, depressed, triangular, distinct from neck; covered in small granules which are inter- mixed with larger conical tubercles; skin of head co-ossified with skull : three deep vertical grooves penetrate dorsal edge of rostral scale (n22), Fig. 7f, two vertical grooves penetrating rostral .scale (n5), Fig. 7g; a single inverted ‘Y’ shaped groove penetrating the rostral scale (nl) (Fig. 7h); rostral excluded from nostril; ear opening elliptical, ver- tical, much less than half as large as eye; supralabials 13-18 (n56, mean 15.6); infralabials 13-16 (n56, mean 14.3). Neck: broad. Body: moderate, depressed, covered in small dorsal granules intermixed with larger conical tubercles; small on back, moderate on flanks, most prominent on sides of neck; basal scales sur- rounding flank tubercles slightly enlarged; ventral surface with or without a small patch of enlarged granules on the pectoral region. Preanal pores: absent. Axilla: shallowly to extremely deeply invaginated (Fig. 29). Limbs: long, covered in moderate-sized pointed tubercles dor- sally; proximal portion of forelimb bearing en- larged tubercles on ventral surface; usually a few enlarged granules on ventral surface of thigh; digits strong, compressed distally; subdigital lamellae (fourth toe) 16-20 (n40, meanl8.2). Original tail: (n9) depressed, broad and flared to carrot-shaped, contracted at base and attenuated at lip; with or without an enlarged series of spinose edge tubercles on the anterior Bared por- tion; covered dorsally with numerous moderate- sized, spinose tubercles which become smaller along the vertebral line; tubercles terminate ap- proximately half-way along the attenuated tip which is long, fine and tenninates with a minute REVIEW OF LEAF-TAILED GECKOS 119 FIG. 28. Distribution of P. ossa. rounded ‘knob’ distally; dorsal tubercles fail to form clearly defined rows anteriorly across the attenuated tip; attenuated tip accounts for 50-60% of total tail length; ventral surface smooth, lack- ing a midline longitudinal groove. Regenerated tail: (n 17) depressed, broad and flared, contracted at base and attenuated at tip; lacking spinose tubercles on dorsal surface and edges. Skeletal Features (X rays) J53390, J53392, J53426-8, J53443-5, J53447, J56766-8, J56770-4; (alizarin stained) J53391. Supraocular portion of frontal flat (n 17); anterior process of interclavicle pronounced - Fig. 2h (n 1 ); epipubic cartilage not expanded - Fig. 1 h (nl); presacral vertebrae 26 (nl7); sacral ver- tebrae 2 (nl7); lumbar vertebrae 2 (nl7); 1st autotomy septum 5 (nl2); abdominal vertebrae bearing reduced ribs 4 (nl); rib free cervicals 3 (n9); sternal ribs 3 (nl); mesostemal ribs 2 (nl). Pattern In spirit, dorsal base colour grey to pale brown; heavily marked with distinct, dark grey to black blotches on head, body and limbs; vertebral zone free from darker blotchings; back with or without obscure pale blotches. Body cream ventrally, slightly translucent. Labials pale grey to brovv'n, mottled with white. Original tail dorsally mid- grey, bearing dark grey blotches; attenuated tip black, with three to four distinct creamish bands which extend to the ventral surface; two prominent broken cream bands across the flared anterior portion of the tail (a faint, very obscure band sometimes lies between these two anterior bands); ventrally cream, faintly to boldly mottled with brown. Regenerated tail totally lacking cream bands; cream to pale grey with obscure black blotching or black with cream or tan blotch- ing; ventrally mottled, but paler than dorsal sur- face. Remarks Populations of P. ossa from Conway Ra., MEQ are smaller than the Mt Ossa/Mt Charlton/St Helen’s Gap, MEQ populations (adult SVL mm 63-80, n II, mean 73.7 vs SVL mm 70-89, n 11, mean 83.1). These populations also differ in the depth of their axillary pits. The axilla is extremely deeply invaginaled, Fig. 29 (Conway Ra.) vs axil- la shallowly invaginaled (Mt Ossa/Mt Charl- ton/St Helen’s Gap). FIG. 29. P. ossa from Conway Ra., MEQ, displaying deep axillary pits, acarodomatia (P. Couper). 120 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 30. P. platurus, Heathcote, NSW. (S. Wilson). Phyllunis platurus (Shaw, 1790) (Figs 7i,j; 30; 31) See Bauer (1990). Material Examined Queensland Museum: J56880-I, J56895, Marara, via Gosford (33^24’, 151^21*) MENSW. Australian Museum: R49185, 28km W, 2km S of Sin- gleton (32^34’, 150®5r) NSW; R6I097, Watagan Ra. (32°57’ , 1 5 1 ° 1 4’) NSW; R55807, Glen Davis (33°08\ 150°17’) NSW; R106495, Barrenjoey Head (33°35M5r200 NSW; R106609, lOkm WNW Mt Ku-ring-gai (33°39M51°02') NSW; RI0649I, Mc- Carrs Ck, Ku-ring-gai NP (33°40, 151 °1 5’) NSW; R107089, Hornsby, Sydney (33M2’, 15r06’) NSW; R10680L Gordon, Sydney (33®45\ 151 W) NSW; R55803, Northbridge, Sydney (33M8\ 151“! T) NSW; R106497, Dobroyd Head (33M9\ 151^16’) NSW ; R55808, Heathcote, Sydney (34°05’, 151^01) NSW. The type description, expanded description, and habitat and distribution data of Covacevich (1975) and the rediagnosis of P, platurus (Bauer, 1990) need no elaboration. The following new data on P. platurus are based on examination of a sample of 18 specimens from the collections of the Australian and Queensland Museums. QM J9054 from Buladelah, was treated as P. platurus by Covacevich (1975), but has been identified as Saltuarius swaini here. Reproduction Two females from Marara, MENSW each produced a clutch of two oval shaped eggs in late December, 1992. One egg from each clutch was preserved. The second was incubated in ver- miculite. The eggs measured 25.03 x 15.85mm, 23.26 X 15.30mm, 20.92 x 15.48mm and 21.83 x 15,28mm. Incubation took 71-72 days at an un- controlled Brisbane temperature. Hatchlings measured (SVL) 30.28mm and 3L48mm. Description The following measurements and characters have not been included in previous descriptions of P. platurus. Proportions as % SVL: LI 39-45 (nl8, mean 42.9); L2 50-58 (nl8, mean 54.0); S 12-14 (nl8, mean 13.1); NL 15-21 (nl8, mean 18.4). Rostral only partially divided, by an inverted ‘ Y’ shaped groove, Fig. 7i (n8), by a single verti- REVIEW OF LEAF-TAILED GECKOS 121 FIG. 31. Distribution of P. platurus. cal line (n8), or by an ‘L’ shaped groove, Fig. 7j (n2). Axilla shallowly invaginated. For skeletal details see Bauer, 1 990. P. platurus stands separate from its congeners in two skeletal features - presacral vertebrae (25 vs 26) and first autotomy septum (6 vs 5). ZOOGEOGRAPHY The species of Saltuarius and Phyllurus are associated with rainforest or healhlands, or both. (5. occultus and S. cornutus are obligatory rain- forest species; 5. salebrosus and S. swaini occur in both rainforests and heathlands. P. ossa, P. nepthys, P. isis and P. caudiannulatus are obligatory rainforest species; P. platurus is a species of heathlands). These associations may reflect the parallel history of the evolution of rainforests and healhlands in Australia. The former evolved on fertile soils, the latter on infer- tile soils. Each habitat was widely distributed prior the Tertiary (65mya). Tertiary climatic fluc- tuations account for alternating contractions and fluctuations in their size and for their occurrence as fragmented, coastal and near-coastal narrow strips. Their island-like occurrence in an ocean of sclerophyllous forests is well documented (Kik- kawa et al., 1979). With this history of alternating expansion/linking and contraction/separating rainforests and heathlands, it is not surprising that many taxa, some of which have narrow ranges and all of which are endemic to Australia, are ‘shared’ between rainforest and heathland today. The recognition of two additional (S. occultus and S. swaini) species from the formerly very widely distributed Phyllurus sensu lato (now Sal- tuarius) cornutus, and of three new species of Phyllurus (P. ossa, P. nepthys and P. isis) from a very small area of isolated peaks/ranges in mideastern Queensland is consistent with con- cepts of separation and isolation of rainforests, and the survival, during such drier periods, of relict patches of rainforest and their faunas. S. occultus is a narrowly endemic species, known only from the type series of four specimens, from rainforests of the Mcllwraith Ra., FNEQ. Its dis- covery is interesting zoogeographically. P. occul- tus appears to be one of only five vertebrate species which are confined to the rainforests of Iron-McIlwrailh Ranges. (The others are An- techinus leo Van Dyck 1980, Cophixalis crepi- tans Zweifel, 1985, Cophixalis peninsularis Zweifel, 1985 and Litoria longirostris Tyler & Davies, 1977). The situation where 1/13 rainforest reptile species is endemic to the area (Iron-Mcll wraith Ranges) contrasts sharply with that of the rain- forest block at the southern part of Cape York Peninsula (the Wet Tropics, between Cooktown, 15°48’, 145°15’, and Mt. Spec, I8°57’, 146°ir, NEQ). Here 20/29 rainforest reptile species are endemic. This pattern of Iron-Mcllwraith Ranges endemic-species-paucity vs endemic-species- richness in the Wei Tropics is not unique to the reptiles, mammals and frogs. Exact parallels have been observed in heteropodid spiders (V. Davies pers. comm.); land snails (J. Stanisic, pers. comm.); and in carabid and aradid insects (Dar- lington, 1961; G. Monteith, pers. comm.). S. cornutus sensu striclo is an addition to the already long list of rainforest reptile species en- demic to the Wet Tropics (e.g. Covacevich & McDonald, 1991 a, b). This area supports the highest percentage of endemic species known in Australia. A similarly high degree of endemism has been reported also in mammals, birds, frogs, and many invertebrate groups including snails, insects, spiders, crustaceans. S. salebrosus and S. swaini are found in a wide variety of rainforests and heathlands. The former is common from dense, moist, notophyll rain- forests to drier semievergreen vine thickets and heathlands, on either granites or sandstones. S. 122 MEMOIRS OF THE QUEENSLAND MUSEUM swaini occurs in notophyll rainforests and heath- lands. No other vertebrate species share the exact ranges of these two species, although many rep- tile (and other) species in coastal centr^ and southern Queensland and northern and central New South Wales range between fragments of rainforest and heathland (eg. Saiphos equalis, Hoplocephalus stephensii, Tropidechis carinat- us, Ophioscincus ophioscincus). The distribution of Saltuarius spp. is paralleled by that of Antechinus spp. (Marsupialia: Dasy- uridae) with only minor variations. Antechinus leo has roughly the same distribution as Sal- tuarius occultus; A. godmani / S. cornutus; A. flavipes / S. salebrosus; A. stuartii /A. swaini (S. Van Dyck, pers, comm.). With the separation of Saltuarius from Phyl- lurus sensu lalo, Phyllurus spp. are confined to the area between mideastem Queensland and mideastem New South Wales. They, loo, occur in rainforests (P. isis, P. nepthys, P. caudiannulat- us) and heathlands (P. platurus) and their dis- tributions undoubtedly reflect the parallel evolution of these habitats in eastern Australia. In the Mackay/Proserpine area of mideastern Queensland, three species of Phyllurus occur in three localities, all within 100km of each other (Mt Blackwood-Mt Jukes / P. isis sp. nov.; Eun- gella = Clarke Ra./ P. nepthys sp. nov.; Mt Charl- ton, St Helenas Gap, Mt Ossa, Conway / P. ossa sp. nov.). Such speciation in a small area appears unusual initially, but can be explained in terms of alternate contraction and expansion of rain- forests; the geology of the area; and the probable antiquity of Phyllurus. Given the multiple chromosome reduction in these species it is also possible that chromosome changes have con- tributed to speciation events (Sites & Moritz, 1987). Rozefelds ( 1 990) has described Oligocene rain- forest plant fossils from near Capella, MEQ to the SW of the Phyllurus localities. This area is presently extremely dry in comparison with them, and does not now support any vegetation remote- ly resembling rainforest. He notes of his site \.. the area was covered in tropical rainforest com- munities during the late Oligocene - early Miocene .... the reduction in and/or migration of taxa to refugial rainforest communities along the northeastern Queensland coast can be correlated with the increasing aridity of the Australian climate from the Miocene onwards.... Remnants of Australia's widespread early Tertiary mosaic of rainforest communities are now restricted to refugia along the eastern and northern coasts...’. The sequence of the many expansions and con- tractions of the rainforests has been ably sum- marised by Archer et al. (1991) who write ‘... the existence of many now isolated islands of rain- forest in Australia is evidence that these ... are remnants of what was once a much wider, more uniformly spread rainforest. Of the islands that remain, those of the tropical and temperate regions are clearly descendants of the ancient, primordial types of rainforest ...’. Genetic studies of reptile species in the Wet Tropics rainforest have revealed major differences attributed to con- tractions of rainforest in the past (Moritz et al., 1993). The present species composition of the relict patches of rainforest on Mt Blackwood-Mt Jukes and the occurrence in them of endemic species result from the geological and botanical history of the area. A quartz syenite intrusive (presently Mt Blackwood) formed in the Oligocene (25^0 mya). A second intrusive of granite (presently Mt Jukes) formed, later, but also in the Oligocene. As weathering progressed, the two peaks have been exposed (Champion, 1984). Moist, sheltered pockets haveenabled rainforests which have been eliminated from intervening drier areas, to sur- vive. These small islands of rainforest (Mt Jukes- Mt Blackwood, St Helen’s Gap-Mt Ossa-Mt Charlton) and the large rainforest islands (Clarke and Conway Ranges) can be viewed as a microcosm of the broader, total present picture of Australia’s tropical and subtropical rainforests with their highly endemic, narrowly distributed faunas and floras. Geckos are poorly known in Australia’s fossil record. However, they are present in Oligocene- Miocene deposits (Archer et al., 1991) when rain- forest was the dominant vegetation. Phyllurus has not been identified from fossil deposits, but Car- phodactylini (Phyllurus, Saltuarius and Car- phodactylus) which are, today, the only obligatory rainforest geckos in Australia, are regarded as a probable part of the Gondwanic fauna (Covacevich & McDonald, 1991a; Kluge, 1967). An ancestral Phyllurus may have occurred widely in the rainforests of the Oligocene- Miocene. Descendants of that ancestral form may have speciated in the ‘islands’ of rainforest present today. CONSERVATION STATUS McDonald et al. 1991 reviewed the conserva- tion status of all Queensland’s reptile species. With this review, the following additions to their REVIEW OF LEAF-TAILED GECKOS 123 list of ‘rated’ species are necessary - .S', occultus 2 (‘... species with a very restricted distribution in Australia and with a maximum geographic dis- tribution of less than lOOkm...’) K (‘... poorly known species P. caudiannulatus, P. isis, P. nepthys and P. ossa 2R (R, ‘species which are rare in Australia .... may be represented by a relatively large population in a relatively restricted area or by smaller populations spread over a wider range ACKNOWLEDGEMENTS This work was funded partly by a grant from the National Rainforest Conservation Programme, administered by the Department of Environment and Heritage (DEH). We are grate- ful for assistance in the preparation of this paper from; Alan Beckman, Simon Blomberg, Shamus Conway, Kate Couper, Bruce Cowell, Gary Cranitch, Michael Cunningham, Valerie Davies, Lynette Dickfos, Jenny Faulkner, Anita Heideman, T. Helden, Barry Heinrich, Harry Hines, Glen Ingram, Lauren Keim, Duncan Lim- pus, Tim Low, Jennifer Mahoney, Keith Mc- Donald, Geoff Monteith, Mary Mulcahy, Bob Murphy, David Nebauer, Simon and Jenny Ormsby, Steve Pearson, Steve Phillips, Mark Read, Gary Rees, Kirra Reynolds, Steve Richards, Janet Russell, Trent Russell, Ross Sad- lier, Paul Stumkat, Peter Thompson, Paul Web- ber, John Weigel, Steve Wilson, Peta Woodgate and Jeff Wright. Associate Professor Aaron Bauer constructively criticized this work. LITERATURE CITED ARNOLD, E.N. 1986. Mite pockets of lizards, a pos- sible means of reducing damage by ectoparasites. Biological Journal of the Linnean Society. 29: 1 - 21 . ARCHER, M., HAND, S.J. & GODTHELP, H. 1991. ‘Riversleigh. The story of animals in ancient rainforests of inland Australia’. (Reed Books: Sydney). 264 pp. BARRET, C. 1950. ‘Reptiles of AuslraIia’(Cas- sell: London). BAUER, A. M., 1990. Phylogenetic systematics and biogeography of the Carphodactylini (Rcptilia: Gekkonidae). Bonner Zoologische Monograph- ien30: 1-217. CHAMPION, D.C. 1984. ‘Geology and geochemistry of the Mt Jukes inlnisive complex'. (Unpublished Honours Thesis, James Cook University of North Queensland, Townsville). COGGER, H.G. 1992. ‘Reptiles and amphibians of Australia’. 5lh ed. (Reed: Sydney). 775pp. COGGER, H.G., CAMERON, E.E. & COGGER, H.M. 1983. ‘Amphibia and reptilia. Vol. 1. Zoological Catalogue of Australia’. (Australian Government Printing Service: Canberra). 313pp. COVACEVICH, J., 1 975. A review of the genus Phyl- lurus (Lacertilia: Gekkonidae). Memoirs of the Queensland Museum. 17(2): 293-303. COVACEVICH, J. & INGRAM, G. 1980. The en- demic frogs and reptiles of Cape York Peninsula. Pp 49-57. In Stevens, N.C. & Bailey, A. (eds). ‘Contemporary Cape York Peninsula’, (Royal Society of Queensland: Brisbane). 100pp. COVACEVICH, J. & MCDONALD, K.R. 1991a. Frogs and reptiles of tropical and subtropical eastern Australian rainforests: distribution pat- terns and conservation. Pp 281-309. In Werren, G. & Kershaw, P. (eds), ‘The rainforest legacy. Australian national rainforests study’, Vol. 2. Special Australian Heritage Publication Series. (Australian Government Publishing Service: Canberra). 414 pp. 1991b. Reptiles. In Nix, H.A.& Switzer, M.A. (eds), ‘Rainforest animals. Allas of vertebrates endemic to Australia’s Wet Tropics’. Kowari 1: 69 (Australian National Parks & Wildlife Service: Canberra), DARLINGTON, P.J. 1961. Australian carabid beetles V. Transition of wet forest faunas from New Guinea to Tasmania. Psyche 68(1): 1-24. GUNTHER, A. 1897. Descriptions of new species of lizards and a tree frog from north-eastern Queensland. Novitates 2Ux)logica 4:403-406. KIKKAWA, J., INGRAM, G.J. & DWYER, P.D. 1979. The vertebrate fauna of Australian heath- lands - an evolutionary perspective. Pp 281-299. In Specht, R.L. (ed.) ‘^osystems of the world. 9A. Heathlands and related shrublands. Descrip- tive studies’. (Elsevier Scientific Publishing Co.: Amsterdam). 497pp. KING, 1987. Chromosomal evolution in the Diplodac- lylinae (Gekkonidae: Reptilia). I. Evolutionary relationships and patterns of change. Australian Journal of Zoology 35: 507-531. KLUGE, A.G. 1967. Systematic.s, phylogeny and zoogeography of the lizard genus Diplodactylus Gray (Gekkonidae). Australian Journal of Zool- ogy 15: 1007-1108. LOVERIDGE, A., 1951. On reptiles and amphibians from Tanganyika Territory, collected by C.J.P. lonides. Bulletin of the Museum of Comparative Zoology at Harvard College. 106: 177-204. MCDONALD, K.R., COVACEVICH, J.A., IN- GRAM, G.J. & COUPER, P.J. 1991. The status 124 MEMOIRS OF THE QUEENSLAND MUSEUM of frogs and reptiles. Pp 338-345. In Ingram, GJ. & Raven, R.J. (eds), ‘An Atlas of Queensland’s Frogs Reptiles, Birds and Mammals’. (Board of Trustees, Queensland Museum ; Brisbane). 391pp. McKenzie, D.A. 1978. ‘Egyptian myth and legend’. (Bell Publishing Co: New York). 404pp. MORITZ, C. 1987. Parthenogenesis in the tropical gecko, Nactus arnouxii. Evolution 41: 1252- 1266. MORITZ, C., JOSEPH, L. & ADAMS, M. 1 993. Cryp- tic diversity in an endemic rainforest skink {Gnypetoscincus queenslandiae) Biodiversity and Conservation 2:412-425. OGILBY, J.D. 1892. Descriptions of three new Australian lizards. Records of the Australian Museum 2:6-11. RIDGWAY, R. 1912. ‘Color standards and color nomenclature’. (The author: Washington D.C.) 43 pp., 53 pis. ROZEFELDS, A. 1990. A new mid-Tertiary rainforest flora from central Queensland. Third Internation- al Symposium of Palaeobotany, Melbourne, 1988. pp 123-36. RUSSELL, A.P. 1980 Underwoodisaurus Wermuth 1965, a junior synonym of Phyllurus Schinz 1822. Journal of Herpetology 14: 415-16. SITES, J.W. & MORITZ, C. 1987. Chromosomal evolution and speciation revisited. Systematic Zoology 36(2): 153-174. S WOFFORD, D.L. 1991. Phylogenetic analysis using parismony. (Illinois Natural History Survey: Champagne Urbana, Illinois). WELLS, R.W. & WELLINGTON, R.C. 1985. A clas- sification of the Amphibia and Reptilia of Australia. Australian Journal of Herpetology, Supplementary Series no. 1., 1-61. (Australian Biological Services: Sydney). 64pp. ECOLOGICAL NOTES ON CARLIA ROSTRALIS IN RAINFOREST AND ASSOCIATED HABITAT IN THE SOUTHERN WET TROPICS J.M. WHITTIER Whittier, J.M. 1993 12 24: Ecolo^cal notes on Carlia rostralis in rainforest and associated habitat in the southern Wet Tropics. Memoirs of the Queensland Museum 34(1): 125-129. Brisbane. ISSN 0079-8835. A study of Carlia rostralis was conducted in November-December, 1988, July, 1990-92, and January-February, 1991-93, between Townsville and Cairns, Queensland, Australia. In this southern portion of its range, C rostralis was locally abundant in vine tickets along creeks as well as in grassy riparian areas in open forest. It was absent from dense rainforest at higher elevations in the Seaview Ra. C. rostralis was restricted to the more mesic eastern localities in this southern area. Current and future conservation status of this species is excellent. Reproductive patterns of C. rostralis were investigated at two locations. Waterfall Ck, Watervicw Shire, and Waterview Ck, Jourama Falls NP. Adult males and females (> 45.0 mm snout vent length) were highly sexually dimorphic in both colour and maximal body size. Juveniles in their first year resembled females in colouration. Reproduction in C. rostralis is seasonal; egg-laying commenced in November-December and continued to February in the study. In each year, nearly all adult females in the populations observed were reproductively active. Reproductive activity coincided with the usual lime of the wet sea.son at the two localities, but seasonal rainfall did not appear to regulate the onset or the maintenance of egg-laying. □ Lizard ecology, lizard reproduction, seasonal breeding, Scincidae. JoanM. Whittier, Department of Anatomical Sciences, University of Queensland, St. Lucia, Queensland 4072, Australia, W September, 1993. Recent reviews of reproductive patterns in lizards that inhabit tropical environments have found that there is little relationship between seasonal environmental conditions and timing of breeding (Shine, 1985; James & Shine, 1985; Auffenberg & Auffenberg, 1989; Vitt, 1990). There appears to be a lack of general patterns of breeding in tropical lizards, despite early predic- tions that breeding in these areas would probably be related to seasonal rainfall. The factors that regulate the diverse patterns of breeding in tropi- cal lizards are virtually unknown (Whittier & Crews, 1987; Whittier, 1993). The genus Carlia represents an excellent model group in which patterns of seasonal breed- ing in tropical areas can be studied. Carlia com- prises a total of 20-*- recognised species endemic to tropical Australia, Papua New Guinea, and nearby islands (Ingram & Covacevich, 1989). No long-term studies of the status and reproductive biology of this group have been conducted pre- viously. A few short-term studies have examined the reproductive and thermal ecology of members of this genus. Reproductive cycles of C. rhom- boidalis and C schmeltzii (originally identified as Leiolopisma rhomboidalis and Leiolopisma fusca, respectively; identification checked by ex- amination of Wilhoft’s labelled specimens in the Los Angeles County Museum, by JMW), in northeastern Queensland, and C. bicarinata, in eastern Papua New Guinea, indicate that these species reproduce during the wet season (Wilhoft 1963a, 1963b; Wilhoft & Reiter, I965;Zugetal., 1982). Similarly, several species of Carlia were reported to breed only in the wet season in the Alligator Rivers region of the Northern Territory, Australia, based on examination of museum specimens (James & Shine, 1985). James & Shine (1985) concluded that taxa such as Carlia, with origins in seasonally wet tropical areas, have maintained a historical association between breeding and the occurrence of seasonal rainfall. However, as in other tropical lizards that breed in association with the seasonal onset of rain, the factors that regulate the timing of breeding are not known in Carlia. At the localities where C. rostralis were ob- served in the present study, a large community of reptiles exists at high densities. In this region, the onset and yearly occurrence of the wet season is relatively unpredictable. This is due to the stochastic occurrence of cyclonic rains; in more northern areas rain docs occur in most years, but in these more southern areas dry summers occur 126 MEMOIRS OF THE QUEENSLAND MUSEUM at irregular intervals, approximately eveiy three to six years. For lizards such as C. rostralis that live for one to five years (unpub. data, JMW), a dry summer can be a significant event in the life cycle. Furthermore, the onset of the wet season can vary up to four months, ranging from Novem- ber to February, although normally commencing in late December or January. In the Wet Tropics, the limited data available on reproductive patterns suggest that most species of lizards breed in the summer, with the onset of breeding commencing just prior to the start of the wet season (December & January; James & Shine, 1985). The conclusions from these observations have generally been that ‘wet season’ breeders reproduce in response to in- creased food availability (insect prey) that ac- companies summer rains. These observations suggest that food availability may not play a critical role in the onset of reproduction, but that it may be important for the maintenance of reproduction later in the season. I became interested in determining whether reproduction in C. rostralis in the Wet Tropics was regulated by seasonal rainfall. Of interest was the timing of reproduction in years when the wet season was delayed or did not occur (the latter is associated with the conditions of the Southern Pacific Oscillation, or El Nino). At the same time that the study of reproductive patterns in this species was conducted, the distribution and habitat preferences of C. rostralis were surveyed. Known localities obtained from records in the Queensland and Australian Museums (Ingram & Covacevich, 1989) between Townsville and Cairns, NEQ, were visited to establish and con- firm the current distribution of this species. METHODS A study of C. rostralis was conducted in November-December, 1988; July, 1990-92, & January-February, 1991-93 at two field sites. Waterfall Ck, Watervicw Shire, Ingham SF Dis- trict, and at Waterview Ck, Jourama Falls NP, both approximately 60 km north of Townsville, NEQ. Both sites supported lowland vine thicket extending down the water course from rainforest at higher elevations. The vine thicket riparian zone extended approximately 100m to either side of the creek. Surrounding these areas was open sclerophyll forest. Several additional sites be- tween Cairns and Townsville were visited to determine the extent of current distribution of C rostralis in the southern portion of the Wet Tropics. These sites were mainly limited to loca- tions in the vicinity of those published in Ingram & Covacevich (1989). To assess sex and reproductive condition animals were hand captured using insect lures. On capture, individuals were measured snout to vent ( ±0.5 mm) and weighed ( ±0.2 gram). At the two field sites individuds were toe-clipped for long term population studies. Adult males were distin- guished from adult females by the presence of a black gular region and bright orange lateral coloration. Minimum size (SVL) at sexual maturity was determined by the observation of the smallest size at which either sex was produc- ing functional gametes. This was indicated by palpation of females in the field and dissection of museum specimens (JMW, unpublished observa- tions). During the winter months, when immature and juvenile (< 35-40 mm SVL) individuals were present, coloration was not reliable as a means of determining sex, and individuals with a white throat less than 40 mm SVL were not sexed. Female reproductive condition was assessed by laparotomy or by palpation. Initially palpation t followed by laparotomy was used to confirm the j presence of previtellogenic follicles (no ovarian masses, < 3 mm in diameter), maturing follicles, j (round, firm bodies > 3 mm in diameter), ovulated eggs (oval soft bodies > 5 mm in length), or shelled oviductal eggs (oval hard bodies > 6 mm in length). Subsequently females were palpated only , as this proved to be an accurate method of assessing reproductive condition. Since some females began developing a subsequent clutch, a i portion of females had boSi maturing follicles and oviductal eggs. Most females in the study were recaptured numerous times before, during and after ovulation and after oviposition so that the timing and stage of the ovarian cycle was recon- firmed. RESULTS In the southern portion of its range, the current , distribution of C. rostralis appears to resemble recent historical distribution as indicated by \ records in the Queensland and Australian * Museums, and in Ingram & Covacevich (1989). At only a few sites were no C rostralis found. At the majority, animals were either common or abundant. C. rostralis was locally abundant in vine thickets along creeks as well as in grassy riparian areas in open forest. This species was absent from dense rainforest such as that at higher elevations in the Seaview Ra. C. rostralis was ECOLOGICAL NOTES ON CARLIA ROSTRALIS 127 restricted to the more mesic eastern localities in this southern area. No changes in habitat preference were noted between summer and winter seasons at the intensively studied localities. In the following list of C rostralis sites surveyed, 1988-1993, the data are: (location sur- veyed; lat/long; habitat; dale; status: A-abundant, C-common, U-uncommon, Ab-absent; AM/QM#, recorded locality). 1. Holloway Beach; 16 51/145 45; beach, residential; 7/90;Ab; AM R97693-4; Holloway Beach, via Casuarina St. 2. Davies Creek NP; 17 00/145 34; riparian open forest, rocks; 7/91; A; AM R53904; Davies Creek Rd., 16 miles SE Mareeba. 3. Palmerston NP; 17 36/145 45; rainforest; 12/88; Ab; 4. Millstream Falls NP; 17 39/145 27; grass in open forest, riparian; 12/88; C; AM R62272; Millstream Falls NP. 5. Murray Falls SF Park; 18 12/145 55; grass in open forest, riparian; 12/88; A; . 6. Wallaman Falls NP; 18 38/145 33; grass in open forest, riparian; 12/88; U; AM R97690-2; 24.1 km ESE Wallaman Falls NP by road. 7. Jourama Falls NP; 18 52/146 07; grass in open forest, riparian vine thicket; 12/88-7/93; A; — ; 8. Jourama Falls NP, upper drainage of east tributary of Waterview Ck; 1 8 52/14607 ; rainforest; 7/9 1 ; Ab; 9. Waterfall Ck, Waterview Shire; 1 8 53/146 09; grass in open forest, riparian vine thicket; 12/88-7/93; A; . 10. Waterfall Ck upper drainage, Waterview SF; 18 53/146 08; rainforest; 12/88; Ab; — . 11. Birthday Falls. Blackfriars SF; 18 59/14609; rain- forest; 12/88; Ab; — ; — . 12. Hencamp Ck northern tributaries, Hinch inbrook SF; 19 01/146 21; rocky dry stream beds, open forest; 12/88; C; QM J27695; QM 32570-32575; Hencamp Creek, 5km N of 1 km E of Rollingstone. 13. Hencamp Ck, Hinchinbrook SF; 19 01/146 21; rocky dry stream bed, vine thicket; 7/93; A; QM J27695; QM 32570-32575; Hencamp Ck, 5km N of I km E of Rollingstone. , 14. Little Crystal Ck, Mt Spec NP; 19 01/146 17; vine thicket, riparian; 12/88; U; AMR97675-6; lOkmW of Bruce Hwy via Paluma Rd. 15. McClellan’s Lookout, Mt Spec NP; 19 01/146 12; grass in open forest; 12/88; U; — ; — . 16. Cloudy Creek upper drainage , Mt. Spec NP; 19 01/146 12; rainforest; 12/88; Ab; -; 1 7 . Pine Ck tributaries, Blackfriars and Kangaroo Hills Shires; 19 01/146 07; dry streams, open forest; 12/88; Ab; 18. Station Ck, Clemant SF; 19 07/146 26; rocks in dry stream bed, vine thicket; 7/93; A; — ; — . There is a high degree of sexual dimorphism in size in C. rostralis. Data from the first and second breeding seasons, in November & December, 1988, and January & February, 1991, are repre- sentative. Males were significantly longer and heavier than females and were heavier per unit of body length, except for the largest females (n, mean ± ISE, of SVL and BW, of males: 21, 61.6 ± 3.5 mm, range, 53.0-69.0 mm; 5.4 ± 1,0 g, range, 4.0-7 .0; of females: 21, 56,0 ± 2.8 mm, range, 50.0-60.0 mm; 4.4 ± 0.9 g, range, 3, 0-7.0 g). At first breeding season, most of the adult females (12/14) were reproductively active during the study, with either preovulatory, post- ovulatory or, in one case, bo^ types of follicles present in the reproductive tract. All females ex- amined had two eggs. Two oviductal eggs, 8x4 mm, were found in two additional females that were kept as specimens. One of these individuals also had two yolking follicles, 1 mm in diameter, present in the ovary, indicating that females are capable of producing multiple clutches. Based on direct observation (1988) or indirect inference from the presence of cohorts of hatc- hlings (1991 & 1992), ovarian recrudescence and egg-laying in this species began in November in all three years of the study. Reproductive activity was maintained through February in at least two years, regardless of the timing of the onset of the rainy season (Table 1). In 1988/89, females began developing mature follicles in early November and the first clutch of eggs was laid in mid- November; hatchlings first appeared in mid- December, suggesting that incubation time under the ambient conditions was 4-5 weeks. The onset of summer rains in 1988/89 occurred earlier than usual, in mid-December. In the second breeding season examined, 1990/91, breeding again com- menced in November, as estimated by the presence of two cohorts of hatchlings in the population in mid-January. However, there was no onset of summer rains in this season, due to an occurrence of a severe drought as a result of a persistent Southern Pacific Oscillation. Finally, in the third year of the study, 1991/92, breeding commenced in November, estimated by the presence of three cohorts of hatchlings observed in late January, and continued through February. In mid-February, females ceased developing ma- ture follicles and the reproductive season came to an end. In this year drought conditions persisted through early January, and heavy wet season rains did not occur until mid-February. DISCUSSION The current and future conservation status of C. rostralis in the southern portion of its range in the 128 MEMOIRS OF THE QUEENSLAND MUSEUM Wet Tropics appears to be excellent. The habitat preferences and distributional requirements of the lizards in this area confirmed previous accounts (Ingram & Covacevich, 1989). C. rostralis is often observed basking on the rocky floor of the vine thicket. This heliothermy probably accounts for the species’ absence in denser rainforest at higher elevations in the Seaview Ra. C. rostralis has a relatively diverse range of habitat preferen- ces, being found also in grassy areas in open sclerophyll forest. However, the animals found in these localities usually are in close proximity to creeks and mesic riparian zones; in drier open forest on the western side of the range the species is absent. Reproductive observations of C. rostralis at two localities (W aterfall & Waterview Cks) indi- cated that this species attains a large index of sexual dimorphism (1.1; ratio of mean SVL of males to females; Stamps, 1983). In addition, C rostralis has a striking degree of sexual dichromatism that develops in the first year. These patterns of sexual dimorphism, although not uncommon amongst Carlia or other Australian scincids, are very different from that found in scincids that have been studied else- where, although few species have been studied (Stamps, 1983). This high degree of sexual di- morphism and dichromatism in C. rostralis ap- pears to be related to breeding, as males do not usually develop the black gular region and bright orange, black and white markings on the lateral head, neck and thorax until breeding commences in the first year (Whittier & Scott, 1 9^89; Whittier, 1991). These patterns of sexual dimorphism and dichromatism may be related to a highly competi- tive and ritualistic dominance hierarchy that is exhibited by male C. rostralis in both the laboratory and the field (Whittier & Martin, 1991). Reproduction in C rostralis is seasonal, with egg-laying beginning in November and Decem- ber, and continuing to February. Egg production and hatchlings were not observed in July. In each year, nearly all adult females in the populations observed were reproductive! y active, having yolking follicles or oviductal eggs. Multiple clutches appeared to be produced by most females; in nearly every case of a recapture in the summer, developing follicles were detected in females that had previously been gravid. From these observations, I suggest that female C. rostralis produce three to four clutches of two eggs each during each breeding season (Whittier & Scott, 1989). Although the breeding period in C. rostralis coincides with the usual time period of the wet season, breeding is not regulated by the onset of seasonal rain. Nor is breeding maintained by any factors such as purported increases in food availability associated with those rains. Although the factors that regulate timing of reproduction in this species are not known, two conclusions can be drawn from the study. First, that the liming of breeding is predictable and regulated in a precise manner, and second, that timing of breeding coin- cides with the most frequent occurrence of seasonal rain. I speculate that the timing of reproduction in this species is regulated either by seasonal changes in photoperiod or by an en- dogenous mechanism. These mechanisms would represent the best predictor of favourable condi- tions over the long term in this unpredictable environment. Many factors, including physiology, phylogeny, zoogeographic history and life his- tory traits may influence seasonal reproduction in squamate reptiles that inhabit subtropical and tropical environments (Shine, 1985; James & Shine, 1985; Auffenberg & Auffenberg, 1989; Vitt, 1990). From a physiological viewpoint, to understand the patterns of breeding in tropical lizard species and communities, an important consideration is the somatic condition of the female lizards (Bradshaw et al., 1991; Whittier, 1991a; 1993; Wiittier & Tokarz, 1992). Patterns of abundance of food resources in the environ- ment, assessed in the context of energy require- ments for subsistence, growth, and reproduction of the species need to be considered. Because reptiles, particularly squamales, are limited by the energetic cost of female egg production, the mechanisms by which females assess somatic condition can be co-opted to regulate reproduc- tion directly (Steams, 1976; Tinkle & Hadley, 1985). Future studies should determine how TABLE 1 . Female egg-laying patterns of C. rostralis, NEQ, in relation to the onset of the wet season. Time Onset of Rain Breeding Onset Breeding Duration Nov. 1988- Dec. 1988 mid-Dec. mid-Nov. unknown Jan. 1992- Feb. 1992 no season mid-Nov.* Feb. Jan. 1993- Fcb. 1993 Feb. mid-Nov.** Feb. ^Estimated from the presence of two cohorts of hatchlings ^j>resent in early January. Estimated from the presence of three cohorts of hatchlings present in late January. ECOLOGICAL NOTES ON CARLIA ROSTRALIS 129 species such as C. rostralis precisely time and maintain reproduction in relatively unpredictable environmentslike the southern portion of the Wet Tropics. ACKNOWLEDGEMENTS J. Scott, J. Martin, K. Verkest, N. Taylor, D. Stewart, D. Moeller, and C. Macrokanis provided enthusiastic field assistance. The Department of Environment and Heritage and Queensland Forest Service gave permission to collect and work in National Parks and State Forests. Por- tions of this study were supported by a Portland State University Faculty Development Grant, a University of Queensland Special Projects Grant, and Supplementary Grants from the Wet Tropics Management Agency. LITERATURE CITED AUFFENBERG, W. & AUFFENBERG, T. 1989. Reproductive patterns in sympatric Philippine skinks (Sauria: Scincidae). Bulletin of the Florida State Museum Biological Sciences 34: 201-247. BRADSHAW, S.D., SAINT GIRONS, H. & BRAD- SHAW, F. 1991. Patterns of breeding in two species of agamid lizards in the arid subtropical Pilbara region of Western Australia. General and Comparative Endocrinology 82: 407^24. INGRAM, G. & COV ACEVICH, J. 1 989. Revision of the genus Carlia (Reptilia, Scincidae) in Australia with comments on Carlia bicarinata of New Guinea. Memoirs of the Queensland Museum 27: 443-90. JAMES, C. & SHINE, R. 1985. The seasonal timing of reproduction: a tropical-temperate comparison in Australian lizards. Oecologia 67: 464-474. SHINE, R. 1985. The reproductive biology of Australian reptiles: A search for general patterns. Pp. 297-303. In Grigg, G.. Shine, R. & Ehmann, H. (eds), ‘Biology of Australasian frogs and reptiles'. (Royal Zoological Society of New South Wales: Sydney). STAMPS, J.A. 1983. Sexual selection, sexual dimor- phism, and territoriality. Pp. 169-204. In Huey, R.B., Pianka, E.R. & Schoener, T.W. (eds). ‘Lizard ecology studies of a model organism’. (Harvard Univ. Press: Cambridge, Mas- sachusetts). STEARNS, S.C. 1976. Life-history tactics: a review of the ideas. Quarterly Review ofBiology 51: 3-47. TINKLE, D.W. and HADLEY, N.F. 1985. Lizard reproductive effort: caloric estimates and com- ments on its evolution. Ecology 56: 427-34. Vnr, L.J. 1990. The influence of foraging mode and phylogeny on seasonality of tropical lizard reproduction. Papeis Avulsos de Zoologia (Sao Paulo) 37: 107-23. WHITTIER, J.M. 1991. Endocrine correlates of female territoriality in the Australian rainbow skink, Carlia rostralis. American Zoologist 3 1 : 2 A. 1991a. Artificial control of reproduction. Pp. 243- 268. In Pang, P., Schreibman, M. & Jones, R. (eds), ‘Vertebrate endocrinology, fundamentals and biomedical applications. Volume IV, Reproduction’. (Academic Press: New York). (in press). Seasonal reproduction in squamate rep- tiles: Endocrine patterns and variations. Proceed- ings of the XII International Conference on Comparative Endocrinology, National Publica- tions Board of Canada, Toronto, Ontario. WHITTIER, J. M. & CREWS, D. 1987. Seasonal reproduction: Patterns and control. Pp. 385-409. In Norris, D.O. & Jones, R.E. (eds), ‘Hormones and reproduction in fishes, amphibians and reptiles’. (Plenum Press: New York). WHITTIER, J. M. & SCOTT, J. M. 1989. Observa- tions on the reproductive biology of an oviparous skink (Carlia rostralis) in NE Queensland, Australia. Abstract, American Society Ich- thyologists and Herpetologists 1989: 161. WHITTIER, J.M. & MARTIN, J.E. 1992. Aspects of social behaviour and dominance in male rainbow skinks, Carlia rostralis. Australian Journal Zool- ogy 40: 73-9. WHITHER, J. M. & TOKARZ, R. R. 1992. Physiologi- cal regulation of sexual receptivity in female rep- tiles. Pp. 24-69. In Gans, C. & Crews, D. (eds), ‘Biology of the reptilia, Vol. 18, physiology E’. (University of Chicago Press: Chicago). WILHOFT, D.C. 1963a. Reproduction in the tropical Australian skink, Leiolopisma rhomboidalis. American Midland Naturalist 70: 442-61. 1963b, Gonadal histology and seasonal changes in the tropical Australian lizard Leiolopisma rhom- boidalis. Journal of Morphology 113: 185-204. WILHOFT, D.C. & REITER, E.C. 1965. Leiolopisma fuscum, a tropical Australian skink. Journal of Morphology 116: 379-88. ZUG, G.R., BARBER, M.M. & DUDLEY, J.C. 1982. Gonadal histology and reproduction in Carlia bicarinata (Scincidae, Sauria, Reptilia) of the Port Moresby area, Papua New Guinea. Her- petologica 38: 418-25. 130 MEMOIRS OF THE QUEENSLAND MUSEUM VARANUS PRASINVS (THE EMERALD GO ANNA) ON MOA ISLAND, TORRES STRAIT, AUSTRALIA. Memoirs of the Queensland Museum 34(1)130. 1993:- A specimen of Varanus prasimts (QM transparency NM 791, E. Mattock) was observed in closed mesic forest on Moa Island (10°irS 142®16*E) recently. This is the only record of this species in Australia this century, and is the southernmost record for the species. Moa Is., formerly Banks Is., is a relatively large continental island(700km approximately) in thecenlralpartof the Torres Strait. There are two small villages on the island Diverse native vegetation, most of which is undisturbed, predominates. At higher elevations on and around Moa Peak (374m), there is a closed mesic forest. This is now one of the few large patches of closed mesic forest in Queensland in which virtually no herpetological survey work has been done. Accompanied by Mr T. Moore. Ms E. Mattock and Mrs A. Torenvwek, we climbed Moa Peak on 28 August, 1993. At about 180m, from a rocky outcrop, we saw a specimen of V. prasinus basking in bright sunlight on a broken tree top, about 10m from the ground. The specimen was bright green, with black chevrons along its back. It was slender and had a snout- vent length of approximately 30cm. Its long, slender tail was curled, not used to hold on to the tree. V. prasinus is well known in Papua New Guinea, but is recorded in Australia only from Dauan (formerly Cornwallis) and Mer (formerly Murray) Islands in the Torres Strait (Gunther, 1877, 1879; Boulenger, 1885). These two islands are in the northern and eastern portions of the Torres^lrait. Cogger (1992) reports V. prasinus in the far northern Torres Strait and Papua New Guinea. There are no recent records of V. prasinus in Australia. No Australian specimens are held in the collections of the Queensland or Australian Museums (Covacevich & Couper, 1991; R. Sadlier pers. comhi.). Records of V. prasinus from the Mcllwraith Range area of Cape York, reported by Czechura ( 1 980), have been shown to be based on specimens of V. teriae (Sprackland, 1991). Sprackland (1991) also described a new species of tree goan- na, V. telenestes, from Rossell Island, Papua New Guinea, from material previously treated as V. prasinus. This raises the possibility that the specimen we observed on Moa Island may also be an endemic island taxon. Future collection and taxonomic evaluation of the green goanna on Moa Island will clarify this. This observation emphasises the importance of the closed mesic forest on Moa Island. Acknowledgments We thank the Torres Strait Inter Island Community Council, Mr O. Bosun of the Kubin Village Community Council, and Mr W. Pedro of the St Paul’s Village Community Council for permission to wewk on Moa Island, and for assistance with transport on the island; Mr T. Moore for accommodation; and the University of Queensland for the Special Studies Al- lowance to JMW. Literature cited Boulenger, G.S. 1885. ‘Catalogue of the lizards in the British Museum (Natural History)’. Vol. 2 (British Museum Natural History: London). 497pp. Covacevich, J.A. & Coup)er, P.J. 1991. The reptile records. Pp 45-140. In Ingram, G.J. & Raven, R.J. (eds) ‘An atlas of Queensland’s frogs, reptiles, birds and mammals’. (Board of Trustees, Queensland Museum: Brisbane). 391pp. Cogger, H.G. 1992. ‘Reptiles and amphibians of Australia’. (Reed: Sydney). 775pp. Czechura, G.V. 1980. The emerald monitor Varanus prasinus (Schlegel): an addition to the Australian mainland her- petofauna. Memoirs of the Queensland Museum 20: 103-109. Gunther, A. 1877. Description of three new species of lizards from islands of Torres Straits. Annals and Magazine of Natural History (4) 19: 4I3-4I5. 1879. Notice of a collection of reptiles from islands of the Torres Straits. Annals and Magazine of Natural History (5) 3: 84-87. Sprackland, R.G. 1991. Taxonomic review of the Varanus prasinus group with description of two new species. Memoirs of the Queensland Museum 30: 561-576. J. M. Whittier & D.R. Moeller, Department of Anatomi- cal Sciences, University of Queensland, Queensland, 4072, Australia; 7 September, 1993. REPRODUCTIVE BIOLOGY OF THE PRICKLY FOREST SKINK, GNYPETOSCINCUS QUEENSLANDIAE, AN ENDEMIC SPECIES FROM NORTHERN QUEENSLAND MICHAEL CUNNINGHAM Cunningham, M. 1993 12 24: Reproductive biology of the Prickly Forest Skink, Gnypetos- ciVicM^ queenslandiaCy an endemic species from northern Queensland. Memoirs of the Queensland Museum 34(1):131-138. Brisbane. ISSN 0079-8835. Examination of morphology, reproductive state and skeletochronological markings in specimens of a rainforest skink, Gnypetoscincus queenslandiae, in the Queensland Museum, reveals a sex ratio of 1: 1.5 (2 mm diameter) ovarian follicles; III number of enlarged follicles, between 1 .5 mm and 2 mm in diameter; IV num- ber of ovarian follicles between 0.5 mm and 1 .5 mm in diameter; and V number of small follicles, less than 0.5 mm in diameter. These classes were deterntined from the ob- served size distribution of developing follicles, and correspond to the follicular growth stages described for the Diamond-backed Water Snake, Natrix rhombifera^ (Betz, 1963) and the latter five classes described for several tropical snakes (Zug, 1979). Oviducal eggs (class 1) were a golden yellow colour and always larger than 5 mm; class II follicles were a creamy yellow colour; classes III, IV and V were white. The diameter of each ovarian follicle was determined using a dissecting microscope. To assess the relationship between size and relative age in G. queenslandiae , the left femur from fourteen specimens (representative of the range of sampling locations and size within each sex), was removed for skeletochronological analysis. Each femur was decalcified in formic acid and formalin, and vacuum embedded in wax. Transverse sections were taken from the mid- diaphysis of these bones, mounted on slides and stained for 15 minutes with Delafield’s haematoxylin and for 10 seconds with Eosin. These were examined under a compound micro- scope, to record the number of haematoxylin stained concentric rings in the compact bone of each femur. ANALYSES A Chi square test was used to compare the observed sex ratio in the total sample, with an expected 1 : 1 ratio. Morphological differences be- tween sexes were assessed by Canonical-Dis- criminant Analysis. This analysis finds the maximum difference between groups (in this case males and females) based on a weighted com- bination of variable characters, measured on each individual. Canonical-Discriminant Analysis may demonstrate separation among groups that appear indistinguishable in analyses of single variables, due to overlap between groups in the range of each morphological character. A series of meristic measurements (SVL, head length, head width, length of the left tibia, and length of the left fourth toe) and scale counts (lamellae under the left and right fourth toes, left and right supra-ocular scales, para-vertebral scales, and nuchal scales) gathered on these skinks for another study were submitted to the procedure ‘CANDISC’ in the SAS statistical analysis pack- age (SAS Institute Inc, 1985). In addition to this multi-variate analysis ‘CANDISC’ provides analysis of variance (ANOVA) comparisons of groups for each morphological variable (SAS institute inc., 1 985). These test the hypothesis that differences between the sexes cannot be distin- guished from variation within each sex. Nix & Switzer (1991) report size variation in populations of G. queenslandiae from different locations. The SAS procedure ‘GLM’ (SAS in- stitute inc., 1985) was used to investigate size (SVL) differences between sampling sites by one-way ANOVA. This analysis included only those locations from which five or more samples had been collected. The mean size of gravid females (with oviducal eggs) was compared with that of all other females from the Atherton Table- land, using Student’s t-test, generated by the SAS ‘TTEST’ procedure (SAS institute inc., 1985). This analysis tests the null hypothesis that gravid females do not differ in size from other females. Pearson product-moment correlation coeffi- cients, generated by the SAS procedure ‘CORR’ (SAS instimte inc., 1985), were used to compare size of female skinks with number of eggs or ovarian follicles carried by those females, for each class defined above. These analyses test the hypothesis that the number of eggs and ovarian follicles is not correlated with SVL. A Chi square test of homogeneity was used to investigate tem- poral variation in the proportion of gravid females in the Atherton Tablelands samples, and to test the probability that this variation could be due to random sampling. Finally, the procedure ‘REG’ in SAS (SAS institute inc., 1985) was used to generate simple linear regressions of the number of femoral bone rings against size, SVL, for ail samples and for those from the Atherton Tableland alone. These regressions are used to predict the expected num- ber of femoral bone rings from a skink of known size. CLUTCH SIZE (I) ENLARGED FOLLICLES (III) SMALL FOLLICLES (V) REPRODUCTIVE BIOLOGY OF THE PRICKLY FOREST SKINK 133 FIG. l(A-E). Plot of clulchsizeandnumber of follicles in each size class against SVL for all females. Sex Ratio, Sexual Dimorphism and Size Variation In the sample there were 64(5 c5 and 94$ 9. Individuals ranged from 30 mm to 84 mm (SVL) (mean±sd: SVL=58.4±10.9 mm), but it was not possible to determine the sex of the two skinks smaller than 34 mm SVL. This gives a sex ratio of 1:1.47 which differs significantly from the expected 79:79 ratio (X^=5.70, df=l, p<0.02), indicating an overall bias towards females in this collection. The canonical discriminant analysis demonstrated significant separation between males and females in muiti-dimensional mor- phological space (Exact F=6.30, df=ll, p<0.0001). This separation was generally not reflected in the univariate tests of sexual dimor- phism, with only the number of nuchal scales (NTOT - defined here as the number of scales posterior to, and in contact with, the parietal 134 MEMOIRS OF THE QUEENSLAND MUSEUM Table 1. Correlation between number of oviducal eggs/ovarian follicles and the snout-vent length (SVL) of all females, mature females (SVL > 55 mm) and gravid females (clutch size > 0). Sample size (n) Pearson product-moment correlation co-efficient (r) Significance probability (p) ALL FEMALES Clutch sized) 91 0.494 0.0001 Vitellogenic follicles (II) 88 0.622 0.0001 Enlarged foIlicles(III) 88 0.472 0.0001 Developing follicles dV) 88 0.508 0.0001 Small follicles (V) 88 -0.349 0.0009 MATURE FEMALES Clutch size (1) 61 0.158 NS Vitellogenic follicles (II) 58 0.257 NS Enlarged follicles (III) 58 0.130 NS Developing follicles (IV) 58 -0.042 NS Small follicles (V) 58 0.124 NS GRAVID FEMALES Clutch sized) 36 0.390 0.019 Vitellogenic follicles (II) 33 0.299 NS Enlarged follicles(ni) 33 0.115 NS Developing foHicles(lV) 33 -0.021 NS Small follicles (V) 33 -0.231 NS scales) showing significant differences between males and females (F=6.11, df=l, p=O.015). Females tend to have fewer nuchal scales (mean±sd, ? 9 : NTOT=16.64±1.83; 6 cJ: NTOT =17.43±1. 66). However, the $ range(NTOT=13 to 21) is contained in that of (5 c3 (NTOT=l 1 to 21) and, as this is a discrete character, the ex- pected value of NTOT for both is 17. Although no significant size differences were detected between the sexes in the above analyses, significant SVL differences were found among sample locations in the anova (F=4.39, df=13, p<0.0001). To further explore these differences the mean of each population was compared with that of each other population using Tukey’s studentized range tests calculated by the SAS procedure ‘GLM’ (SAS institute inc., 1985). The overall type 1 error rate for these comparisons (the rale at which results are falsely accepted as sig- nificant) was restricted to 0.05. Several samples were found to differ significantly in SVL. The skinks from Dowlings’ Fragment (17°23’S 145°4rE) are smaller on average (n=7: SVL=43.6±6.3 mm) than those from other loca- tions, whilst those from Major’s Mountain (17°38’S 145°32’E) were generally larger (n=5: SVL=72.9±3.3 mm). Size and Reproduction All 6S larger than 50 mm SVL displayed enlarged epididymides (approximately 4.5 mm in diameter), which presumably indicates sexual maturity. None smaller than 45 mm SVL pos- sessed an enlarged epididymis. Among the 6 S larger than 50 mm SVL, epididymides were either turgid or flaccid with no clear seasonal or geographic trends. This may indicate a stage in the reproductive cycle, such as the production and storage of sperm, or may be an artefact of collec- tion and fixation. Of the 71 $ 9 from the Atherton Tableland, 25 were gravid (mean SVL=66.8±5.0 mm) and 46 were not (mean SVL=55.5±1 1 .9 mm). The small- est gravid skink 55.5 mm (SVL); the largest (non- gravid) was 78.7 mm (SVL). Using an estimate of T, modified to account for unequal variance between groups, gravid 9 9 were found to be significantly larger than others (modified t=-5,48, df=65.9, p<0.()()01 ). No specimen smaller than 55 nun (SVL) had any class I or II ovarian follicles. Class III follicles were found in only three 9 9 (12%) smaller than 55 mm (SVL), Fig. 1C. The correlation analysis was performed on three nested groups: all 9 9, to investigate general size related trends in reproductive state; only 9 9 >55 mm SVL, to exclude immature skinks 9 9; and only gravid 9 9 (clutch size > 0), to compare skinks in a similar physiological FIG. 2. Temporal variation in percentage of females from the Atherton Tableland carrying oviducal eggs. Sample sizes are given for each month from which collections were available. REPRODUCTIVE BIOLOGY OF THE PRICKLY FOREST SKINK 135 condition. Pearson product-moment correlation coefficients for each of these comparisons are presented (Table 1). In the comparisons including all ? $, SVL was significantly correlated with clutch size and each class of follicles (Table 1). Clutch size and the numbers of class II, III and IV ovarian follicles increase with SVL, the number of class V follicles decreases with SVL, however, there is considerable variation between similarly sized lizards (Fig. lA-E). When immature $ V are excluded from the correlation analysis, varia- tion in the number of eggs and follicles appears to be independent of SVL (Table 1). Among gravid $ 9 clutch size is significantly and posi- tively correlated with SVL (Table 1); large gravid $ $ generally carry more eggs than smaller gravid 9 9 (Fig. lA). Although the number of vitellogenic follicles (II) was not significantly correlated with SVL in gravid females (Table 1), it was significantly correlated with clutch size (n=33, =0.536, p=0.001). Female Reproductive Cycle, Atherton Tableland At least 50% of 9 $ collected from sites on the Atherton Tableland in September, November and December were gravid (Fig. 2). Neither of the two 9 9 collected in February were gravid. Only one of thirteen 9 9 collected in April, and one of seventeen 9 9 collected in June, were gravid (Fig. 2). The Chi square test of homogeneity indicates statistically significant temporal varia- tion in the proportion of gravid females (X^=22.628, p=0.000). A temporal trend was also evident in the development of eggs within gravid females. The single gravid specimen collected in June carried small eggs (average egg length 6.0 mm). Several non-gravid 9 9 collected in June carried very large, class II, follicles (between 4 mm and 6 mm), which were of similar size to small oviducal eggs and were not found in females collected at other times. In gravid 9 9 collected in Septem- ber, eggs were larger (average length =10.7±L2 mm) but showed no obvious signs of fertilisation. Eggs in gravid females collected in November and December were of a similar size to those in September (average egg length 1 1 .8±1 .7 mm) and carried developing embryos, up to 5 mm long, surrounded by yolk. The gravid female from April (J31057) was the only individual in the entire sample canying well developed young, and the only gravid individual to have a clutch size of one. The single foetus found in this female ap- peared to be fully developed and, although still encased in the egg membrane, was not sur- rounded by yolk. This unborn skink had a SVL of 25.9 mm, which was 4 mm smaller than the smallest juvenile skink examined in this study. This apparent seasonality of reproduction sug- gests temporal variation in the average size of G. queenslandiae on the Atherton Tableland, result- ing from an annual pulse of recruitment. The average SVL of G. queenslandiae collected from the Atherton Tableland and examined here, shows no such pattern, being highest in April and November, and lowest in September. In some seasonally breeding species the observed sex ratio varies with the reproductive cycle, as gravid females are often more obvious than other in- dividuals (Schwarzkopf & Shine, 1992; Sim- botwe, 1985). In this study the observed proportion of females varies between 50% in February and 73% in December, however, both months have small sample sizes (n=4 and n=ll respectively). A chi square test of homogeneity shows that there is no evidence for temporal variation in the sex ratio ()c2=l .28 df=5 p=0.937). Bone Histology and Skeletochronology The bone structure of G. queenslandiae is similar to that of other squamates (Castilla & Castanet, 1986; Ricqles, 1976; Enlow, 1969). At mid-diaphysis a weakly vascular or non- vascular cortex of periosteal bane surrounds an almost hollow medullary cavity lined with endosteal bone. The periosteal bone consists of a series of peripherally deposited lamellae, among which are regularly arranged, haematoxylinophilic rings or ‘rest lines’. Endosteal bone results from the resorption and reworking of the inner cortex and so the amount of endosteal increases with the diameter of the femur. The process of erosion of periosteal bone and substitution with endosteal bone results in the obliteration of some rest lines. Linear regressions, predicting the number of rest lines from SVL, were statistically significant for both the total sample and for the seven in- dividuals from the Atherton Tableland alone (all samples - R^=0.76, p=0.0001; Atherton samples -R2=0.77, p=0.01)(Fig 3). DISCUSSION The results of this study allow broad estimates of parameters which are vital to the successful application of population models, and which would be difficult to measure under field condi- tions. These factors include the sex ratio, the clutch size produced, the size at which maturity 136 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Plot of estimated number of haematoxylin stained femural bone rings against SVL for in- dividuals from all locations. Open symbols ~ 66; closed symbols = $ $ . The least squares regression for all individuals (bone rings = 0.12 ±0.02 X SVL- 1.49 ±1.19) depicted above is significant (R^ = 0.76, p < 0.001). The regression including only samples from the Atherton Tableland was also significant (bone rings = 0.09 ±0.02 X SVL+ 0.13 ±1.3, R^ = 0.77, p< 0.01). is attained and the relationship between size and age. These results and others, such as the degree of sexual dimorphism and spatial variation in size, would complement ecological and genetic studies of G. queenslandiae. Sex Ratio There appears to be a slight bias in the sex ratio towards females. As there is distinct temporal variation in the proportion of gravid females, albeit on small numbers, but no similar variation in the sex ratio, it seems that this observed sex ratio bias can not be attributed to the capture of more gravid females than other individuals. The sex ratio of G. queenslandiae would be difficult to determine in the field as there do not appear to be any clear morphological differences between the sexes, although one possible method is the eversion of male hemipenes described by Schwarzkopf (1992). Variation in Size Significant size differences were found be- tween several populations. Although these may be the result of non-random sampling, it does appear that skinks from the southern Atherton Tableland, particularly from Majors Mountain, are larger than those elsewhere. No significant size differences were found between latitudinally separated upland populations or between upland and lowland populations, however, the small sample sizes from areas other than the Atherton Tableland may be insufficient to detect such dif- ferences. Considerable genetic variation has also been detected between geographically separated populations of G. queenslandiae (Moritz et al., 1993). The degree of congruence between this genetic variation and morphological variation could not be examined because of limited sample sizes from matched sites and possible collector bias. Reproductive Biology Sexual maturity in females is attained at ap- proximately 55 mm SVL. Males probably mature at a slightly smaller size. Skinks smaller than 50 mm were uncommon in this collection. This may reflect collection bias, rapid growth to maturity, that juvenile skinks are more difficult to locate than mature animals, or a combination of these. In mature females ovarian follicles are always present but vitellogenesis is seasonal. Developing follicles greatly outnumber the number of ovidu- cal eggs, presumably with the remaining vitel- logenic follicles being resorbed after the breeding season. According to Shine (1977) the production of more vitellogenic follicles than are ovulated is of adaptive value, allowing an individual to adjust clutch size according to environmental conditions at the time of ovulation. These additional en- larged follicles may also allow multiple clutches Vitetlogenesis OviJation Fertilisation and development Parturition FIG. 4. Reproductive cycle of G. queenslandiae on the Atherton Tableland. Vitellogenesis peaks early in the mild, dry season, and is followed by ovulation and fertilisation mid-year. Development of young occurs late in the dry season and throughout the warm wet season, with parturition late in the wet season. REPRODUCTIVE BIOLOGY OF THE PRICKLY FOREST SKINK 137 within a breeding season. The latter suggestion is supported by the observed correlation between clutch size (I) and the number of vitellogenic follicles (ID. The correlations between the numbers of ovarian follicles and SVL, when all females are considered, and the lack of any correlations with SVL when only females larger than SVL=55 mm are considered, indicates that the number of en- larged follicles (III & IV) produced by immature females increases with size until maturity. The negative correlation between the number of small follicles (V) and SVL, when immature females are included, suggests that the total number of follicles is relatively constant throughout life. As females grow towards maturity some class V follicles become enlarged, and so the number of follicles that remain small decreases. As in studies of other squamates (Schwarzkopf, 1992; Simbotwe, 1985; Shine, 1977), clutch size in G. queetislandiae is correlated with size of females. Life history theory predicts that females should maximise lifetime reproductive output, and that it may be advantageous for a female to use energy reserves for growth rather than breed- ing, if the probability of survival is high and additional growth will substantially increase fu- ture clutch sizes (Schwarzkopf & Shine, 1992). The relationship between clutch size and SVL (Fig. 1 A) is consistent with this prediction in that larger females produce either small or large clutch sizes whilst smaller females only produce small clutches. In each of the samples of G. queenslan- diae considered here a proportion of ‘mature' females (SVL>55 mm) were not gravid. In the related seasonally breeding, viviparous skink Eulamprus tympanum, Schwarzkopf (1992) found that variability in reproductive output per year was expressed as variation in offspring size, and possibly the proportion of non-breeding females, whilst the average clutch size remained constant across years. The lack of replicate samples from the same sites precluded such an analyses of between-year variation in this study. Suggested Reproductive Cycle The skinks considered in this study were col- lected not only in different months, but also in different years, so it is not possible to distinguish within-year patterns from between-year varia- tion. The predictable climate of the Atherton Tableland and the pattern of temporal variation in the proportion of gravid females and egg develop- ment suggest the following seasonal reproductive cycle in G. queenslandiae (Fig. 4). Active vitel- logenesis becomes apparent in June and is fol- lowed by ovulation in June and July. Fertilisation and development of foetuses occurs between Sep- tember and January. Parturition occurs between February and April and is probably accompanied by regression of the vitellogenic follicles that were not ovulated. The time at which mating occurs could not be inferred from the results of this study, as females may store sperm for a considerable time before fertilisation (Fitch, 1982), in this case ovulation. Reproducti ve eye les i n other tropical ski nks are highly variable both between and within species (Fitch, 1982). The primary determinant of seasonality of reproduction in tropical skinks ap- pears to be the seasonality of precipitation (Fitch, 1982). Precipitation on the Atherton Tablelands is distinctly seasonal with average rainfall over 250 mm in January, February and March, and below 50 mm in June, July, August, September and October (measured at Kairi experimental sta- tion; AGPS 1988). Temperature is also seasonal on the Atherton Tableland and varies between a mean daily minimum of 10.9°C, with occasional frost, in July, and a mean daily minimum of 19.5®C in February (Kairi experimental station; AGPS 1988). In the reproductive cycle outlined above, embryonic development on the Atherton Tableland would occur during the warm, rainy season, with parturition occurring just before the onset of the mild, dry weather (Fig. 4). In other parts of the Wet Tropics, particularly the Bellen- den Ker Ra. immediately to the east of the Ather- ton Tableland, precipitation and temperature are less distinctly seasonal and populations of G. queenslandiae from these areas may have dif- ferent reproductive cycles. Skeletochronology The number of ‘rest lines’ in the femurs of G. queenslandiae shows a clear relationship with size which is roughly approximated by: rest lines = 0.1 X SVL (Fig. 3). Although these rest lines can not be assumed to represent a true index of age as a result of remodelling of the bone, they may be a useful measure of age differences be- tween skinks (Enlow, 1%9). Mature individuals had at least five rest lines and no more than nine. If it is assumed that the intervals between rest lines represent constant periods of growth, these results suggest that maturity in G. queenslandiae is delayed until quite late in life. In other species, mainly those from temperate regions, rest lines are formed semi-annually or annually but not less frequently (Castanet et al.. 138 MEMOIRS OF THE QUEENSLAND MUSEUM 1988). The observed number of rest lines in G. queenslandiae could be used in demographic models as a rough estimate of maximum age, where skinks reach maturity at five years of age and live a maximum of ten years. A sex ratio of 1:1.5 could be used in this model with 75% of mature females breeding each year (maximum number of gravid females with SVL>55 mm) and each gravid female producing a clutch size of (3.11=average clutch size). Field studies of size specific mortality and population density would be required to complete this basic model. ACKNOWLEDGEMENTS This project was funded by a Queensland Museum Student Research Scholarship. Thanks are extended to Patrick Couper and Jeanette Covacevich for support and access to the museum collections; to Patrick Couper and Craig Moritz for suggestions on possible comparisons and helpful advice on the literature; to Lina Daddow and Zenab Kalel for assistance with the bone histology; and to Damien Broderick, Leo Joseph, Nicole Lee, Craig Moritz, Jeff Miller and an anonymous referee for constructive comments on the manuscript. LITERATURE CITED AUSTRALIAN GOVERNMENT PUBLISHING SERVICE (AGPS). 1988. ‘Climatic averages Australia’. (AGPS Press: Canberra). BETZ, T. W. 1963 The gross ovarian morphology of the Diamond-backed Water Snake, Natrix rhom- bifera, during the reproductive cycle. Copeia 4:692-697 CASTANET. J. , NEWMAN, D. G. & SAINT GIRONS, H. 1 988 . Skeletochronological data on the growth, age, and population structure of the tuatara, Sphenodon punctatus^ on Stephens and Lady Alice Islands, New Zealand. Herpetologica 44(1): 25-37 CASTILLA, M. A. & CASTANET. J. 1986. Growth, age and longevity of Lacerta lepida assessed by skeletochronology. Pp33I-336. In Rocek, Z. (ed), ‘Studies in herpetology’. (Charles Univer- sity: Prague). COGGER, H. G. 1992 Reptiles and amphibians of Australia. Pp 498^99 (Reed Books: Sydney) GREER, A. E. 1989 The biology and evolution of Australian lizards. Pp 154-155 (Surrey Beatty: Sydney) ENLOW, D. H. 1969. The bone of reptiles. Pp 45-80. In Gans, C. , Bellairs, A. & Parsons, T. S. (eds), ‘Biology of the Reptiha. Volume 1’. (Academic Press: London). FITCH, H. S. 1982. Reproductive cycles in tropical reptiles. Occasional papers of the Museum of Natural History 96:1-53. The University of Kan- sas, Lawrence, Kansas. MORITZ, C. . JOSEPH, L. & ADAMS, M. 1993 Cryptic diversity in an endemic rainforest skink (Gnypetoscincus queenslandiae). Biodiversity and Conservation. 2:412-425. NIX, H. A. & SWITZER, M. A. 1991 Tropidophorus queenslandiae. P.82. In ‘Rainforest animals. Atlas of vertebrates endemic to Australia’s wet tropics.’ (Australian National Parks and Wildlife Service. Canberra). RICQLES, A. J. 1976. On bone histology of fossil and living reptiles with comments on its functional and evolutionary significance. Pp 123-151. In Bellairs. A. & Cox, B. (eds), ‘Morphology and biology of reptiles. Linnean Society Symposium Series 3’. (Academic Press: London). SAS INSTITUTE INC 1985 SAS/STAT user’s guide, release 6.03 edition. (SAS Circle: Cary North Carolina) SCHWARZKOPF, L. & SHINE, R. 1992. The evolu- tion of reproductive effort in lizards and snakes. Evolution 46:62-75. SCHWARZKOPF, L. 1992. Annual variation in litter size and offspring size in a viviparous skink. Herpetologica 48:390-395. SHINE, R. 1977. Reproduction in Australian elapid snakes II. Female reproductive cycles. Australian Journal of Zoology 25:655-666. SIMBOTWE, M. P. 1985. Sexual dimorphism and reproduction of Lampropholis guichenoti (Lacertilia: Scincidae). Pp 11-16. In Grigg, G. , Shine, R. & Ehmann, H. (eds), ‘Biology of Australasian frogs and reptiles’. (Royal Zoologi- cal Society of New South Wales: Sydney). WINTER, J.W., BELL, F.C., PAHL, L.l. & ATHER- TON, R.G. 1987 Rainforest clearfelling in north- eastern Australia. Proceedings of the Royal Society of Queensland 98:41-57 ZUG, G. R., HEDGES, S. B. & SUNKEL, S. 1979 Variation in reproductive parameters of three neotropical snakes, Coniophanes fissidens, Dip- sas catesbyi, and Imantodes cenchoa. Smith- sonian Contributions to Zoology No. 300. TAXONOMY AND DISTRIBUTION OF THE SCINCID LIZARD SAPROSCINCUS CHALLENGERI AND RELATED SPECIES IN SOUTHEASTERN AUSTRALIA ROSS A. SADLIER, DON J. COLGAN AND GLENN M. SHEA Sadlier, R.A., Colgan, DJ. & Shea, G.M. 1993 12 24: Taxonomy and distribution of the scincid lizard Saproscincus chalkngeri and related species in southeastern Australia. Memoirs of the Queensland Museum 34(1):139-158. Brisbane. ISSN 0079-8835. The skink Saproscincus chalkngeri is here recognised as comprising three species on the basis of allozymic and morphological variation. S. chalkngeri is redefined and restricted to the McPherson Range region, southeastern Queensland. The names Saproscincus galli Wells & Wellington (1985) and Saproscincus Wells &WeIlington (1985) apply to two widespread species. All species occur almost exclusively in closed forest. Mocoa spectabilis De Vis. 1888 is shown to be a senior synonym of Saproscincus basiliscus (Ingram & Rawlinson, 1981). □ Scincklae, Saproscincus, taxonomy, electrophoresis, distribution, rainforest. Ross A. Sadlier, DonJ. Colgan, Australian Museum, POBoxA285, Sydney South, New South Wales 2000, Australia: Glenn M. Shea, Department of Veterinary Anatomy, University of Sydney, New South Wales 2006, Australia; 15 September, 1993. As our knowledge of the taxonomy and relationships of the Australian scincid lizard fauna has been refined in the past two decades, a number of widespread species have been found to be composite. One such example is the species Lygosoma challengeri Boulenger (1887), which was formerly regarded as widespread in closed forests of northern New South Wales and Queensland (Worrell, 1963; Dale, 1973; Cogger, 1975). This taxon has variably been assigned to the genus Leiolopisrna (Cogger, 1975), Lampropholis (Greer, 1974) and the challengeri species group within Lampropholis (Greer & Kluge, 1980). The latter species group was regarded as generically distinct by Greer (1980) and subsequently given the generic name Sapros- cincus by Wells & Wellington (1984). Greer (1980) and Greer & Kluge (1980) listed two additional species in the complex from the north- ern end of the distribution. One of these, Lampropholis tetradactyla^ was a new discovery (only two records were known prior to 1974), while the other, which Greer & Kluge diagnosed but left undescribed, represented northern populations previously assigned to L. challengeri (WorrelPs 1963 diagnosis of challengeri was at least partly based on the northern taxa). The second species was subsequently described as Lampropholis basiliscus (Ingram & Rawlinson, 1981), with a third northern species in the chal- lengeri group, Lampropholis czechurai. Conse- quently the name challengeri was restricted to soiithem (SEQ-NENSW) populations within the complex, although no author had redefined that species. The possibility of the restricted S. challengeri being composite was suggested by Wells & Wel- lington (1985), who named two additional species, 5. galli and S. rosei, both from single specimens, and by Wilson & Knowles (1989), who figured several distinctive morphotypes from SEQ. Neither gave evidence to distinguish these taxa at the species level. An undescribed member of the challengeri species group from the Sydney region has been known for many years (e.g. Griffiths, 1987). Several specimens of the same taxon (one figured by Wilson & Knowles, 1988, pl.502) were col- lected (RAS) in the Bellingen region, NENSW, in 1983, a northern extension of the known dis- tribution of over 400km. The discovery of this species in regional sympatry with what was then considered typical S. challengeri initiated the in- vestigations reported here. MATERIALS AND METHODS The species recognized are identified by pos- sessing unique combinations of both morphologi- cal and electrophoretic characters, and their species-level distinction is supported by field ob- servations relating to distribution, habitat preferences, and the occurrence of sympatry be- tween species. Electrophoretic procedures: Electrophoresis of liver samples was performed on ‘Titan Ilf (Helena, Austin) cellulose acetate gels according 140 MEMOIRS OF THE QUEENSLAND MUSEUM to standard procedures (Hebert & Beaton, 1989). Gels were run for 60 minutes, with a constant potential drop of 200V between electrodes. Twenty-one enzyme systems encoded by 24 loci were scored. Staining protocols were adapted from Harris & Hopkinson (1977) and Hebert & Beaton (1989). Fluorescence methods were used for esterase and negative stains for superoxide dismutase. The enzymes stained, abbreviations used herein. Enzyme Commission numbers, run- ning buffer and number of presumptive genetic loci are given in Table 1. Tissue was ground in 1 volume of tissue to 1 volume of homogenising buffer (l(X)ml tris-HCI, pH 7.0, ImM Naz EDTA, 0.5mM NADP and 50|Xi/100mI (i-mercap- toethanol) in hand-held glass homogenisers. Al- lozymes are designated in order of their relative anodal mobility, as are different loci encoding the same enzyme. Results were analysed using the BIOSYS-1 package of Swofford & Selander (1981). Morphological studies: All specimens of S.challengeri in the Australian and Queensland Museums were examined. Specimen registration numbers for Australian Museum (AMS) specimens are prefixed R and Queensland Museum (QM) specimens, J. From this material 23 series of specimens, corresponding to most of the samples analysed biochemically, but in some cases enlarged by the addition of specimens from the same or nearby localities, were examined for the full suite of characters listed below. Measure- ments (axilla to groin, hindlimb, and tail lengths) are expressed as percentages of snout to vent length (SVL) in the taxon accounts. The following characters were scored for each specimen where possible: Axillato groin distance (AGL). Hindlimb length (HLL) - measured from TABLE 1. Enzymes stained (I), abbreviations (2), Enzyme Commission numbers (3), running buffer (4) and number of presumptive genetic loci (5). 1 2 3 4 5 Adenylate kinase AK A^artale aminMransferase AAT Esterase EST Fructose btsphosphatase FBP Fumarate hydratase FH Glucosephosphale isomerase GPI Glucose-b'phosphale dehydrogenase G6PD Glyceraldehydc phosphate dehydrogenase GA-3-PDH ^Glycerophosphate dehydrogenase GPD Isociirate ^hydrogenase IDH Lactate tlchytlrogenase LDH Malate deh]^rogetta.se MDH Mannosephospivtie isootcrase MPl PepiKlase (ku-ula .vuhslrate) PEP- la Peptidase (leu gly-gly substrate) PEP-Igg Peptidase (plie-prt? substrate) PEP pp PhoNphoglucumuia-se PGM 6- Phcsphogluconate dehydrogenase 6-PGDH Superoxide dismutase SOD Triosephosphate isomerase TPl UDP glucose pyrophosphorylase UDPG 2.7.4.3 TEM 50 2.6.1. 1 TC 100 3.1. 1.1 TBM50 3.1.3.1ITEM 50 4.2. 1.2 THM50 5.3. 1.9 TEM 50 1.1.1.49TC 100 I .2.1.12TEM 50 I 1.1.8 TEM 50 1.1 I .42TEM 50 1.1.1.27TC 100 I 1.1.37TEM50 5.3 1.8 TEM 50 3.4.11 TEM 50 3.4.11 TEM 50 3.4 11 TEM 50 2.7.5. 1 TEM 50 1.1.1.44TEM 50 I.15.MTEM 50 5.3.1. 1 TEM 50 2.7.7.9 TC 100 1 2 2 2 2 groin to tip of fourth toe including nail. Tail length (TL) - measured from caudal edge of anal scales to tip of tail , on complete original tails only. Supraciliaries (SCIL) - first row of enlarged scales above eye posterior to prefrontal scales and bordering supraocular scales. Last scale in series is that abutting the posterior edge of the fourth supraocular. Midbody scale rows (MB) - number of longitudinal scale rows around body counted midway between axilla and groin. Paravertebral scales (DSR) - number of scales in a paravertebral row from first scale posterior to parietal scale to last scale anterior to level of vent opening. Fourth finger (FS) and toe (TOES) scales - number of dorsal scales on fourth digit of foot and hand. Distal scale contains claw and basal scale broadly contacts adjacent basal scale of third finger or toe. Fourth finger (FL) and toe (TOEL) lamellae - number of ventral scales on fourth digit of foot and hand. Distal scale contains claw and basal scale is last largely undivided scale at a point level with intersection of third and fourth digits. Bilaterally scorable characters (SCIL, FS, FL, TOES, TOEL) were scored on both sides and the mean value used. Apart from tail length, which was not subjected to analysis, all characters had significant geographic variation using one-way analysis of variance. The metric characters AGL and HLL both showed allometric growth in comparison to SVL. To remove the effects of varying size and allometric growth the values for these two vari- ables were log-transformed and adjusted to the values they would assume at a constant SVL using the formula (Thorpe, 1975) V=e where y is the adjusted dependent variable, yi is the raw dependent variable, xi is the SVL for that individual, x is the mean SVL across all samples and a the allometric coefficient from the regres- sion ln(y) = a*ln(x)+b. Values were adjusted to mean SVL49.287mm. As an estimate of a the mean value of the al- lometric coefficients for the three best sampled populations (populations 6, 9, and 23 correspond- ing to the three putative biochemical species) was used: 1 . 1 27 for AGL and 0.778 for HLL. Canonical variates analyses were run with the nine characters, adjusted AGL, adjusted HLL, SCIL, MB, DSR, FS, TOES, FL, and TOEL, using S YSTAT (Wilkinson, 1 987). Four analyses were run. In the first analysis, all twenty-three samples were treated as a priori groups (opera- tional taxonomic units). In the second analysis, TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 141 the three putative species were used as groups. The final two analyses treated males and females separately, again using the three putative species as groups. In total, 210 animals were run in the first two analyses, while 88 males and 120 females were run in the last two analyses. Two features of osteology were also surveyed but because of the more limited samples ex- amined were not included in the main analysis above. These were number of presacral vertebrae (anterior to sacrum), and postsacral vertebrae (posterior to sacrum). RESULTS The results of combined electrophoretic and morphological analyses on 27 populations iden- tified three major groups. The presence of one or more fixed allelic differences between regionally sympatric populations of each group was regarded as clear evidence of species level dif- ferentiation. Regional sympatry is here def ^ned as species occuring in the same general vicinity but not necessarily within the same habitat or al- titudinal range. From among these 27 populations sampled for electrophoretic analysis regional sympatry was observed between all possible pair- ings of the three major groups at one or more localities. On this basis, three species are recog- nised to which the following names apply: S. challengeri from the McPherson Ranges, SEQ and hinterland far NENSW; S. rosei from the eastern edge of the Great Dividing Range and associated ranges, NENSW and SEQ; and S. galli from isolated areas of the central and northern coast and adjacent ranges, NSW, and the Mc- ' Pherson Ra., SEQ. Interspecific Variation 1. Electrophoresis: The electrophoretic results are presented in Table 2. A phonogram based on Nei’s unbiased genetic distance is shown in Fig. 1. Three main groups of populations (separated at the 0.2 distance) can be recognised. There are fixed allozymic differences between all of these, including cases of sympatry or near sympatry, for each pair of groups. All populations of S. challengeri and S. galli examined electrophorctically, including two in- stances where populations of each species oc- curred in ecological sympatry (syntopy), show fixed allelic differences for the Ak, G-6-pdh and Idh-J loci, and nearly fixed differences in Est~J and Gpi, In addition S. challengeri has only one observable Mdh locus, whereas 5. galli and in- FIG. 1. Phenogram of genetic similiarity based on Nei’s unbiased genetic distance, populations 1-14 S. rosei, populations 15-19 5. challengeri, populations 20-27 5. gain. n FIG. 2. Canonical variates analysis of 23 Saproscincus populations. Ordination of 23 populations on first two canonical vjiriates. Points represent individuals as follows: dots = S. galli; squares = S. rosei; triangles = S. challengeri; open circles represent type speci- mens (1 = S. challengeri; 2 = S. galli; 3=5. rosei). Polygons enclose scatter of points for each taxon. Population Population MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 2: Gene frequencies in Ihe smdied Saproscincus populations. Populations 1 - 14 are S. rosei, 15 - 19 are 5. challengeri and 20 - 27 are 5’, gallL given in the text. Numbers in the same row as locus identifications indicate sample sizes. Where no frequency is listed for IDH-2 or MDH-2, the locus is not present in the population. TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 143 n FIG. 3. Canonical variates analysis of Saproscincus populations. Same axes and polygons as Fig. 2. Group centroids (numbered) of populations. Dots = 5. galli; squares = S. rosei\ triangles = S. challengeri. Popula- tions 4-5, 12 and 26 analysed biochemically but not included in canonical variates analysis. deed all other examined Saproscincus taxa have two. 5. challengeri (populations 17-19) and region- ally sympatric S. rosei from NENS W (population 12 separated by less than 10km from population 17) and SEQ (population 13) showed fixed allelic diferences at the Ak, G-6~pdh, Gpi, ldh-1, Mpi, Pep-la and 6-Pgdh loci, in addition to the absence of a scorable Mdh-2 from S. challengeri and a scorable Idh-2 from S. rosei. With the exception of Pep-la, the differences listed above also distin- guish S. challengeri from allopatric S. rosei. S. galli and S. rosei show fixed allelic differen- ces for the Mpi and 6-Pgdh loci for all populations examined, including three instances of regional sympatry between S. galli. and S. rosei. 2. Morphology: The initial canonical variates analysis, using populations as OTUs, identified three largely distinct groups of populations on the first two functions (Figs 2-3; Table 3). These three groups corresponded exactly with the groupings determined biochemically. Factor 1 separated two clusters of populations (groups 1 +2 and group 3) largely on the basis of subdigital lamellae (FL, TOEL) while factor 2 separated a third population (group 2 from group 1) largely on the basis of midbody scale rows, supraciliaries, and supradigital scales (MB^ SCIL, TOES; Table 3), Canonical scores for primary type specimens (see below) of the avail- able names within the 5. challengeri complex were calculated using the functions: the lectotype of S. challengeri was placed among group 1; the holotype of S. galli lay among group 2; and the holotype of 5. rosei lay among group 3. One other name has been previously placed in the synonomy of S. challengeri (Cogger et al,, 1 983). Mocoa spectabilis De Vis (1888) was described from specimens from Gympie, SEQ. Of the four identifiable types in the Queensland Museum (Covacevich, 197 1; Cogger etal., 1 983) one (QM J244) (Fig. 4) is S. galli, while the other three (QM J255, J 19742-43) agree in all respects with S. basiliscus (Ingram & Rawlinson, 1981), a species not known from further south than the Mackay district, MEQ. Diagnostic characters of the latter species shown by these three specimens include the presence of a divided nuchal scale bordering the parietal and two tertiary temporals bordering the posterior edge of the lower secon- dary temporal scale. Designation of J 19743 (Fig. 5) as lectotype by Wells & Wellington (1985) means that Mocoa spectabilis becomes a senior synonym of, and the available name for the taxon previously known as Saproscincus basiliscus. Treating the three species as OTUs in the second canonical variates analysis resulted in 96.2% of specimens being correctly identified (96.7%, n=63 of S. challengeri', 98.2%, n=58 of S. galli; and 94.8%, n=101 ofS. (Canonical coefficients and loadings for characters for the two functions extracted are given in Table 4. Group classification coefficients are presented in Table 5, allowing unknown specimens to be as- signed to species. TABLE 3. Standardised discriminant function coefficients (and correlations with discriminant functions) for the first seven discriminant functions of nine characters from the Saproscincus challengeri complex. Twenty-three of the 24 populations biochemically sampled are used as a priori groups. Group 12, represented by a single individual, not used in determining functions. Variable I II III IV V VI VII AGL (adj) 0.019(0.012) 0.162(0.164) -0.119(0.004) -0.037(0.097) 0.482(0.408) -0.007(0.032) 0.551(0.646) HLL(adj) 0.151(0.216) -0.234(-0.173) 0.162(0.170) 0.359(0.357) -0.617(-0.601) 0.305(0.323) -0.137(-0.061) MB -0.L59(-0.131) -0.609(-0.484) 0.267(0.383) 0.204(0.158) 0.128(0.159) 0.196(0.105) 0.288(0.347) DSR -0.215(-0.138) 0.215(0.145) 0.605(0.659) 0.267(0.312) -0.175(-0.045) -0.665( -0.629) 0.059(0.123) SCIL -0.254{-0.106) 0.577(0.487) -0.135(-0.125) -0.164(-0.23I) -0.529(4).522) -0.036(0.003) 0.292(0.376) FL 0.403(0.715) 0.032(0.117) -0.284(0.029) 0.135(0.066) 0.209(0.109) -0.536(-0.350) -0.492(-0.151) FS 0.065(0.311) 0.079(0.200) 0.641(0.664) -0.788( -0.525) 0.080(0.107) 0.254(0.256) -0.192( -0.173) TOEL 0.685(0.856) -0.170(0.068) 0.055(0.141) -0.068(-0.012) -0.206(-0.133) 0.039(-0.052) 0.640(0.386) TOES -0.020(0.272) 0.581(0.547) 0.102(0.299) 0.648(0.461) 0.242(0.287) 0.497(0.429) -0.133(-0.I11) 144 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. QM J244 syntype of Mocoa spectabilis De Vis, a specimen of S. galli. Treating both sexes seperately did not offer any noticeable improvement in resolution. Overall, 96.6% of males and females were correctly as- signed to groups. INTRASPECIHC VARIATION 1. Electrophoresis: The electrophoretic results clearly demonstrate that there are at least three species level taxa in the S. challengeri group. The Nei distances separating lineages within these groups are, however, quite high, the distances approaching, in the case of S. galli, the figure of 0.15 which has been suggested (Thorpe, 1982; Nei, 1987) as a criterion for determining the specific status of a population when this is other- wise unresolvable. Tbis figure is probably too low for lizards, as sister species in this group, with the exception of some iguanids (Gorman & Kim, 1976; Adesl, 1977; Case & Williams, 1984), generally exhibit Nei distances of more than 0.2 (Kim et al., 1983; Milton et al., 1983; Busack, 1986; Daugherty et al., 1990). Nevertheless, the possibility of taxonomic structuring within S. rosei, S. galU, and S. challengeri should be con- sidered . On the basis of genetic (Nei distance) similarity two subgroups of S. rosei are recognised. One includes populations from central eastern and NENSW, the other populations from SEQ. In- cluded in the SEQ subgroup are distinctive regionally restricted high altitude populations in the eastern border ranges. Tbe first subgroup has fixed differences from the second for Pep-la and 6-Pgdh, and a nearly fixed difference for Est. Another distinguishing feature of the SEQ sub- group is the presence, at a relatively high frequen- cy of the GPI A allozyme, which is absent from the central-eastern and north-eastern subgroup of FIG. 5. QM J 19743 syntype of Mocoa spectabilis De Vis designated as lectotype for the species by Wells & Wellington, 1985. TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 145 TABLE 4.Standardised discriminant function coeffi- cients (and correlations with discriminant functions) of nine characters from the Sapwscincus challengeri complex. The three putative species based on biochemical data used as a priori groups. Discriminant Function Variable AGL (adjusted) HLL (adjusted) MB DSR SCIL FL FS TOEL TOES I -0.131(-0.075) 0.290(0.290) 0.146(0.060) -0.363(-0.146) -0.485(-0.306) 0.283(0.607) 0.048(0. 177) 0.725(0.732) -0.254(0.059) n 0.247(0.165) -0.305(-0.109) -0.683(-0.537) -0.040(-0.039) 0.472(0.419) 0.272(0.423) 0.028(0.180) 0.126(0.418) 0.431(0.435) S. rosei. Within the SEQ subgroup of S. rosei there is a fixed difference for between the high altitude populations in the eastern Border Ranges and those populations to the west and north. In these high altitude populations the GPI A and PEP-LA C allozymes are found in low frequen- cies but occur at frequencies of more than 50% in other populations of the SEQ subgroup of 5. rosei Two subgroups of S. galli were identified, separated at a Nei unbiased distance of about 0. 1 . One subgroup comprises the populations from the Dorrigo region in the central part of the species range, the other subgroup, separated by several hundred kilometres from the first, com- prises the northern and southern populations. No morphological differences were observed to sup- port differentiation of the Dorrigo populations of S. gain from those to north or south. The electrophoretic differentiation is due to a fixed difference between the subgroups for P ep-pp and the relatively high frequencies of allozymes AK C, IDH-1 C and D, and GPI B which are found at lower levels in other S. galli. The Pep~pp B al- lozyme which distinguishes Dorrigo S. galli from other populations of this species is fixed in S. rosei and nearly fixed in S. challengeri. The presence of the allozyme may be due to its reten- tion in Dorrigo S. galli rather than evolution in TABLE S.Fisher group classification function coeffi- cients for the three species in the Saproscincus chal- lengeri complex. Individual animals are assignable AGL (adjusted) challengeri 11.352 HLL (adjusted) 18.156 MB 20.325 DSR 10.064 SCIL 43.111 FL 13.693 FS 0.264 TOEL 0.555 TOES -2.373 Constant -1124.175 galli rosei 12.269 11.895 16.279 16.614 17.051 19.114 10.304 10.805 50.560 49.498 14.325 12.654 0.311 -0.047 -0.063 -1.773 0.803 -0.376 -1130.669 -1112.650 situ. The frequency differences for other al- lozymes may be due to local adaptations or genetic drift but do not suggest that the Dorrigo S. gain is greatly genetically isolated from other populations of this species. ilie populations of S. challengeri showed little genetic variation across their range, a notable exception being the fixation of the MPl D al- lozyme in the Mt Warning population, this form being at low frequencies elsewhere. The range of the species is, however, small in comparison to that of S. galli or S. rosei so the same degree of genetic structuring should not be expected. 2. Morphology: Within each of the three major species there was little additional resolution of populations on subsequent discriminant func- tions in the first analysis. Factor 3 partially separated the regionally restricted high altitude populations of S. rosei (Population 13 - Toolong Falls) in the eastern McPherson Range from other populations of S. rosei^ although much overlap remained. Presacral and poslsacral vertebrae number was surveyed for a sample of individuals (n=88) rep- resenting most of the populations examined. S. challengeri, S. galli, and populations of S. rosei from central eastern and NENSW usually had 27, rarely 28 or 26 presacral vertebrae, while popula- tions of 5. rosei from SEQ (including population 13) usually had 28, occasionally 29 presacral vertebrae. SPECIES DESCRIPTIONS Saproscincus challengeri (Boulenger, 1887) Figs 6-8 Lygosoma challengeri Boulenger, 1887; 575. Type material. Lectotype BMNH 1946.8.16.55 (Fig. 6), Paralectotype BMNH 1946.8.16.56 Queensland. SPECIMENS Examined The following specimens were used in the description, md include those used in the canonical variates malysis and electrophoretic analysis (morphological uialysis only electrophoretic analysis only !). Border Ranges NP, Lophostemon Falls on Brush Box ^alls track (population 17), 28.24’S, 153.04’E R133463-66;R133467-71*,R138{)62-66); MtWarn- ng NP (population 19), 28.’24S, 153.18 E R133450*; R133451-53; R133454-56*; R133457; fl33458-61*; R138005-06*, RI38017-18; R1380I9*R138020!; R138021-22; R138023-24* R 138026*, R138028-30*); Nightcap NP, vicinity of Terania Ck picnic area (population 18), 28.34’S, \53 WE (R138077-78; R138079-82*; R138083- 146 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. BMNH 1946.8. 16.55 syntype of Lygosoma challengeri Boulenger designated as leclotype for the species by Wells & Wellington, 1985. 87*); Dome Mt. area, Yabbra SF (population 16), 28.28’S, 152.40’E (R135469, R13547I); 28.27’S, 152.39’E (R135470); 28.27*S, 152.34’E (R135472- 73); Tooloom Ra., Yabbra SF (population 15), 28.38’S, I52.29^E (R135461-62; 135465-66); 28.35’S, 152.29^E(R135459-60*); 28.34*S, 152.29*E (R135463* R135468*). Other specimens used to map the distribution of this species. S. galli and S. rosei are listed in appendix 1. Diagnosis The following features in combination general- ly distinguish S. challengeri from other members of the S. challengeri species group: maximum adult size 57mm; supraciliaries usually 6; lamel- lae beneath fourth finger 15-19; lamellae beneath fourth toe 22-27; presacral vertebrae usually 27; postsacral vertebrae 45-48; dorsal surface uniform brown; ventral surface with irregular brown spotting; dorsal surface of tail usually with several moderately large, pale vertebral blotches anteriorly; abdomen of adult males uniformly cream or with a pale yellow wash. The first, fourth, and ninth characters will dis- tinguish S, challengeri from S. rosei which reaches a greater adult size (maximum SVL 64mm), usually has fewer fourth toe lamellae (17-24), and lacks moderately large tail blotches. The fifth and eighth characters will also distin- guish S. challengeri from regionally sympatric northern populations of S. rosei which have 28-29 presacral vertebrae and the markings on the ventral surface faint and regularly aligned. The second, and sixth to tenth characters will distinguish S. challengeri from 5. galli which usually has 7 supraciliaries, 48-53 postsacral ver- tebrae, a mottled dorsal colour, the markings on the ventral surface regularly aligned, pale mark- ings on the dorsal surface of the tail present as isolated spots a single scale in size, and the ab- domen of adult males a bold lemon yellow. Description Measurements: Maximum SVL 57mm; TL 137-168% of SVL (x=155.8%, n=27); AG 51- 59% of SVL (x=55.6%. n=62); HL 36-44% of SVL (x=4L3%, n=60). Scalation: Nasals widely separated; prefrontals moderately to narrowly separated; supraciliaries 6-7 (x=6.0, sd=0.1 6, n=63); upper labials 6, rarely 7; midbody scale rows 23-26 (x=24.6, sd=1.0, n=63); paravertebral scales 54-63 (x=57.6, sd=L6, n=63), fourth finger scales 9-1 1 (x=10.0, sd=0.3, n=63); fourth finger lamellae 15-19 (x=17.3, sd=0.9, n=63); fourth toe scales 11-13 (x=l 1 .2, sd=0.4, n=63); fourth toe lamellae 22-27 (x=24.3, sd=1.0, n=63). Osteology: Presacral vertebrae 27-28 (x=27.1, sd=0.3, n=16); postsacral vertebrae 45-48 (x=46.8, sd=Ll.n=12). Colour and pattern: The populations of S. chal- lengeri are generally similar. Two forms of sexual dichromatism, both at low frequencies, occur in certain populations. Dorsal surface usually over- all mid brown (occasionally lighter or darker), TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 147 uniform or with scattered pale brown scales. Dor- solateral region with dark flecking along scale rows 3 and 4 forming a rough-edged stripe at least anteriorly, and variably extending partly or whol- ly along the body and basal portion of the tail. >^ere dark dorsolateral markings continue to level of hindlimb these may occasionally be bor- dered above by a pale, poorly defined, brown to russet hip stripe. Head with a bold dark brown loreal streak between nares and eye, becoming narrower and obscure behind eye and generally not continuous with dark dorsolateral stripe. Dor- sal and lateral surfaces of tail similar to body, usually marked with several moderately large, pale vertebral blotches anteriorly, rarely uniform, and occasionally defined by a continuation of the fine dark dorsolateral flecking on scale rows 3 and 4 of body. Dark lateral and pale ventral surfaces of tail in bold contrast and the ventrolateral mar- gin of tail defined by a fine but obvious black stripe. Ventral surface white with sparse to heavy scattering of mid brown spots positioned in either the centre or edge of the individual ventral scales. In life adults males generally have a pale yellow wash to the posterior half of the abdomen, oc- casionaly extending to the underside of hind! imbs and basal portion of tail, whereas adult females generally lack such colour or only occasionally have a very weak yellow flush to the posterior half I of the abdomen. I Two colour patterns were observed in females of this species. A small proportion of females (approximately 10-20%) from all populations had a uniform mid brown dorsal surface bordered by a smooth, well defined, pale edged, dark dor- solateral stripe continous along the body and tail and contrasting with the darker brown lateral colour to the Ixxly and tail. In the Nightcap Range population a small percentage of females had a relatively plain mid brown dorsal surface and boldly contrasting uniformly darker brown upper laterd surface bordered below by a broad, bold white stripe which occupied most of the mid and lower lateral region, particularly in the region of the forelimb. One of these ‘white-striped’ specimens had the dorsal and lateral surfaces further distinguished by a narrow, pale brown laterodorsal stripe along the entire length of the body and basal portion of the tail. Distribution, Habitat 5. challengeri is restricted to the McPherson Ra. and its hinterland (Fig. 8). In NSW it occurs in the ranges of the Mt Warning caldera (Mt Warning, Nightcap Ra.), southern edge of the McPherson Ra., and theTooloom and Richmond Ras. In SEQ it occurs throughout the McPherson Ra. from its eastern margin west to Cunningham’s Gap and adjacent Mt Tambourine, and has been recorded from coastal lowland at Beenleigh and Scotts Is. in the lower reaches of the Tweed R. Throughout most of its range ij inhabits closed gully forest from sea level to 500 m. S. chal- lengeri is a conspicuous, surface active, diurnal species that inhabits the forest floor and edges of streams in closed forest, where it can be relatively abundant. It is sympatric with S. galli at two localities. Breakfast Ck at Mt Warning and Sheepstation Ck in the border ranges. Both FIG. 7. S. challengeri, Lamington Plateau, SEQ. 148 MEMOIRS OF THE QUEENSLAND MUSEUM localities are small gully creeks running through closed forest at mid to low (<500 m) altitude. At these sites S. chailengeri is widespread and moderately abundant on the forest floor bordering and adjacent to the streams, whereas 5. galli is rarely encountered away from the stream and, in contrast, difficult to observe, tending to shuttle between sheltering sites and remain secretive, rarely appearing to bask in the open. Saproscincus galli Wells & Wellington, 1985 Figs 9-12 Saproscincus galli Wells & Wellington, 1985:37. Holotype: AMS R1 16964 (AMH 16800) (Fig. 9) Specimens Examined The following specimens were used in the description, and include those used in the canonical variates analysis and electrophoretic analysis (morphological analysis only *; electrophoretic analysis only !). Lamington NP, Toolona Gorge area on Toolona Falls track (population 21-part), 28.15’S, 153.10’E (R140^0-62); Lamington NP, Toolona Falls (popula- tion21-part),28.15’S. 153. 10*E(R 140667, R140674); Border Ras NP, vicinity of Brindle Ck rest area (population 22). 28.22’S. 153.03’E (R138031-33, R 1 38042; R 138043-44*); Mt Warning NP, vicinity of Breakfast Ck carpark (population 20), 28.23’S, 153.17’E (RI33456!; R133458!; R138000-01*, R 1 38007-09, R 1 380 1 6*, R 138070, R 1 38072); Border Ras NP, Lophostemon Falls on Brush Box Falls track (population 23), 28.24’S, I53.0rE (R138047-48, R 1 38049*; R 1 38050-5 1 ); Twelve Sixty Flora Reserve, Coffs Harbour district (population 26). 30.07’ S, 152.55’E (R138207!, R138409-10!); Mobong Falls, Wild Cattle SF (population 25). 30.10’S, 152.47’E (R134986-87, R138191-92); Donrigo NP, Never Never picnic area (population 24), (R138174*; RI38175-76); Cooper’s Park, Bellevue Hill, Sydney (population 27). 33.5rS. 15L17’E (R71622-26*, R71628*, R71709-14*, R93768-69*. R132041-44; R132045*; RI32046, R138404-07; R138408*). Diagnosis The following features in combination general- ly distinguish 5. galli from other members of the 5. chailengeri species group: maximum adult size 59mm; supraciliaries usually 7; lamellae beneath the fourth finger 16-21; lamellae beneath the foulh toe 22-28; presacral vertebrae usually 27; postsacral vertebrae 48-53; dorsal surface a mosaic of lighter and darker scales; ventral sur- face with regularly aligned faint brown spotting tending to form longitudinal rows; lateral and dorsal surfaces of tail barely differentiated, the markings tending to be a continuation of the overall dorsal colour pattern; abdomen of adult males bold lemon yellow. The second, and sixth to tenth characters will distinguish S. galli from S. chailengeri as outlined in the diagnosis of S. chailengeri. The first, third to fourth, and tenth characters will distinguish S. galli from all populations of S. rosei which reach a greater adult size (maximum SVL 64mm). generally have fewer lamellae beneath the fourth finger and toe. and lack bold ventral colour. Characters seven and eight further distinguish S. galli from sympalric southern populations of S. rosei which have a more more uniformly coloured dorsal surface and the ventral surface marked with irregular brown spots. Char- acters five and nine will further distinguish S. galli from sympatric northern populations of S. rosei which have more presacral vertebrae and in subadults and females a bold russet hipstripe which define the dorsal and lateral surface of the TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 149 fll HG. 9. AMS R1 1694 holotype of 5. galli Wells & Wellington. tail anteriorly, this latter character though present in subadult and female S. rosei from southern populations is less bold. Description Measurements: Maximum SVL 59mm; TL 139-192% of SVL (x=165.0%, n=28); AG 53- 62% of SVL (x=56.8%, n=58); HL 35-44% of SVL (x=39.7%, n=57). Scalation: Nasals widely separated; prefrontals moderately to narrowly separated; supraciliaries 6-8 (x=6.8, sd=0.3,n=58); upper labials 6, rarely 7; midbody scale rows 22-24 (x=22.7, sd=0.9, n=58); paravertebral scales 54-61 (x=57.9, sd=L7, n=58), fourth finger scales 9-1 1 (x=10.2, sd=0.4, n=57); fourth finger lamellae 16-21 (x=17.9, sd=L0, n=57); fourth toe scales 11-14 (x=12.3, sd=0.9,n=58); fourth toe lamellae 22-28 (x=24.8, sd=L4, n=58). Osteology: Presacral vertebrae 26-28 (x=27.0, sd=0.3, n=41); postsacral vertebrae 48-53 (x=:50.3, sd=L2, n=31). FIG. 10. S. galli, 6 Lamington Plateau, SEQ. 150 MEMOIRS OF THE QUEENSLAND MUSEUM I i FIG. 11.5. gain, 9 , with white midlateral stripe. Border Ranges, NENSW. Colour and pattern: The various populations of S. gain are similar in colour and pattern (Fig. 10). Dorsal surface mid to dark brown with numerous scattered pale brown to cream scales. Dorsolateral region with dark flecking along scale rows 3 and 4 forming a rough-edged stripe at least anteriorly, and variably continuing partly or wholly along the body and basal portion of the tail. Lateral and dorsal surfaces of tail barely differentiated, the markings tending to be a con- tinuation of the overall dorsal colour pattern. Ventral surface of tail pale with scattered brown spotting, contrasting with darker lateral surface but tending to grade into it at the ventrolateral margin. Head with a bold dark brown loreal streak between the naris and eye and continuous past the eye with the dark dorsolateral stripe. Ventral sur- face white with sparse to dense brown flecks which are generally aligned along the edge of the individual ventral scales, giving the appearance of rough longitudinal streaks on boldly marked individuals. In life adult males have a bold lemon yellow enamel flush to the abdomen. A low frequency of two forms of sexual dichromatism occurs in females of this species, the combinations of which vary between popula- tions. In populations from the Sydney region a small proportion (10-20%) of females have a uniform mid brown dorsal surface bordered by a smooth, well defined, pale edged, dark dor- solateral stripe continuous along the body and tail, contrasting with a darker brown lateral colour of the body and tail. In the McPherson Ranges a small percentage of females have a relatively plain mid brown dorsal surface and boldly con- trasting uniformly darker brown upper lateral sur- face bordered below by a moderately broad, bold white, midlateral stripe (Fig. 11). Distribution, Habitat Saproscincus galli, although widespread is known from a limited number of sites over much of its range. It extends from Mt Tambourine and the McPherson Ra., SEQ to the Sydney region, NSW (Fig. 12). In the northern and central parts of its range it has been recorded mainly from closed forest in gullies. At only one locality in the central part of its range, BeUinger Is., has S. galli been recorded on the coastal plain. In the Sydney region this species is found in remnant patches of low, closed forest in the sandstone hills adjacent to Port Jackson, one of the more densely popu- lated areas in the region. It is also known from urban gardens in near suburbs. Saproscincus rosei Wells & Wellington Figs 13-21 Saproscincus rosei Wells & Wellington, 1985:38. Holotype: AMS R1 16963 (AMH 16801) (Fig. 13) Specimens Examined The following specimens were used in the description, and include those used in the canonical variates analysis and electrophoretic analysis (morphological analysis only *; electrophoretic analysis only !). Conondale Ra., Boolournba Ck xing ca 15km by rd from SF camp (population 14), (R140655-56; R140657*;RI40658-59);MtGlorius,5.6kmN village (population 10), 27.16’S, 152.45’E(R140651-54);Mt Nebo, L5km N village (population 11), 27.33’S, 152.48*E (RI4064546*; R140647^9); Lamington NP, Toolona Falls (population 13), 28.15’S, 153.10’E (R140665; R140666*, R140668-70, R140672-73, R140676-77); Border Ras NP, Tweed Valley Lookout (population 12),28.22’S, 153.05’E(R 133485); Wash- pool NP, Coonibadjha Ck, Coachw'ocxi Pool (popula- tion I-part), 29.28’S, 152.18’E (R1 38098-100; R139101*; R138102); Washpool SF, 5.0km S Hayden’s trig (population 1-part), 29.19’S, 152.18’E TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 151 Washpool SF (population 1-part), 29.23’S, 152.22’E ’ (R96827*, R96877*, R96923-24*); Washpool SF, btn ^ forks of Oorowin road 4km N The Sugarloaf (popula- tion 1-part), 29.24’S, 152.23’E (R96878-79*, R96921*); Washpool SF, off Old Coombadjha road, 9.5km N The Summit (population I -part), 29.25’S, 152.2rE (R96803*); Gibraltar Ras NP, Cedar Valley (population l-part), 29.28*S, 152.20’E (R96792*, R96888*. R96930*); Gibraltar Ras NP, Hakea picnic area (population l*part). 29.28’S, 152. 2rE (R96808*); Washpool, 29.30’S, 152.22’E (R9291 1*); Bruxner P, Coffs Harbour district (population 4), 30.15’S, 153.06’E (R138208!); Wild Cattle Ck SF, Measuring Hut rd, l-6km N Cascade (population 5- part), 30.10’S, 15Z47’E (R138218!); Dorrigo NP, Never Never picnic area (population 2), 30.2 1’S, l52.48’E(RI38163-67; RI38168*); Dorrigo NP, T^e Glades picnic area (population 3-part), 30.22’S, 152.43’E (R1381 84-86); Dorrigo NP, Crystal Shower Falls track (population 3-parl), 30.23’S, 152.43’S (R138180)*; Chaelundi SF (population 5-part), (R135281-83!); Styx R. SF, Softwood rd (population 7), 30.32’S, 152J9’E (R130029-33; R130034*; R130035-36, R130038-39*, R138198-99); Plateau Beech, Werrikimbe NP (population 6), 31.10’S, 152.15’E (R130019-22); Mt Banda Banda Flora Reserve (population 8), 31.10’S, 152.25'E (R13(X)23- 27); Williams R., nr Barrington House (population 9), 32.10’S, 151.3rE (R130069-71, R130780-82; R-130783, R130786-90; R130791*; R138092; R130793*;R130794-95. Diagnosis The following features in combination general- ly distinguish S. rosei from other members of the S. challengeri species group: maximum adult size 64mm; supraciliaries usually 7, occasionally 6; lamellae beneath the fourth finger 14-18; lamel- lae beneath the fouth toe 17-24; presacral ver- tebrae 27-29 (usually 27 in southern populations and usually 28 in northern populations); postsacral vertebrae 46-54; dorsal surface uniform brown (southern jx>puIations) or with a mosaic of lighter and darker scales (northern populations); ventral surface with irregular brown spotting (southern populations) or regular- ly aligned faint brown spotting tending to form longitudinal rows (northern populations); dorsal surface of tail of subadults and adult females with a bold, russet dorsolateral hipstripe; abdomen of adult males usually with a pale yellow wash. See accounts for S. challengeri and 5. rosei for direct comparison between S. rosei and these species. Description Measurements: Maximum SVL 64mm; TL 137-192% of SVL (x=168.1%, n=31); AG 46- 61% of SVL (x=57.1%, n=97); HL 34-44% of SVL (x=38.2%, n=97). Scalation: Nasals widely separated; prefrontals 152 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 13. AMS R1 1693 holotype of S. rosei Wells & Wellington. FIG. 14. S. rosei, 6 , Mt Glorious, SEQ. FIG. 15. S. rosei, 9, Mt Glorious, SEQ. TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 153 FIG. 16. S. roseiy 9, with white midlaleral stripe, Mt Nebo, SEQ. moderately to narrowly separated; supraciliaries 5-8 (x=6.6, sd=0.5, n= 1 0 1 ); upper labials 6, rarely 7; midbody scale rows 22-26 (x=24.1, sd=0.9, n=101); paravertebral scales 55-67 (x=58.8, sd=2.2, n=101), fourth finger scales 8-1 1 (x=9.7, sd=0.8, n=100); fourth finger lamellae 14-18 (x=15.3, sd=0.9, n=100); fourth toe scales 10-14 (x=l 1.2, sd=0.6, n=100); fourth toe lamellae 17- 24 (x=21.0, sd=1.2, n=]00). Osteology: Presacral vertebrae 27-29 (x=27.7, sd=0.7, n=31); poslsacral vertebrae 46-53 (x=49.6, sd=2.3, n=20). Colour and pattern: The following description applies to populations of 5. rosei from MENSW (i.e, Bellinger R. region south to the Williams R.) including the type locality for S. rosei. Variation in colour and pattern elsewhere throughout the species range are discussed with respect to dif- ferences from the type population. In specimens from MENSW the dorsal surface is usually overall mid brown (occassionally lighter or darker), uniform or with a few scattered pale brown scales. Dorsolateral region with dark flecking along scale rows 3 and 4 forming a broken, rough-edged stripe along the body and basal portion of the tail. Females with a poorly defined pale brown to russet hip stripe. Head with a bold dark brown streak between the naris and eye, becoming narrower and obscure behind the eye and generally not continuous with the dark dorsolateral stripe. Dorsal and lateral surfaces of tail in males generally similar, tending to be broken only by fine dark dorsolateral flecking where scale rows 4 and 5 overlap. The darker lateral and pale ventral surfaces of tail are in bold contrast, and defined by an obvious black stripe along the ventrolateral margin. Ventral surface white, with sparse to heavy scattering of mid brown spots. In life adult males generally have a pale yellow flush to the posterior half of the abdomen and a similiar or lx)lder yellow flush to the underside of hindlimbs and basal portion of tail, whereas adult females generally lack such colour or only occasionally have a very weak yellow flush to the very posterior edge of the abdomen, underside of hindlimbs and basal por- tion of tail. A low frequency of sexual dichromatism occurs in populations around the Dorrigo-Bellinger region. Some females had a relatively plain mid brown dorsal surface and boldly contrasting uniformly darker brown upper lateral surface bordered below by a broad, bold white mid lateral stripe. A narrow, pale brown laterodorsal stripe further defined the dorsal and lateral surfaces of the body. Specimens from the Clarence River region north to the NSW-QLD border are intermediate in coloration between MENSW populations and those from SEQ. The dorsal surface is usually marked with a mosaic of lighter and darker scales; 154 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 17. S. rosei, 9, with dark vertebral stripe, Lamington Plateau, SEQ. the russet hip-stripe is bold and well defined and the ventral surface is marked with regularly aligned faint brown spotting tending to form lon- gitudinal rows. Adult ventral colour in life similar to MENSW populations. A colour photograph of an adult female S. rosei from Mt Glorious, SEQ is figured by Wilson & Knowles (1988). The russet hip stripe of subadult and adult females tends to dominate the dorsal coloration. The ventral surface is marked with regularly aligned faint brown spotting tending to form longitudinal rows rather than with sparse to heavy scattering of mid brown spots (Figs 14-16). Adult ventral colour in life similar to central eastern NSW populations. A low frequency of sexual dichromatism oc- curs in most SEQ populations. Some females have a relatively plain mid brown dorsal surface and boldly contrasting uniformly darker brown upper lateral surface bordered below by a broad, bold white mid lateral stripe. A narrow, pale brown laterodorsal stripe further defines the dor- sal and lateral surfaces of the body. Four coloration forms are recorded on the E McPherson Ra. population (Figs 17-20). The most common variant occurs in both sexes where darker flecking on the dorsal surface tends to concentrate down the middle of the body to form a rough, dark vertebral stripe (Fig. 17). The pale ventrd surface is marked with regularly aligned faint brown spotting tending to form longitudinal rows. The yellow ventral colour in live adult males is more prominent than in other popula- tions being present as a moderate to pale flush over the ix)Sterior part of the venter, underside of hindlimbs, and basal portion of tail (where it is boldest), and sometimes extending over most of the venter. It is similarly well developed in live adult females being present as a pale yellow wash to most of the venter in all but one individual. A low frequency of two forms of sexual dichromatism was observed in females and one in males. One female had a relatively plain grey- brown dorsal surface (Fig. 18) the other a mid- dark brown dorsal surface with a mosaic of lighter and darker scales (Fig. 19). Both had boldly con- trasting uniformly darker brown upper lateral sur- faces bordered below by a broad, bold white mid lateral stripe, and bold, russet dorsolateral hip stripes which tend to dominate the dorsal colora- tion. A colour photograph of one of these forms from Lamington NP, SEQ is figured by Wilson & Knowles (1988). One male was completely patternless, being uniform grey-brown (Fig. 20). Distribution, Habitat S. rosei has a broad distribution extending from just south of Gympie (26°irS), SEQ, to the Barrington Tops region (32°10’S) in central east- ern NSW (Fig. 21). Over much of its range S. rosei occurs along the eastern edge of the Great Dividing Ra. and its associated near coastal ran- ges, but is noticeably absent from most of the McPherson Ra. and adjacent Richmond and Tweed Ranges, except for two isolated high al- titude populations in the eastern McPherson Ra. In SEQ, S. rosei occurs on the Great Dividing Ra. along the western edge of the Brisbane R. drainage, and north of the Brisbane R. drainage from the D’ Aguilar, Conondale and Jimna Ran- ges and associated near coastal ranges. It is not known from the more inland Bunya Mts on the Great Dividing Ra.. or from ranges south of the Brisbane R. drainage, except for the two isolated populations in the McPherson Ra. 5. rosei is a conspicuous, surface active, diurnal species that inhabits the edge of closed forest or open, sunlit, patches in the forest. It lends to be more abundant in the former situation where it is commonly observed basking among debris piles. TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 155 FIG. 18. S. rosei, 9, with uniform dorsal surface and white midlateral stripe, Lamington Plateau, SEQ. FIG. 19. S. rosei, 9, with dark flecked dorsal surface and white midlaleral stripe, Lamington Plateau, SEQ. FIG. 20. S. roseiy S y with uniformly coloured dorsal and lateral surface, Lamington Plateau, SEQ. 156 MEMOIRS OF THE QUEENSLAND MUSEUM In MENSW it has been recorded regionally sym- patric with S. galli at a number of locations in the Dorrigo region. On the western edge of the Mc- Pherson Ra.. SEQ. S. rosei has been recorded regionally sympatric with S. challengeri (Cunningham’s Gap) and with S. galli (Mt Super- bus). In the eastern McPherson Ra. it is restricted to high altitude closed forest, being found in creekside vegetation at the headwaters of Toolong Ck, where it was sympatric with S. galli, and from beside a foot-track through closed forest on a ridge on the adjacent Tweed Ra. KEY l.Supraciliaries usually 6; dorsum uniform brown; venter with irregular brown spotting; dorsal sur- face of tail usually with several moderately large, pale vertebral blotches anteriorly S. challengeri Supraciliaries usually 7; dorsum uniform brown or with a mosaic of lighter and darker scales; venter with irregular brown spotting or regularly aligned faint brown spotting tending to form longitudinal rows; dorsal surface of tail without several moderately large, pale vertebral blotches anteriorly 2 2. Dorsum unifonn brown; venter with irregular brown spotting; abdomen of adult males usually with a pale yellow wash S. rosei southern populations Dorsal surface of body with a mosaic of lighter and darker scales; ventral surface with regularly aligned faint brown spotting tending to form lon- gitudinal rows 3 3. Lamellae beneath the fourth toe 22-28; abdomen of adult males bold lemon yellow ; lateral and dorsal surfaces of both sexes undifferentiated S. galli Lamellae beneath the fourth toe 17-24; abdomen of adult males usually with a pale yellow wash; dorsal surface of tail of subadults and adult females with a bold, russet dorsolateral hipstripe S. rosei northern popula- tions ACKNOWLEDGEMENTS Many people contributed to this project through either assisting in field collection of specimens or providing live specimens for tissue samples used in the electrophoretic studies, in particular we would like to thank: Mr Harry Ehmann, Mr James Goodsell, Dr Allen Greer, Mr Harry Hines, Prof. Arnold Kluge, and Mr Alan Manning. The fol- lowing people kindly provided specimens in their care for examination: Dr E.N. Arnold, British Museum of Natural History; Ms Jeanette Covacevich & Mr Patrick Couper, Queensland Museum. Mr Stuart Humphreys photographed the type specimens illustrated in Figs 4, 5, 6, 9, and 13. Ms Tina Goh assisted with typing tables. TAXONOMY, DISTRIBUTION OF SAPROSCINCUS 157 Permits for collection of specimens in NSW were provided by the NSW National Parks & Wildlife Service. In Queensland, permits were issued by the Queensland National Parks & Wildlife Ser- vice of the Department of Enviroment and Heritage. The assistance of Mr Peter Starkey in the Brisbane office with permit applications is appreciated. LITERATURE CITED ADEST, G.A. 1977. Genetic relationships in the genus Uma (Iguanidae). Copeia 1977: 47-52. BOULENGER, G.A. 1887. ‘Catalogue of the Lizards in the British Museum (Natural History). 3.’ (London : British Museum) 575 pp. BUSACK, S.D. 1986. Taxonomic implications of biochemical and morphological differentiation in the Spanish and Moroccan populations of the three-toed skinks, Chalcides chalcides (Lacer- tilia, Scincidae). Herpetologica 42: 230-36. CASE, S.M. & WILLIAMS E.E. 1984. Study of a contact zone in the Anolis distichus complex in the Central Dominican Republic. Herpetologica 40: 118-37. CGGGER H.G. 1975. ‘Reptiles and amphibians of Australia' . (A.H. & A.W. Reed: Sydney). 584 pp. DALE, F. 1973. ‘Forty Queensland Lizards’. (Queensland Museum Booklet: Brisbane). No.8, 64pp. DAUGHERTY, C.H., PATTERSON G.B., THORN C.J., & FRENCH D.C., 1990. Differentiation of the members of the New Zealand Leiolopisma nigriplantare species complex (Lacertilia, Scin- cidae) Heipetological Monographs 4: 61-76. DE VIS, C.W. 1 888. A contribution to the Herpetology of Queensland. Proceedings of the Linnean Society of N.S.W. (2) 2:81 1-826. GREER, A.E. 1974. The generic relationships of the scincid genus Leiolopisma and its relatives. Australian Journal of Zoology - Supplementary Series 31: 1-67. 1980. Critical Thermal maximum Temperatures in Australian Scincid Lizards: their Ecological and Evolutionary Significance. Australian Journal of Zoology 28:91-102. GREER, A.E. & KLUGE, A.G. 1980. A new species of Lampropholis (Lacertilia: Scincidae) from the rainforests of northeast Queensland. Occasional Papers of the Mu.seum of Zoology University of Michigan No.691:l-12. GRIFFITHS,K. 1987. ‘Reptiles of the Sydney region’. (Three Sisters Productions Pty. Ltd.: Winmalee, NSW). 120 ppf HARRIS, H.&HOPKINSON,D. A. 1977. ‘Handbook of enzyme electrophoresis in human genetics.’ (North-Holland: Amsterdam). HEBERT, P.D.N. & BEATON, M.J. 1989. ‘Methodologies for allozyme electrophoresis using cellulose acetate electrophoresis*. (Helena Laboratories: Beaumont, Texas). 32 pp HILLIS, D.M. 1985. Evolutionary genetics of the An- dean lizard genus Pholidobolos (Sauria: Gym- nophthalmidae): Phylogeny, biogeography, and a comparison of tree construction techniques. Systematic Zoology 34: 109-26. INGRAM, G. & RAWLINSON, P. 1981. Five new species of skinks (genus Lampropholis) from Queensland and New South Wales. Memoirs of the Queensland Museum 20(2): 311-17. KIM, Y.J.. GORMAN, G.C.. & HUEY R.B., 1978. Genetic variation and differentiation in two species of the fossorial African skink Typh- losaurus (Sauria: Scincidae). Herpetologica 34: 192-94. MILTON. D.A., HUGHES, J.M., & MATHER, P.B. 1983. Electrophoretic evidence for the species distinctness of Egemia modesta and E. whitii (Lacertilia, Scincidae). Herpetologica 39: 100- 105. S WOFFORD, D.L., & SELANDER, R.B. 1981. BIOSYS-1: a FORTRAN program for the com- prehensive analysis of electrophoretic data in population genetics and systematics. Journal of Heredity 72: 281-83. MURPHY. R.W., COOPER, W.E., & RICHARDSON, W.S. 1983. Phylogenetic relationships of the North American five-lined skinks, Genus Eumeces (Sauria, Scincidae). Her- petologica 39: 2(X)-21 1. NEI, M, 1987. ‘Molecular Evolutionary Genetics.’ (Columbia University Press: New York). 512pp. THORPE, J.P. 1982. The molecular clock hypothesis: biochemical evolution, genetic differentiation and systematics. Annual Review of Ecological Systematics 13: 139-68. THORPE, R.S. 1975. Quantitative handling of charac- ters useful in snake systematics with particular reference to intraspecific variation in the Ringed Snake Natrix natrix (L.). Biological Journal of the Linnean Society 7(I):27-43. WELLS, R.W. & WELLINGTON. R.C. 1985. A Clas- sification of the Amphibia and Reptilia of Australia. Australian Journal of Herpetology - Supplementary Scries I. 1-66. WILKINSON, L. 1987. SYSTAT: the system for statistics. (SYSTAT Inc: Evanston). WILSON, S.K. & KNOWLES, D.G. 1988. ‘Australia’s Reptiles. A photographic reference 158 MEMOIRS OF THE QUEENSLAND MUSEUM to the terrestrial reptiles of Australia’. (Collins: WORRELL, E. 1963. ‘Reptiles of Australia’. (Angus Australia). 447 pp. & Robertson: Sydney). 207 pp. APPENDIX 1 Specimens examined (AM and QM) to prepare the distribution maps of S. challengeri, S. galli, and S. rosei. Saproscincus challengeri Queensland Museum specimens: J303740: J3042; J7497; J12I4344; JI2I51-57; JI216I-70: JI3362-63: J13635; J16456; J18020; J18023; J21997: J22224; J2294I42; J24836; J249I0; J26032-33; J2668I-85; J26723; J27705-06; J27769-71; J27840-42; 328297-99; J30808; 332274-75; 334951-56; 334978-86; 335258-59; 342434-37; 34%29; 349632; 34%36: 349641; 349649; 349662; 349664; 349667; 349686; Australian Museum specimens; R8780; RI5747-48: RI8299; RI830I-02; R18726; R20513-I5; R52745-50: R55088-90; R85947-48; R87048-50: R92I3I-32; R95382: R96440-43; R97819*2I;R97885; R1(M104; R! 11207; R130855; R130857; RI30865; R130867; R130881; RI30905; R131869-80; R132436-37; RI32450; RI32460-61; R132464; R133462; R135464; R138003-04; R138027; RI38062-69; R138088-89; R138092-94; R138265-68; RI38272-78; R139I24-25; R139514-I5; R139521-22; RI40640-44. Saproscincus galli Queensland Museum specimens: 33041; 33043; 313636; 330564-67; 330569-74; 33067 1-72; 332276. Australian Museum specimens: R16997: R18540-44; R18565; R20319; R53742; R54341; R64519; R71627; R71629-32; R76549; R85724; R90604; R906I4; R92I33; R95613; R96588-96; R97817; R97836-46; R104284-87; R104304; Rin73740; Rn8999-n9000; RI27378: RI30018; R132033-37; R132CV47-48; R132434; R132462; R13344849; RI38002; R138010-15; R13803441; RI38052-61; R138071: RI38073-75; RI38177; RI38193-94; RI3841I-12; R139551; R140663-64; R 140675. Saproscincus rosei Queensland Museum specimens: 31581; 33754; 312254; 314318; 316915; 318022; 320229; 320655; 321409; 322471; 322684; 324049-50; 324334-35; 324346; 325534; 326501; J26679;3269 11; 328278; 329035; 329944; 330575; 330809; 332189; 332277;33691 9; 336958; 337934; 3421 22; 349663; 349735; 351620; 351983. Australian Museum specimens; R 169%; R 17002; R43738: R43750; R49I73-77: R52730-44; R53746; R54470-71; R54622-23; R54890-9I; R61 167-68; R61303; R62786; R85938-46; R90603; R96893; R97818; R99475; R103007-08; R1030I2; RI03024-26; R103066; R104I10; R104113; RI07678; RI08693-97; R108705-06; R108738; R108752; RU 1566-69; R1 12253-59; Rn2273;R130037;RI30784-85;R130796-97;R132000-28;RI32049-72;RI32826:R137685;RI377CM;RI377U; R137715-16;R137727:R137753;R137760;RI37763-64;R137868-71;R138162;RI38169;RI3825I-52:R138269-7I;R138279-96;RI38356; R138413-17;R139031;R139054-56;R139063;R139076;R139079;R13908I;RI39097:R139112;R139I38;R139217;R139220;Rl39226-27; R139234;R139236-42;R139271;R139275-77;R139282;R]39284;R139313-15;R139341;RI39346-49;R139366-67;R139484-85:R139489; R139499;RI395n;R139558-63;R13%80-81;RI39702-03;R139786;R139791;RI40650;RI40671;RI41050;R141071;R141395:R141437; R141463; R141544; R14 1552-54; R14I556; RI41622; RI4I653. A NEW SKINK, NANGURA SPINOSA GEN. ET SP. NOV., FROM A DRY RAINFOREST OF SOUTHEASTERN QUEENSLAND J.A. COVACEVICH, PJ. COUPER AND C. JAMES Covacevich, J.A., Couper, P.J. & James, C. 1993 12 24: A new skink, Nangura spinosa gen. et sp. nov., from a dry rainforest of southeastern Queensland. Memoirs of the Queensland Museum 34(1);159-167. Brisbane. ISSN 0079-8835. Naiigura spinosa gen. et sp. nov. is a very distinct member of the Sphenomorphus group. It resembles Gnypetoscincus queenslandiae superficially in being very spiny. It is, however, larger than G. queenslandiae^ has supratemporal fossae (absent in G. queenslandiae), and a karyotype 2n=28 (vs 30). It also has an interparietal which totally separates the parietals. Nangura spinosa is the first reptile species known to be confin^ to the remnant dry rainforests (= semievergreen vine thickets) of southeastern Queensland. It is known only from Nangur State Forest (Figs la,b), which is now, like most semievergreen vine thickets in southeastern Queensland, in the care of the Queensland Forest Service, □ Nangura spinosa, Sphenomorphus group, dry rainforest, Scinidae, southeastern Queensland, J.A. Covacevich, P.J. Couper, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; C. James, Queensland Department of Environment and Heritage, PO Box 3130, Rockhampton Shopping Fair, Rockhampton, Queensland 4701, Australia; 20 July, 1993. The reptiles of the tropical and subtropical moist rainforests (=micro, meso and notophyll vine forests) of Queensland are now relatively well known as far as species and broad distribu- tions are concerned (Covacevich & McDonald, 1991). These species have been the subject of intense taxonomic research in the last decade or so. No less than 1 1 species have been described from Queensland’s moist rainforests since 1980. Many of these species are endemic to Australia and have extremely narrow ranges. By contrast, the reptiles of the dry rainforests (=semievergreen vine thickets, inter alia. Webb, 1978) are not well known. Semievergreen vine thickets occur in a broken band to the west of the better-known moist rainforests of the near coastal uplands and plains. Some reptile surveys were undertaken in semi- evergreen vine thickets in the mid 1970s (Anonymous, 1976; Queensland Museum, 1977). They form an important habitat for some recently described species from north and mid- eastern Queensland (e.g. Anomalopus brevicollis Greer & Cogger 1985; Glaphyromorphus cracens (Greer 1985); andLerato vittata Greer et al., 1985), but endemism is not a feature of their reptile (and other vertebrate) fauna. The reptiles from semievergreen vine thickets surveyed to date are either wide-ranging species recorded from moist rainforests to sparse, open woodlands (e.g. Varanus varius, Rhinoplocephalus nigres- cens, Boiga irregularis), or from purely open forests (e.g. Carlia mundivensis, Carlia schmeltz- a, Heteronotia binoei, Lialis burtonis). In southern Queensland, ‘.... vineforests ... in- clude a number of vegetation communities that are known by a variety of names, including rain- forest scrub, hoop-pine scrub, vine scrub, vine thicket and softwood scrub. These communities have been largely cleared in southern Queensland because of their rich soils and value for agricul- ture. The remai ning stands are remnants ... ‘of less than 1 hectare to large stands of up to SOGO ha, however most are small and less than IGO ha in size’ .... (Forster etal., 1991). Nangur State Forest (Fig. la-b) is a patch of semievergreen vine thick- et near Murgon, SEQ, only 250km northwest of Brisbane. Survey work on the vertebrates of this and other similar forests was undertaken in 1992. The results of those surveys are reported in this volume (Covacevich et al, 1993; Horsup et al., 1993). In June, 1992, one of us (C.J.), working with Mr Mark Fletcher, excavated a burrow ‘which looked as if it might have been made by a lizard’. A very distinct, spiny skink, here described as Nangura spinosa gen. et sp. nov., was found in a small chamber about 60cm from the burrow entrance. N. spinosa is only the second reptile species known to be confined to semi- evergreen vine thicket. (The other is Leris ta vit- tata). It is a large, distinctive skink. That it remained unknown till 1992 and has been dis- covered close to Brisbane, where the reptiles are generally well-known, is remarkable. 160 MEMOIRS OF THE QUEENSLAND MUSEUM A NEW GENUS OF RAINFOREST SKINK 161 N.spinosa ‘fits’ closely, but not perfectly, into the definition of Greer (1979) for members of the Sphenomorphus group. It has 8 premaxillary teeth; an open Meckel’s groove; each parietal bordered posterolateral ly by the temporals; en- larged medial preanals, two rows of scales on the basal half of the dorsal part of the fourth digit; an iris and pupil which are equally dark; a thin bilobed hemipenis. It lacks pterygoid teeth and a postorbital bone. (The latter is present in some members of the Sphenomorphus group, e.g. Eulamprus spp. and absent in others, e.g. Ctenotus spp. Its presence is regarded as primitive for the group, Greer, 1979; Hutchinson, pers. comm. In N.spinosa the fonner postorbital has been incorporated in the postfrontal. A short, incomplete suture is the only remaining trace of this bone.) N.spinosa is secretive, has a scaly eyelid and fully-developed pentadactyl limbs, and appears not to be sexually dimor- phic/dichromatic (our sample size is of five specimens only). The latter suite of characters used also by Greer (1989) for the Sphenomorphus group are primitive characters for all skinks, and are generally true for the Sphenomorphus group (M. Hutchinson, pers. comm.). N.spinosa has greatly enlarged preanal scales, an iris virtually as dark as its pupil, and a bifurcate hemipenis - a combination present only in members of the Sphenomorphus group (Hutchinson, pers. comm.). In two characters N.spinosa is unusual amongst members of the Sphenomorphus group. Nangura is the only member of the Spenomorphus group karyotyped to date that has 2n=28. (Genera karyotyped include Anomalopus, Calyptotis, Ctenotus, Eremiascincus, Glaphyromorphus, Gnypetoscincus, Hemiergis, Lerista, Ltpinia, Lobulia, Notoscincus, Papuascincus, Prasino- haema, Saiphos and Sphenomorphus). All, ex- cept Lipinia (which has 2n=42), have 2n=30 (which is seen also in Scincella laterale of the USA). Karyotype thus strongly emphasises the uniqueness of N.spinosa in comparison with Gnypetoscincus queens landiae and Eulamprus spp. with which it shares some characters (Don- nellan, 1985; Donnellan, pers. comm.). Further, its parietal scales are totally separated by the interparietal. This character is present in only one species group (Eulamprus quoyii) of the Sphenomorphus group. It seems more reasonable to expand definitions of Greer (1979, 1989) slightly to include N.spinosa in the Sphenomor- phus group, than to separate it from the group because of these distinctive features. The most striking feature of N.spinosa is the presence of well-defined spines all over its dor- sum. In this feature and in general body propor- tions, it bears some resemblance to G. queenslandiae (of the Sphenomorphus group) and to some of the Egernia species (of the Mabuya group). In the Sphenomorphus group, Greer (1989) recognises two subdivisions based on ecology, surface-dwelling species (e.g. Ctenotus spp.) and semi-fossorial or fossorial species (e.g. Coeranoscincus spp.). He notes the unique case of G. queenslandiae which, despite its surface-dwelling habits, is a sedentary lizard that spends much of its life under rotting rain- forest logs. He places it with semi-fossorial or fossorial species. The following characters are used: snout to vent length (SVL); tail length (T); length of front leg (LI); length of hind leg (L2); head length (HL); head width (HW). Character definitions and measurements follow Couper et al., 1993. Nangura gen. nov. Head scales rugose. Most other dorsal and lateral scales keeled, and not, or only just over- lapping. Keels on the tail are most pronounced, almost pyramidal. Dorsal keels form longitudinal lines. Axillal and inguinal scales bead-like to granular. Ventral scales weakly carinate. Iris very dark, almost as dark as the pupil which is black. Rostral in broad contact with the frontonasal, which thus separates the nasals widely. Prefron- tals large, moderately spaced. Frontal elongate, almost 1.5 times as long as broad. Fronto-parie- tals in broad contact, and in contact with supraoculars 2, 3 and 4. Interparietal elongate, almost rectangular, about twice as long as broad, and about as long as the parietals. Parietals widely separated by the interparietal and bordered by two temporals. Supraoculars 4. Supraciliaries 5-7. Suboculars in a distinct row, in contact with supralabials and granules of the lower eyelid. Lower eyelid scaly, without a disc. Loreal scales 2, the posterior one the larger. Ear opening large, about three times as long as wide; vertical, lack- ing lobules, and with a superficial tympanum. FIG. la Nangur State Forest from the track bisecting it, showing low, closed canopy of typical semievergreen vine thicket.b Bed of small unnamed creek in Nangur State Forest. All known specimens of N.spinosa h^\e been collected or seen on the banks of this creek.c-d Entrances of burrows of N.spinosa. e N.spinosa (J57247). 162 MEMOIRS OF THE QUEENSLAND MUSEUM FIG.2 Ectopterygoid, pterygoid and palatine contacts in (a) N.spinosa (J57247), (b) Gnypetoscincus queenslan- dice (J56824). Medial pairs of preanal scales greatly enlarged. Dorsal scales of the fourth toe paired basally. Palatine bones in broad medial contact. Palatal rami of the pterygoid bones narrowly separated. Ectopterygoid bones contacting both the palatines and the pterygoids (Fig. 2a). No postor- bital. Supratemporal fossae present. Eight conical teeth on the premaxilla. Pterygoid without teeth. Meckel’s groove open (Fig, 3). Karyotype 2n=28. Hemipenis smooth, bilobed. Nangura spinosa sp. nov. (Figs 1-5) Material Examined Holotype: QMJ55424 ?, Nangur State Forest (SF74), 26°07\ 151°58’) SEQ, C. James & M. Fletcher, 5 June 1992. Parattpes: QMJ56029 9, J56031 , J57246-7 , Nan- gur State Forest, SEQ. Diagnosis N.spinosa is distinguished from other members of the Sphenomorphus group (except some Eulamprus spp., all of which lack keels)by the state of the parietal shields (parietals not in con- tact behind the interparietal vs parietals in contact behind the interparietal). It is distinguished from G. queenslandiae, the species to which it bears most resemblance, by midbody scale count (28- 30 vs 32-36, Cogger 1992); and by karyotype 2n (28 vs 30);degree of caudal keeling (very pro- nounced vs not pronounced); degree of lateral scale overlap(some vs none); supratemporal fos- sae (present vs absent); contact between ectop- terygoid, palatine and pterygoid bones (ectopterygoid contacting both vs contacting only the pterygoid, Fig 2a,b); and size (max SVL 95.10 mm vs 90.80 mm). As Nangura and Gny- petoscincus are monotypic, the characters which separate them generically also separate them at the species level. (The latter was redefined by Ingram, 1985, following separation of the Australian 'Tropidophorus' queens landiae from Southeast Asian Tropidophorus spp., as Gny- petoscincus queenslandiae. Wells and Wel- lington, 1985). Distribution Known only from the type series. All specimens were collected in Nangur SF (SF74), approximately 20km north of Murgon, SEQ. Habitat Semievergreen vine thicket on dark, basaltic soils. Forster et al,, 1991 describe Nangur State Forest as having ‘canopy intact, exotic weeds present ....’. Estimates of the size of the forest range from < 5(X) ha (Forster et al., 1991) to 822 ha (P. Flower, pers. comm.), Holotype SVL (mm): 88.9. Proportions as % SVL: T = 85.0, LI = 30.2, L2 =38. 1 , HL = 24.5, HW = 1 6.9. Very spinose, dorsal and lateral scales, save those of the head (which are rugose) and of the axillal and inguinal area (which are bead-like to A NEW GENUS OF RAINFOREST SKINK 163 FIG, 3 Skull of N. spinosa (J57247)- granular), keeled.(Fig. 5). Keels most pronoun- ced, almost pyramidal, laterally at the base of the tail; strongly developed dorsally and laterally (in- cluding the limbs); forming pronounced dorsal longitudinal ridges. Scales of the venter weakly carinate. Midbody scale rows, 30. Paravertebral scales, 40. Preanal scales 4, the middle two great- ly enlarged. Head: moderate, with rugose scalation on dor- sal and lateral surfaces; parietals not in contact behind interparietal; interparietal free, elongate; frontoparietals paired, in broad contact; fron- tonasal single; prefrontals large, moderately spaced; frontal elongate, almost 1.5 times as long as broad; nasals separated by broad rostral/fron- tonasal contact; supralabials 6; infralabials 4, 2nd largest; supraoculars 4, 2nd largest; supraciliaries 6/7; loreals 2, 2nd largest; presuboculars, 1; sub- ocular scales between the supralabials and scales of the lower eyelid, 3; temporals, an irregular series of small scales; eye dark, pupil not clearly defined; lower eyelid scaly, lacking palpebral disc; ear opening large, vertical without lobules; tympanum superficial, 1st and 2nd chin shields enlarged, 3rd chin shields divided Limbs: well developed, markedly overlapping when adpressed; plantar scales rounded, slightly keeled; 4th toe markedly longer than 3rd toe; subdigital lamellae, 16/17, bluntly keeled. Tail: cylindrical, tapering to a point; subcaudal scales slightly enlarged in relation to those ad- joining them; vertebral scales not enlarged. Colour: dorsal surface brown with irregular black crossbands from nape to base of tail; venter cream; lips pale, heavily barred with black; upper lateral zone predominantly black, broken by cream bars; flanks cream with dark flecks. 164 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4 a-b Nangura spinosa. (J57246, J57247) Paratypes SVL (mm): 84.2-95.1 . Proportions as % SVL: T 88-94 (n3, mean 89.8); LI 30-33 (n4, mean 31.9), L2 37-42 (n 4, mean 39.7), HL 24-29 (n4, mean 26.8), HW 1 7-20 (n4, mean 1 8.7). Midbody scale rows, 28 (J57246-7), 29 (J56031), paraver- tebral scales, 38 (J57247), 41 (J56029). Head: infralabials 3, right side only, 3rd largest A NEW GENUS OF RAINFOREST SKINK 165 (J57246); supraciliaries 5, right side only (J56031); subocular scales between the supralabials and scales of the lower eyelid 2, right side only (J56029, J56031); 2, left side only (J57246). Limbs: subdigital lamellae, 15/15 (J56029), 18/17(156031), 16/16(157246), 15/13(157247); lamellae broadly callose (J56031, J57246). Field Notes All known N.spinosa specimens have been col- lected from burrows in dry, hard, black, basaltic soil. J57246 was taken from a burrow 33.5cm long, with a diameter of roughly 2.0cm. Other burrows excavated have ranged to 60cm. The entrances of burrows are concealed generally by association with tree bases or surface roots (Fig. lc,d). One (the first found and excavated by C.J. and M. Fletcher) was in the open, several metres from any cover. Two of four burrows excavated terminated in an oval chamber with a length of about 5cm. N.spinosa occurs in small colonies. The type locality has been searched methodically and ex- tensively. All burrows known lie on the gently sloping bank of a seasonal, unnamed small creek, above what we surmise to be the wet season water flow level, but close to it in one case. Two colonies of solitary or near solitary specimens were found 500 metres apart. (All but one burrow housed only one adult N.spinosa. The exception was ‘home’ to two specimens - an adult observed, and a neonate ‘fished’ from the burrow using a meal worm as bait and a cotton thread as line. 14 April 1993. This specimen was released after being measured, S VL 4. 1 cm, T 3.4cm. It retained an ‘umbilicaT scar). Burrows located are roughly 20-30 m apart. In June, 1992, and April, 1993 the ground was dry and very hard. No sign of digging was seen, and each burrow appeared to have a smooth resting platform of about 6x4cm. The platforms appear to be compacted. In November, 1992, C.J. observed that burrows appeared to be ‘active’. Small screes of friable soil near burrow entrances suggested digging was either current or very recent. With each burrow there is a defeca- tion site about 30cm from the burrow entrance. As well as faeces, fragments of shed N.spinosa skin were found on lhe.se sites. Faeces found in April, 1993, contained hemip- teran abdomen, thorax, other remains, coreid (Pachycolpura) remains; coleopteran legs, scarab elytra, carabid beetles (Notonomns sp., Crasp- edophorus sp.), dung beetle {Canthonosoma sp., Cephalodesminus sp.) remains, ground weevil FIG 5. Head scales of the holotype (J55424) of Nan- gura spinosa. remains, click beetle remains, a large cricket head; large ant remains, a wasp head; millipede remains, spider legs. Dr. G.B. Monteith who identified the remains, notes ‘.... The sample was of a diverse range of well-chewed anthropod remains. No plant material was present. Most common were remains from beetles and spiders, with fewer from bugs, millipedes, ants and a cricket. The largest animal in the sample is the cricket which would have measured 3-4cm. Some of the prey animals have very distasteful defence secretions (e.g. millipedes; carabid beetles; and the coreid bug, Pachycolpura sp.). Some of the animals found would be expected to be nocturnal (e.g. Canthonosoma sp., Notonomns sp. and the cricket). All taxa represented in the sample are forest-floor dwellers '.All would be wanderers and thus susceptible to ambush by N.spinosa (G. Monteith, pers. comm.). Of the two females, J55424 collected in winter (June, 1992), has slightly enlarged follicles. J56029 collected in spring (September, 1992) also has enlarged follicles. These are larger than those of J55424. Members of the Sphenomorphus group are both egg-layers (many Glaphyromor- phus spp.) and live-bearers (all 8 spp. of Eulamprus examined and G. queenslandiae). 166 MEMOIRS OF THE QUEENSLAND MUSEUM Greer, 1989. N.spinosa is not typical of the Sphenomorphus group. It has some features in common with the latter, and the combination of burrow-dwelling habits and a very young specimen in a burrow with an adult suggest that N.spinosa is probably a live-bearer. CONSERVATION By any standard, a species known from only five type specimens from a single locality, which is a remnant of a forest type previously widely distributed, is in need of special conserva- tion/management attention. Under the system devised by Thomas & McDonald (1989) and applied to all then-known species of reptiles in Queensland by McDonald el al. \99\j N.spinosa is a ‘species known only from the type collection’(l), and is ‘rare in Australia, but not currently considered endangered or vul- nerable....’ (R). N.spinosa has already received special atten- tion. Nangur State Forest is protected from further clearing. In addition, following the discovery of N spinosa , the Queensland Forest Service took action to ‘minimise disturbance to the animal’s habitat...’ (T. Ryan, in litl., 23 September, 1993) by restricting access, applying special precau- tions regarding fire prevention, and advising staff to strive to locate more colonies of N.spinosa. Like all other reptiles in Queensland, N.spinosa is fully protected under the The Nature Conserva- tion Act of 1992 and The Nature Conservation Regulations, 1993. Two different approaches can be followed regarding the conservation of N.spinosa. A non- interventionist approach is based on the fact that this species has survived much alteration to its habitat. (It is an extremely secretive species, vir- tually impossible to see accidentally and this un- doubtedly accounts for its remaining unrecognised till 1992). Nangur State Forest has been subjected to many assaults, which N.spinosa has survived. Much of the original Nangur semi- evergreen vine thicket was cleared for agricul- ture, leaving only a core area of about 5()0ha. This has been bisected by a road, grazed by cattle, and altered on a smaller scale by fires, timber-getting and invasion by Bufo marinus, Sus scrofa, and many weeds. The second approach could evolve from recog- nition of a second important fact about N.spinosa. It is, at least as far as present knowledge is con- cerned, rare and extremely narrowly restricted. Should research effort to extend knowledge of its ecology with a view to devising an ‘active management’ scheme be treated as a matter of urgency? Rare, endangered and threatened species research is a very popular field at present (along with research on rainforest taxa). N.spinosa qualifies on both fronts for a special place in the current hierarchy of ‘worthwhile’ research projects. Perhaps it can be considered extra ‘worthy’, because semievergreen vine thickets are a very poorly known type of rain- forest, herpetologically and otherwise. We favour a thorough documentation of the life history, oc- currence and habitat requirements of this species, along with continued measures to ensure total protection of its habitat. ACKNOWLEDGEMENTS We have received generous assistance in the field from Terry Beutel, and Steve Wilson; in the laboratory from Anita Heideman and Lauren Keim; and in discussion about the status of N.spinosa or its habitat from Michael Cunnin- gham, Steve Donnellan, Alan Horsup, Mark Hutchinson and Glen Ingram. Photographs for this description were taken by Bruce Cowell, Gary Cranitch and Jeff Wright. Identifications of insect remains in the faeces of N.spinosa were provided by Geoff Monteith. The karyotype was determined by Craig Moritz. Most of the field work for this description was funded by the Na- tional Rainforest Conservation Programme, ad- ministered by the Queensland Department of Environment and Heritage. LITERATURE CITED ANONYMOUS, 1976. Fauna of eastern Australian rainforest: preliminary report on sites surveyed by the Queensland Museum in mid-eastern and northeastern Queenland.(Queensland Museum: Brisbane). 78pp. COGGER, H.G. 1992. ‘Reptiles and Amphibians of Australia’. (Reed Books: Chatswood, Sydney). 776pp. COUPER, P.J., COVACEVICH, J. & MORITZ, C. 1993. A review of the leaf-tailed geckos endemic to eastern Australia: a new genus, four new species and other new data. Memoirs of the Queensland Museum 34(1): 95-124. COVACEVICH,!. & McDONALD, K.R. 1991 . Frogs and reptiles of tropical and subtropical eastern Australian rainforests: distribution patterns and conservation. Pp.281-309 In Werren, G. & Ker- shae, P. (eds). ‘The rainforest legacy, Australian A NEW GENUS OF RAINFOREST SKINK 167 National Rainforests Study, Volume 2 - Flora and Fauna of the rainforests. Australian Heritage Publication Series Number 7(2)\ (Australian Government Publishing Service: Canberra). 414 PP- COVACEVICH, J., COUPER, PJ., &1NGRAM, GJ. 1993. New reptile records from rainforests of south and mideastem Queensland. Memoirs of the Queensland Museum. 34(1):185-188. DONNELLAN, S. 1985. The evolution of six chromosomes in scincid lizards. (Unpublished PhD thesis. Macquarie University, Sydney). 142pp FORSTER, P.I., BOSTOCK, P.D., BIRD, L.H. & BEAN, A.R. 1991. ‘Vineforesl plant atlas for south-east Queensland. An assessment of conser- vation status’. (Queensland Government, Queensland Herbarium: Brisbane).466 pp. GREER, A. 1979. A phylogenetic subdivision of Australian skinks. Records of the Australian Museum. 32(8): 338-71, Figs. 1-18. GREER, A.E. 1989. ‘The Biology and evolution of Australian lizards’. (Surrey Beatty and Sons Pty Ltd: Chipping Norton, Sydney). 264 pp. HORSUP, A., JAMES, C., & PORTER, G. 1993. Surveys of rainforest vertebrates, south and mideastern Queensland. Memoirs of the Queensland Museum. 34(1): 215-228. McDonald, k.r., covacevich, j.a. Ingram, G.J. & COUPER, P.J. 1991. The status of frogs and reptiles, Pp. 338-345. In Ingram, G.J. and Raven, R.J. (eds). An atlas of Queensland’s frogs, reptiles, birds and mammals. (Board of Trustees, Queensland Museum: Brisbane). 391 pp. INGRAM, G J. 1985. Implicit technique in taxonomy: the scincid lizards of Cape York Peninsula. (Un- published PhD Thesis. University of Queensland: St Lucia, Brisbane).366 pp. QUEENSLAND MUSEUM, 1977. Fauna of eastern Australian rainforests II. Preliminary report on sites surveyed by the Queensland Museum in southeastern and far northeastern Queensland, with additional results from sites surveyed pre- viously in northeastern Queensland. (Queensland Museum: Brisbane). 102pp. THOMAS, M.B. & McDONALD, W.J.F. 1989. Rare and threatened plants of Queensland. 2nd ed. (Department of Primary Industries: Brisbane). WEBB, L.J. 1978. A general classification of Australian rainforests. Australian Plants 9: 349-363. WELLS, R.W. & WELLINGTON, R.C. 1985. A clas- sification of the Amphibia and Reptilia of Australia. Australian Journal of Herpteolgy. Sup- plementary Series No. 1 -61 . (Australian Biologi- cal Service: Sydney), pp.64. 168 MEMOIRS OF THE QUEENSLAND MUSEUM THE ULTRASTRUCTURE OF SPERMATOZOA OF NANGURA SPINOSA (SCINCIDAE, REPTILIA) B.G.M. JAMIESON AND D.M. SCHELTINGA Jamieson, B.G.M. & Scheltinga, D.M. 1993 12 24: The ultrastructure of spermatozoa of Nangura spinosa (Scincidae,Reptiiia). Memoirs of the Queensland Museum 34(1) 169-179 Brisbane. ISSN 0079-8835. Spermatozoa of Nangura spinosa are filiform, and approximately 85 pm long. The acrosome vesicle in the form of an elongate hollow, concentrically zoned cone is compressed near its tip, and basally overlies a subacrosomal cone. Axial within the acrosome vesicle is a slender rod, the putative perforatorium. The subacrosomal cone is paracrystalline and invests the tapered anterior end of the nucleus. The perforatorium is a slender, slightly oblique rod extending anteriorly from the subacrosomal material. A conspicuous laminated structure forming a wing-like projection on each side of the proximal centriole contacts with the first of the dense ‘ring structures’ of the midpicce. The midpiece contains four dense ring structures in longitudinal succession, pexsterior to which lies the much smaller annulus, all being sep^ated by mitochondrial regions. The mitochondria mostly form 12 or more elongate, sinuous columnar structures, with numerous predominantly longitudinal cristae. Nine peripheral dense fibres are associated w ith the 9 triplets of the distal centriole and the doublets of the axoneme. 1 lowever, within the midpiece, only those peripheral fibres adjacent to doublets 3 and 8 remain conspicuous to the level of the annulus, each as a double structure associated with the annulated fibrous sheath. All peripheral fibres iue absent from the principal piece. Similarity to the sperm of Ctenotus supports placement of Nangura in the Sphenomorphus group. Comparisons with other amniote sperm are made. □ Nangura spinosa, Scincidae, spermatozoon, ultrastructure, phylogeny. B.GM. Jamieson and D.M, Scheltinga, Department of Zoology, University of Queensland, Queensland 4072, Australia; J4 October, 1993. Published descriptions of the male gametes of the Scincidae are limited to a description of the mature spermatozoon of Chalcides ocellatus tiligugu by Furieri (1970); an account of sper- miogenesis, with some description of mature, epidydimal sperm, in the same subspecies (Car- cupino et al., 1989); and a very brief account of the development of the midpiece in Eumeces laticeps by Okia (1990). A description of the spermatozoon of the newly described genus and species, Nangura spinosa (see Covacevich et al., 1993), allows the addition of morphological char- acters from the spermatozoon to the description of this species and extends knowledge of skink spermatozoa. Comparison with the sperm of other reptiles will be limited chiefly to that neces- sary to determine which characters or character states appear to be, from the small sample, dis- tinctive of the Scincidae and of this species. Al- though the Reptilia is an invalid, paraphyletic grouping (e.g. Jamieson & Healy, 1992), the term ‘reptile’ is here retained for convenience. The ultrastructure of spermatozoa or sper- miogenesis has been studied in the major groups of squamate reptiles, in addition to skinks: Lacer- tidae - Butler & Gabri (1984), Courtens & Depeiges (1985), Furieri (1970); Agamidae - Al- Hajj et al. (1987), Dehlawi et al. (1992), Chamier et al. (1967); Chameleonidae - Tuzet & Bourgat (1973); Iguanidae - Furieri (1974), Saita et al. (1988a); Anolidac - Clark (1967); Gekkonidae - Furieri (1970), Phillips & Asa (1993); Teiidae - Del Conte (1976), Newton & Trauth (1990, 1992); Tropiduridae - Da Cruz-Landim & Da Cruz-Hofling (1977), Furieri (1974); and Serpen- tes - Austin (1965), Boissin & Mattel (1965, 1966), Furieri (1970), Hamilton & Fawcett (1968), Saita et al. (1988b), Phillips & Asa (1993). Non-squamate reptiles which have been inves- tigated are: Chelonia - De et al. (1987), Furieri (1970), Hess et al. (1991), Sprando et al. (1988), Yasuzumi & Yasuda (1968), Yasuzumi et al. (1971); Sphenodontida - Healy & Jamieson 0992), Jamieson & Healy (1992); and Crocodilia - Saita et al. (1987). Material and Methods Small pieces of testis were taken from a single specimen of Nangura spinosa Covacevich, Couper and James, 1993, collected from Nangur State Forest, near Murgon, SEQ. These samples 170 MEMOIRS OF THE QUEENSLAND MUSEUM were diced into l-2mm^ portions, and fixed for transmission electron microscopy (TEM), in 3% glutaraldehyde in 0.1 M sodium phosphate buffer (pH 7.2), at 4°C for 2 hours and agitated for the first hour. The material was then rinsed in 0.1 M phosphate buffer; post-fixed for 80 min in similarly buffered 1% osmium tetroxide; rinsed in buffer; dehydrated through an ascending ethanol series; and infiltrated and embedded in Spurr’s epoxy resin. Sections were cut with diamond knives, on an LKB 2128 UM IV microtome. Thin sections, 50-80 nm thick, were collected on carbon stabilized, colloidin-coated, 2(X) |im mesh copper grids, rinsed in distilled water, stained for 30 s in Reynold’s lead citrate, then in 6% aqueous uranyl acetate for 4 min and for a further 2 min in lead citrate before final rinsing. Electron micrographs were taken on an Hitachi 300 electron microscope at 75 kV and a JEOL 100-s electron microscope at 60 kV. Light microscopic observations of spermatozoa, from glutaraldehyde-fixed tissue squashes, were made under Nomarski contrast using an Olympus BH2 microscope. RESULTS Spermatozoa of Nangura spinosa (Fig. 1) are filiform, and approximately 85 |im long (mean of 10 = 85.4 )am, S.D = 2.8). A cytoplasmic droplet seen in some sperm, by light microscopy and by transmission electron microscopy, is located im- mediately behind or slightly overlapping the base of the nucleus. Dimensions (for one or two sperm) are: 5.5 |j,m for the length of the acrosome com- plex; 6.6 pm for the nucleus posterior to the acrosome;7.6 )am for the midpiece, from trans- mission electron microscopy, and, from light microscopy 66 |xm for the flagellum behind the midpiece (principal piece). The head (acrosome and nucleus), and often the midpiece and flagel- lum, is curved (Fig. 2A). As a result of this curvature it has not been possible to obtain a complete longitudinal section through the head. The sperm is circular in cross section with the exception of the acrosome. Although the acrosome is circular at its base, anterior to this it develops a unilateral ridge and anterior to the tip of the subacrosomal cone it becomes increasingly compressed and elliptical in transverse section (Figs 2D-I). FIG. 1 Nangura spinosa. A diagrammatic summary of the spermatozoon viewed in longitudinal section, with the corresponding transverse sections. SPERMATOZOA OF NANGURA SPINOSA 171 Acrosome Complex The acrosome complex consists of an acrosome vesicle in the form of an elongate hollow cone, an underlying subacrosomal cone and, axial within the acrosome vesicle, a slender rod, the putative perforatorium. The acrosome complex is 5.5 pm long (Fig. 4A). The anterior end of the vesicle, comprising slightly less than half its total length, forms a thick walled hollow cone with a narrow lumen housing the perforatorium (Fig. 2B). The longer, posterior region of the vesicle is a thin walled continuation of this hollow cone, the vesicle here being no more than a sleeve-like investment (acrosome sleeve) of the sub- acrosomal material, as seen in longitudinal (Figs 2B, K, 4A) or transverse section (Figs 2F-I). The underlying subacrosomal material forms a thick relatively pale layer. The material of the subacrosomal cone is paracrystalline, its matrix having fine obliquely longitudinal and less distinct transverse stria- tions, indicating that it forms a fine lattice (Fig. 2C). For most of its length, from its posterior end anteriorly, the subacrosomal cone invests the tapered anterior end of the nucleus (nuclear point). Although corresponding with the sub- acrosomal cone of other amniote and amphibian sperm, it is not strictly conical but consists of material with a poorly defined outer border which fills the posterior space within the acrosomal sleeve (Figs 2B, C, F-H, K). The nuclear point terminates within the anterior limit of the sub- acrosomal material at an epinuclear electron lucent region (Fig. 2K). The perforatorium is a slender, moderately electron dense rod, with some signs of internal longitudinal fibres. It extends anteriorly from the subacrosomal material, lying in a narrow lumen internal to the inner acrosome membrane (Figs 2B, D, 4A). It has been observed to extend through approximately the posterior half of the thick walled part of the acrosomal vesicle. Whether or not a pale, central axial tube-like structure which extends to the tip of the acrosome and displays some internal longitudinal fibres is a forward continuation of the perforatorium has not been determined (Fig. 2B). The perforatorium makes contact at its posterior end with the sub- acrosomal material. Even allowing for the pronounced curvature of the acrosome, the lon- gitudinal axis of the perforatorium appears to be slightly oblique relative to that of the acrosome vesicle (Figs 2B, D, 4A). In transverse sections of the acrosome vesicle through the nuclear point and perforatorium (Figs 2D-I), the vesicle is seen to have a concentric zonation which in sequence from the per- foratorium outwards is: a narrow space around the perforatorium; a wide, dense, homogeneous zone; a narrow zone with radial striations; a thin, dense, homogeneous layer apposed to the plasma membrane. Nucleus The nucleus is curved and tapers to a point within the basal region of the acrosome (acrosome sleeve). The transition from the tapered region (nuclear point) to the much longer cylindioid region is abrupt but the ‘shoulders’ seen in many other reptile sperm are represented only by a gentle curvature on each side. The length of the nucleus from the base of the acrosome vesicle to the base of the nucleus is 6.6 pm with a further 2.9 pm for the nuclear point which is surrounded by the subacrosomal cone (Fig. 2K). The nucleus is almost parallel sided, showing only a slight increase in width posteriad, from 0.7 to 0.9 pm, reaching its greatest width shortly before its posterior end. The cross section of the nucleus is circular throughout (Figs 2F-J, 3C). The chromatin is condensed and strongly electron dense. Basally the nucleus has a compact conical fossa which houses dense material ex- tending from the proximal centriole (Figs 3A, C, L, 4B, D). Neck Region The neck region (Figs 3A, L, 4B, D) is the region where the nucleus joins the midpiece and is here recognized by virtue of its internal com- ponents although the anterior end of the midpiece, as here defined, directly abuts the posterior end of the nucleus. The neck region includes the proximal and distal centrioles and associated den- sities, including the first of the ring structures of the midpiece. Each centriole consists of 9 triplets. The proximal centriole lies immediately anterior to the distal centriole and with its long axis at slightly less than a right angle to it (Fig. 4B). The long axis of the distal centriole, which forms the basal body of the flagellum, is in the long axis of the axoneme. The centrioles do not lie in the basal nuclear fossa but the proximal centriole, immedi- ately behind this, is surmounted by a hollow conical density (dense cone) which conforms in shape with the nuclear fossa which it occupies. An electron lucent space separates it from the wall of the fossa (Figs 3C, L, 4B, D). Compact dense material extends from the base of the dense cone to cover the more axial end of the proximal 172 MEMOIRS OF THE QUEENSLAND MUSEUM centriole and insinuates itself as a large mass between the proximal and distal centrioles (Figs 3A, L, 4B, D). The two central singlets of the axoneme extend anteriad at least into the region of transition between the distal centriole and the axoneme. In this region there is a density connect- ing triplet 3 with the adjacent central singlet in addition to the peripheral dense fibre connected to each triplet or doublet (Fig. 3D). The peripheral dense fibre at doublet 3 is detached in some sections, indicating the commencement of the corresponding longitudinal column. A conspicuous stratified laminar structure forms a wing-like projection on each side of the proximal centriole, near its anterior limit and is continuous around its axial pole (Figs 3A, L, 4B, C, D). It is therefore seen in some longitudinal profiles of the sperm, which are parallel to but not through the long axis of the axoneme, as a con- tinuous wide lamina spanning much of the width of the nucleus behind the nuclear fossa (Fig. 4C). It is deduced, therefore, that the lamina forms a thick disc around the proximal centriole but that the disc is interrupted at the peripheral end of the proximal centriole. Evidence for this interpreta- tion is also seen in some transverse sections of the neck through the proximal centriole (Fig. 4E). The outer edges of the laminar structure make contact with the first of the dense ‘ring structures' of the midpiece, described below (Figs 3A, L, 4B, C, D) as does the peripheral end of the proximal centriole (Figs 4B, E). Midpiece The midpiece includes the neck, described above. It consists of mitochondria, ring structures and the contained axoneme with its fibrous sheath and ends posteriorly with the annulus. There are four ring structures (rs 1-4) in lon- gitudinal succession, posterior to which lies the much smaller annulus (an). The ring structures, with the annulus, are separated by mitochondrial regions (mi 1-4). In terms of the pattern recog- nized for the teiid lizard Cnernidophorus by New- ton and Trauth (1992), the fonnula for Nangura is rsl/mil, rs2/mi2, rs3/mi3, rs4/mi 4, an. Each ring structure appears in longitudinal section as an approximately kidney-shaped density on each side of the fibrous sheath of the axoneme (Figs 3A, N, 4D). The profile on one side is staggered relative to that on the other, though always over- lapping it, but in transverse section the ring is complete, however, when sectioned near its anterior or posterior borders interruption to the ring can be seen (Figs 3E, 4F). This indicates that each structure is a ring which is tilted relative to the axonemal axis. The mitochondria mostly form elongate, sinuous columnar structures, with numerous predominantly longitudinal crislae, each of which extends from one ring structure to the next (Figs 3A, B, N, 4D). There are 12 or more around the axoneme as seen in transverse section (Fig. 3F), Occasional single, ovoid mitochondria are seen. A few small mitochondrial profiles are some- times present lateral to the ring structures, the outer surface of which may be scalloped by them (Figs 3E, 4F). The axoneme has the usual 9+2 pattern. Each doublet has two dynein anns. The A subtubule is occluded by dense material. Around the axoneme almost as far anteriorly as its junction with the distal centriole, there is a fibrous sheath. In lon- gitudinal section (Figs 3A, B, M, N, 4D) the fibrous sheath exhibits rather regularly arranged, ' approximately square dense blocks which, from glancing longitudinal sections (Figs 3L, N) and transverse sections (Figs 3E-I) are shown to form rings around the axoneme. They show a tendency | to tilt relative to the axonemal axis and there are interruptions in the cross sections but that they form a spiral is questionable. Occasional anas- tomoses of adjacent rings are seen in tangential longitudinal sections. Nine large peripheral dense fibres are as- sociated with the transition between the distal centriole and the axoneme (Fig. 3D) and continue posteriorly, though much narrower, along the axoneme into the midpiece (Figs 3E, F, 4D). One is attached externally to each triplet or doublet. Within the midpiece, at an undetermined level, all but two of the peripheral fibres become greatly FIG. 2 Nangura spinosa. A.Whole spermatozoon (Nomarski contrast light microscopy). B. Longitudinal section (L.S) through the apical end of the acrosome showing the perforatorium. C. L.S through the basal region of the acrosome showing the nuclear point and paracrystalline matrix of the subacrosomal cone. D-I. A series of transverse sections (T.S) through the acrosome. Note that anteriorly, in D and E, the acrosome is compressed in transverse sections, while further posteriorly, in F-H, it is unilaterally ridged, and at its posterior limit, in 1, it is circular. J. T.S through the nucleus. K. L.S through the basal region of the itcrosomc showing the epinuclear electron lucent region. B-K to the same scale, as indicated. Abbreviations a = acrosome; av = acrosome vesicle; et = epinuclear electron lucent region; mp = midpiece; n = nucleus; p = perforatorium; pm = plasma membrane; pp = principal piece; sc = subacrosomal cone. pm 174 MEMOIRS OF THE QUEENSLAND MUSEUM SPERMATOZOA OF NANGURA SPINOSA 175 reduced in size. Only peripheral fibres adjacent to doublets 3 and 8 remain conspicuous, as a double structure nearer the fibrous sheath than it is to its doublet (Figs 3E-G). An unspecified peripheral fibre is seen in longitudinal section at the centriolar end of the axoneme to be cross striated (Fig. 4D). The only well developed, though small, peripheral fibres at the level of the annulus are the double fibres at doublets 3 and 8 (dense columns in Fig. 1 ). At the beginning of the principal piece all nine dense fibres are already vestigial or absent (Figs 3G, H). They are absent from the remainder of the principal piece (Fig. 31). The annulus (Figs 3B, G, N) is a small dense ring with an irregular oval cross section. It is closely applied to the inner surface of the plasma membrane. Principal Piece The principal piece, the longest part of the spermatozoon, consists of the continuation, be- hind the midpiece, of the axoneme with its sur- rounding fibrous sheath and plasma membrane. It begins, immediately behind the annulus, with a short region in which a wide zone of cytoplasm intervenes between the fibrous sheath and the plasma membrane (Figs 3B, H). The cytoplasm is finely granular, giving the region some resemblance to a glycogen piece but the presence of glycogen has not been determined. Posterior to this the plasma membrane is closely ap- proximated to the fibrous sheath (Figs 31, M). Endpiece The axoneme projects behind the fibrous sheath as an endpiece of undetermined length (Figs 3J, K). DISCUSSION Extension of the fibrous sheath into the mid- piece in the sperm of N. spinosa is an autapomor- phy of the Squamata, unknown in the sperm of other reptiles (Healy & Jamieson, 1992; Jamieson & Healy, 1992) or other amniotes. Newton & Trauth (1992) are incorrect in suggesting that in Tropidurus sperm (Da Cruz-Landim & Da Cruz- Hofling, 1977), the fibrous sheath does not extend into the midpiece. Nangura is placed by Covacevich et al. (1993) in the Sphenomorphus group of Greer (1979) which also includes Ctenotus. Sperm of Ctenotus robustus and C. taeniolatus have been examined by Jamieson & Scheltinga (in preparation). In the Eugongylus group of Greer (1979), Oliver & Jamieson (unpublished data) have examined the sperm of Cryptoblepharus virgatus, in the Eugongylus subgroup, and Lampropholis deli- cata and Carlia pectoralis, in the Lampropholis subgroup. The sequence of ring structures in the midpiece of Nangura make it more similar to Ctenotus than it is to any other examined reptile. Species of the Eugongylus group differ in having scattered intermitochondrial bodies (considered homologous with the ring structures) in the form of small dense irregular spheres, tortuous rods or large plates (Oliver & Jamieson, unpublished data). The sperm of Chalcides ocellatus ex- amined by Furieri (1970) resemble Nangura in having four regularly placed intermitochondrial rings but, unlike Nangura, each ring consists of a circlet or small juxtaposed spherules rather than a continuous ring. Varanus gouldiiflavirufus also has 4 dense intermitochondrial structures in lon- gitudinal sequence. However, each ‘ring’ is made of many loosely aggregated large granules that do not form a continuous ring (Oliver & Jamieson, unpublished data). In all amniote classes the acrosome plesiomor- phically forms an elongate, narrow cone sym- metrically located on the lip of the nucleus which it overlaps and constricts. The acrosome vesicle, with this form, encloses a similarly shaped sub- acrosomal cone the margins of which arc poorly defined in Nangura. The pointed form of the acrosome, presence of the subacrosomal cone. FIG. 3 Nangura spinosa. A. Longitudinal section (L.S) through the neck region showing, as a squaniate autapomorphy, that the fibrous sheath penetrates the midpiece almost to the junction between the axoneme and the distal centriole. B. L.S through the midpiece-principal piece junction showing the annulus. C. Transverse section (T.S) through the nuclear fossa. D. T.S through the distal centriole-axonemal transition showing the 9 peripheral fibres associated with the triplets or doublets. E. T.S through a ring structure. F. T.S through the midpiece showing 12 mitochondria surrounding the axoneme. G. Oblique T.S through the annulus. H and 1. T.S through the principal piece. J and K. T.S through the endpiece. L. L.S through the neck region showing the dense cone occupying the conical nuclear fossa. M. L.S through the principal piece. N.L.S through the niidpiece showing the four ring structures and annulus separated by four sets of columnar rnitcx:hondna. A-M to the same scale, as indicated. Abbreviations an = annulus; d = dense cone; dc = distal centnole; fs - fibrous sheath; las - laminar structure; mi = mitochondria; n = nucleus; nt = nuclear fossa; pc - proximal centnole, pf- penpheral fibre; pm = plasma membrane; rs = ring structure. 176 MEMOIRS OF THE QUEENSLAND MUSEUM and constriction of the nuclear tip are seen in the Chelonia, Crocodilia, Sphenodon, Squamata (as in Nangura), non-passerine birds (the sub- acrosomal cone is lost in ratiles), and mono- tremes. These states are also seen in the Lissam- phibia, including the primitive frog Ascaphus (Jamieson et al., 1993) and presumably charac- terized the common ancestor of Amphibia and Amniota. A paracrystalline substructure of the sub- acrosomal cone, as in Nangura, has been recog- nized in other squamates (Butler & Gabri, 1984; Carcupino et al., 1989; Furieri, 1970), including Ctenotus (Jamieson & Scheltinga, in prepara- tion). It is probably a synapomorphy, and autapomorphy, of the Squamata. All classes of amnioles possess one or more endonuclear canals, containing one or more per- foratoria, which penetrate the anterior end of the nucleus to varying depths. This condition is also seen in basal lissamphibians: urodeles (Picheral, 1967) and primitive anurans (see Jamieson et al., 1993; Sandoz, 1970), There are also endonuclear canals in Lathneria, and Neoceratodus (see Jam- ieson, 1991), therefore the presumed common ancestor of Amphibia and amniotes probably pos- sessed one or more endonuclear canals. Absence of endonuclear canals is a synapomorphy of the squamates and a homoplasy of these with non- passerines and monotremes (Jamieson & Healy, i 1992). However, squamates, including Aawgwra, retain a perforatorium anterior to the nucleus. The nucleus is an elongate cylinder narrowly constricted within the base of the acrosome in some members of all amniote classes (Jamieson t& Healy, 1 992). This form, also seen in Ascaphus and urodeles (Jamieson et al., 1993), is clearly plesiomorphic for amniotes. It is widespread in reptiles, including Nangura. Representation of the basal nuclear fossa is variable in amniotes. It is poorly developed in the sperm of the caiman, and is small and compact in iurilcs, Sphenodon (tuatara), rooster, guinea fowl, and squamates excepting the skinks. In skinks it ¥\G A Nangura spinosa. A. Longitudinal section (L.S) through the full length of the acrosome. B-D. L.S through the neck region showing, B, the oblique angle of the proximal centriole relative to the distal centriole, C, the laminar structure, and D, a cross striated peripheral fibre. E. T.S through the proximal centriole showing that the microtubules of the proximal centriole make contact with the ring struc- ture. F. T.S through a ring structure showing the outer surface scalloped by small mitochondria. Abbrevia- tions av = acrosome vesicle; d = dense cone; dc = distal centriole; fs = fibrous sheath; las = laminar structure; mi = mitochondria; n = nucleus; nf = nuclear fossa; p = perforatorium; pc = proximal centriole; pf = peripheral fibre; rs = ring structure; sc = subacrosomal cone. SPERMATOZOA OFNANGURA SPINOSA 177 is narrowly funnel-shaped or conical, as in Nan- gura. In the ratites it has a triple profile. The small and compact form may be plesiomorphic for amniotes (Jamieson & Healy, 1992). A dense ring, the annulus, at the posterior end of the midpiece is a feature of many metazoan sperm and is clearly plesiomorphic for amniotes. It has been demonstrated, inter alia, in turtles, crocodile, tinamou, rhea, rooster, guineafowf tuatara and monotremes. Squamates were con- sidered exceptional in absence or at most negli- gible development of an annulus (Jamieson & Healy, 1992). However, an annulus has been demonstrated for Lacerta vivipara by Courtens & Depeiges (1985), Cnemidophorus sexUneatus by Newton & Trauth (1992) and for Nangura, and may be more widely present in squamates than previously suspected. The number of mitochondria seen in transverse section of the midpiece, where possible near its anterior end, is very variable in amniotes. The number in chelonians, here considered the most basal amniotes, is six. A trend towards reduction to four in birds and monotremes has been ob- served. There are eight or nine in the tuatara, the caiman and the skink Chalcides ocellatus (Jamieson & Healy, 1992). In the teiid Cnemi- dophorus sexUneatus there are 8 to 10 (Newton & Trauth, 1992). In snakes, the number shows apomorphic increase to as many as 14. The num- ber, approximately 12 in Nangura, is also apomorphically high. The predominantly linear arrangement of cris- tae in the mitochondria of Nangura sperm is a usual feature of amniotes, including most reptiles. Only turtles, caiman and the tuatara, are Imown to be exceptional. The mitochondrial cristae in these three taxa are concentric and usually sur- round a large central dense body. In all other anuiiotes, excepting the Wooly opossum which also has concentric cristae, the cristae have a ‘conventional’ appearance, being linear or curved but never concentric, and do not surround a dense body . The concentric arrangement around a dense body is here considered to be an apomorphy acquired early in amniote evolution, as evidenced by its occurrence in Chelonia but later lost (Jamieson & Healy, 1992). The intermitochondrial rings, ‘ring structures’ in Nangura, which are limited to squamates are regarded as derivations of the dense bodies of basic amniotes (Jamieson & Healy, 1992). Car- cupino et al. (1989), independently concluded that the rings in Chalcides ocellatus tiligugu, were mitochondrial derivatives. Origin of inter- mitochondrial material from mitochondria has been confirmed by demonstration ontogenetical- ly in the sperm of some squamates by Oliver & Jamieson (in preparation). The distal centriole, forming the basal body of the axoneme, is plesiomorphically short in ver- tebrates, including the Lissamphibia, as in most Metazoa, and in Nangura. In contrast, the distal centriole extends the entire length of the long midpiece in the tuatara, turtles, crocodiles, and ratites, an apparent basal synapomorphy of am- niotes. The shorter, though still elongate distal centriole in the rooster and the somewhat shorter centriole in guinea fowl, the short centriole in squamates, and the vestigial, possibly absent, centriole in monotremes possibly represent secondary reduction in length of the centriole (Healy & Jamieson, 1992). A cross striated dense body lateral to the proximal centriole, represented by the laminated structure in Nangura, appears to be a basal synapomorphy of amniotes but its homology across the various groups requires confirmation. It is seen in tuatara, and the caiman where homol- ogy with the striated columns of eutherian sperm has been suggested (Healy & Jamieson, 1992), It has not been reported for sperm of birds or monotremes and the squamates but in view of its presence in Nangura further examination of squamates is needed. The annulated, dense fibrous sheath seen in Nangura must have developed in the earliest am- niotes as it is present in all amniote classes (though it is absent in some birds). Occasional anastomoses of adjacent rings are seen in tangen- tial longitudinal sections and similar ‘branching’ in the snake Lampropeltis getulus, led Austin (1965) to propose that the annuli are linked together along one or both sides of the tail. A fibrous sheath is absent from amphibian sperm. The isolated peripheral fibres 3 and 8 in am- niotes may well be homologous with columns at this point in other sarcopterygians, the coelacanth, Latimeria, and Dipnoi. However, such modifications at doublets 3 and 8, which are approximately in the plane of the two central singlets, could be independent acquisitions as they presumably are in chondrichthyan sperm (references in Jamieson, 1991). In contrast with reptiles, in eutherian mammals the outer coarse fibres at 3 and 8 are the smallest and they ter- minate first, their place being occupied throughout most of the length of the principal piece by inward prolongations of the dorsal and 178 MEMOIRS OF THE QUEENSLAND MUSEUM ventral portions of the fibrous sheath (Hamilton & Fawcett, 1968). Nine longitudinal dense fibres peripheral to the 9 axonemal doublets, as in Nangura^ are a fun- damental feature of amniote sperm, being found in all classes. They are an autapomorphy and simultaneous symplesiomorphy of the amnioles. The peripheral dense fibres are small in most amniotes investigated: turtles, the caiman, tuatara, squamates and monotremes. The peripheral fibres are described as ‘tiny’ for the rhea. They are present in suboscine and the more apomorphic oscine passerines, being larger in the latter. They are large and diverse in shape in marsupials above the didelphids, and in eutherian mammals. There thus appear to be trends to en- largement of the peripheral fibres in passerines and non-monotreme mammals, with diversifica- tion in the latter. The peripheral fibres are usually situated in the midpiece with some extension into the principal piece as in turtles, the caiman, non-passerines, tuatara, and monotremes. In eutherians and mar- supials they extend far into the principal piece. However, in Nangura, as in other squamates, the only well developed, though small, peripheral fibres at the level of the annulus are the double fibres at doublets 3 and 8 and by the beginning of the principal piece all nine dense fibres are al- ready vestigial or absent. In non-passerine birds the fibres may be restricted either to the midpiece or to the principal piece or occur in both or, as in doves, are absent (Jamieson & Healy, 1992; Asa & Phillips, 1987; Jamieson, unpublished data). In turtles, the tuatara and the skinks Nangura and Chalcides see Furieri (1970), the nine peripheral dense fibres are partly displaced from the radii of the doublets into the gaps between adjacent doublets. Their exact radial position is unknown in the crocodile and in most squamates. In contrast, the peripheral dense fibres lie in the same radius as the doublets in the rhea, at least some non-passerines (turkey) and monotremes (Jamieson & Healy, 1992). ACKNOWLEDGEMENTS We are grateful to Mr P.J. Couper for giving us the opportunity to obtain testicular material from a specimen of A. spinosa. Mrs Lina Daddow and Mr Tom Gorringe are thanked for excellent tech- nical assistance. LITERATURE CITED AL-HAJJ, H., JANAKAT, S. & MAHMOUD, F. 1987. 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SPRANDO, R. L. & RUSSELL. L. D. 1988. Sper- miogenesis in the red-ear turtle (Pseudemys scripta) and the domestic fowl (Callus domes- ticus): A study of cytoplasmic events including cell volume changes and cytoplasmic elimina- tion. Journal of Morphology 198: 95-118. TUZET. O. & BOURGAT, R. 1973. Recherches ultrastructurales sur la spermiogenese de Chaniaeleo senegalensisDaudin, 1802. Bulletin Biologiqiiede la France et de la Belgique 107(3): 195-212. YASUZUMI. G. & YASUDA. M. 1968. Sper- matogenesis in animals as revealed by electron microscopy. XVIII. Fine structure of developing spermatids of the Japanese freshwater turtle w'ith potassium permanganate. Zeitschrift ftir Zellforschung 85; 18-33. YASUZUMI, G.. YASUDA, M. & SHIRAI, T. 1971. Spermatogenesis in animals as revealed by electron microscopy. XXV. Fine structure of spermatids and nutritive cells during sper- miogenesis in the Japanese freshwater turtle. MonitoreZoologico Italiano 5: 1 17-132. 180 MEMOIRS OF THE QUEENSLAND MUSEUM TWO NEW SPECIES OF STRIPED BLINDSNAKES GJ INGRAM AND J.A. COVACEVICH Ingram, G.J. & Covacevich, J.A. 1993 12 24: Two new species of striped blindsnakes. Memoirs of the Queensland Museum 34(1):181-184. Brisbane. ISSN 0079-8835. Ramphotyphlops Silvia sp.nov. occurs in sand in rainforest mostly within the Great Sandy Region, SEQ. It is striped to black-and-white with 20 midbody scale rows and differs from R. minimus (midbody scale rows 16) and R. chamodracaena sp.nov (18). R. hroomi also has 20 midbody scale rows, but has a nasal cleft that distinguishes it from R. silvia sp.nov. R. chamodracaena sp.nov. is a striped blindsnake found in the w'est and far north of Cape York Peninsula. Midbody scale count separates it from other striped species. □ Reptilia, Serpentes, Typhlopidae, Ramphotyphlops, new species, blindsnake, Queensland, Australia. Glen Ingram & Jeanette Covacevich, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; 24 September, Blindsnakes are usually dull, virtually pattern- less snakes. However, some species are strongly marked, sometimes striped, e.g. Ramphotyphlops minimus (Kinghom, 1929) and R. broomi (Boul- enger, 1898). To these we add R. silvia sp.nov. and R. chamodracaena sp.nov. Striped blind- snakes are found mainly in woodlands of northern Australia. The latter new species is from northern woodlands. R. silvia sp.nov., however, inhabits rainforests on sand, southeastern Queensland. Measurements (mm) and scale counts here follow Storr (1981). Specimens prefixed by ‘J’ and ‘R’ are housed in the Queensland and Australian Museums respectively. Regions for Queensland follow Ingram & Raven (1991). Ramphotyphlops silvia sp.nov. (Figs 1,2) Material Examined Holotype: J27387 Seary’s Scrub, Cooloola NP (25°58’S, I53W’E), SEQ. CollectedbyJ. Covacevich & P. Filewood, 3-6 February, 1976. Paratypes: J31579 Fraser Is., NP HQ; J3 1576-7 Fraser Is. NP, HQ on ‘A’ rd; J35872 Tuan SF, Firetower 6; J27386 Seary’s Scrub, Cooloola NP; J43785 Cooloola NP; J23620 Cooloola, on Freshwater rd, 5km from junction; J46128 Pomona, 25km N, on rd to Rainbow Beach; J8521 Nambour. All localities in SEQ. Diagnosis A small (maximum total length 175), striped or black-and-white Ramphotyphlops with 20 mid- body scale rows. The nasal cleft extends up from the nostril, to about the level of the eye and nearly divides the nasal (Fig. 1). 1993. Description Total length: 72-175 (N 10, mean 143.6). Length of tail (% of total length): 2-5 (N 1 0, mean 3.6). Rostral (from above) elliptic, a little longer than wide, about half as wide as head and extend- ing back to just in front of, or just between, the level of the eyes. Nasals narrowly separated be- hind rostral. Frontal smaller than prefrontal. Snout rounded in profile. Nostrils inferior, closer to rostral than preocular. Nasal cleft proceeding from second labial (close to the juncture of the 1 st and 2nd labial) and extending vertically from nostril to about the level of eye, nearly dividing the nasal. Midbody scale rows 20 (N 10). Ventrals 272-320 (N 9, mean 293.9). Subcaudals 14-21 (N 9, mean 17.6). Dorsal and lateral surfaces with 1 1 thick, purplish brown stripes along body on a creamy yellow background. Mostly however, the stripes merge and and the dorsal and lateral sur- faces appear evenly purplish black. Ventral sur- face cream to creamy yellow — one specimen has two black bands across the throat. The ventral colour contrasts strongly with the lateral colour (Fig- 2). Distribution Coastal rainforest on Quaternary sands, from Fraser Is. to Noosa NP, SEQ (S. Wilson, pers. comm.), except for an old specimen from Nam- bour, SEQ. Etymology There are two allusions. Rhea Silvia was the mother of Romulus and Remus, legendary found- ers of Rome. Hannah Sylvia Ingram is the mother of one of us. 182 MEMOIRS OF THE QUEENSLAND MUSEUM Remarks R. Silvia appears to be endemic to the Quater- nary sands of coastal SE Queensland and most of its distribution falls within the Great Sandy Region. For discussions of these faunas see Kik- kawa et al. (1979) and Dwyer et al. (1979). FIG. 1. Ramphotyphlops silvia sp.nov. (paratype J23620). Above: dorsal view of head. Centre; lateral view of head. Below: ventral view of head. Ramphotyphlops chamodracaena sp.nov. (Fig. 3) 1992 Ramphotyphlops sp. Cameron & Cogger, p.60, pl.25. Material Examined Holotype: J40233 N Camp ‘Beagle*, ca.40km N Aurukun (B'^OS’S. I4B59*E), Cape York Peninsula. Collected by G. Ingram & P. Webber, 18 March, 1982. Paratypes: J31963 Weipa, Andoom Mine; J41550 Weipa; R91631 Rocky Pt, Weipa; R93164 Weipa; J28082 Lockhart R. Settlement; J39673 N Camp ‘Beagle’, 15km from camp on rd to Watson R.; J51980 Inkerman Stn. All localities are on Cape York Peninsula. Diagnosis A small (maximum total length 210), striped Ramphotyphlops with 18 midbody scale rows. The nasal cleft extends diagonally up from nostril to terminate about halfway between nostril and rostral (Fig. 3). Description Total length: 1 14-210 (N 8, mean 164.3). Length of tail (% of total length): 1-3 (N 8, mean 1.8), Rostral (from above) elliptic, a little longer than wide, about half as wide as head and extending back to just in front of, or just between, the level of the eyes. Nasals narrowly separated behind rostral. Frontal smaller than prefrontal. Snout rounded in profile. Nostrils inferior, closer to rostral than preocular. Nasal cleft proceeding from second labial and extending diagonally up from nostril to ter- minate shortly about half-way between nostril and rostral. Midbcxly scale rows 18 (N 8). Ventrals 464-523 (N 7, mean 486.1). Subcaudals 14-16 (N 8, mean 15.1). Dorsal, lateral and ventral surfaces usually totally striped (18 stripes along body, but one specimen has 16). Stripes are ctok to light brown on a yellowish cream to off-white back- ground. Tail black and, in some specimens, the head is black. For a colour photograph, see Cameron & Cogger (1992, pl.25). Distribution Western Cape York Peninsula from Weipa south to frikerman Station. Also recorded from Lockhart R. Settlement on the east coast. Inhabits woodland, but Cameron & Cogger (1992) also note that its occurrence on lawns after watering. Etymology ‘Chamodracaena’ (earth snake) was one of the 12*/2 names of the female demon Gello (Stoneman, 1991). TWO BLIND SNAKES 183 FIG. 2. Ramphotyphlops silvia sp.nov. Pile Valley, Fraser Is., August, 1993 (S. Wilson). Remarks Most of the publications on Australian blindsnakes have followed Waite (1918). His paper was an important beginning, but his ‘species’ are sometimes not easily recognised. It is also difficult to recheck his work, because no registration numbers are given for his specimens and there are no data on where they are housed. The identity ofR. broomi (Boulenger, 1898) is a case in point. Waite examined five specimens and referred them to Boulenger’ s taxon. These came from Queensland, Victoria and Western Australia. However, Storr (1981) made no men- tion of broomi, or any other striped species, in his revision of the Western Australian blindsnakes. As well, we cannot be confident that we have examined any specimens of Waite’s broomi. Thus we are unsure of his concept of the taxon. Even so, in northeastern Queensland, there is a small blindsnake known only from the area be- tween the western edge of the Atherton Tableland and Cooktown. It matches Boulenger’s descrip- tion (‘20 scales round the body. Pale buff above, with 11 brown streaks following the series of scales, white beneath’) and is found near the type locality of R. broomi, Muldiva, NEQ. However, we cannot be completely sure of the identity until the holotype of Typhlops broomi in the Natural History Museum, London, can be examined. For this paper, we have assumed that this taxon is R. broomi. If it is, considering the localities Waite gave, we are confident that none of his specimens were broomi, because that species appears to be restricted to the dry belt along the western edge of the Wet Tropics. The confusion with the identity of R. broomi is evident in Wilson & Knowles (1988). Their broomi' (pl.827) is not R. broomi as we understand it. However, their photograph of Ramphotyphlops sp. (pl.847) appears to be true R. broomi. This is J47254 from Cooktown — S. Wil- son, pers. comm. R. chamodracaena is most similar to R. broomi and R. minimus. However, besides the smaller number of midbody scale rows (18 vs 20), R. chamodracaena is usually striped all round the body while R. broomi is not striped ventrally (number of stripes 16-18 vs 11-15). Cameron & Cogger (1992) have discussed the similarities and differences between R. chamodracaena (as Ram- photyphlops sp.) and R. minimus, a species with 16 midbody scale rows. 184 MEMOIRS OF THE QUEENSLAND MUSEUM Other Material Examined R. hroorni: J46849 Cooktown, James Cook Museum Building; J47254 Cooktown; J20315 Mt Molloy; J2953-4, 47503 Atherton Tblds, Stannary Hills, nr Herberton; J53305 Mt Rosey, 15 mile Ck camp; R 128220 Mandalee, Innot Hot FIG. 3. Ramphotyphlops chamodracaena sp.nov. (paralype J39673). Above: dorsal view of head. &ntre: lateral view of head. Below: ventral view of head. Springs. AH localities in Cape York Peninsula and NEQ. R, minimus: R9692 Groote Eylandt, NT (holotype); R9693, R61025-6 (paratypes) same data as holotype. ACKNOWLEDGEMENTS We thank Patrick Couper (drawings), Steve Wilson (photograph) and Ross Sadlier (Australian Museum specimens). LITERATURE CITED BOULENGER, G.A. 1898. Description of two new snakes from Queensland. Annals and Magazine of Natural History (2)7; 414. CAMERON, E.E. & COGGER, H.G. 1992. The her- petof auna of the Weipa region. Cape Y ork Peni n- sula. Technical Reports of the Australian Museum 7: 1-200. DWYER, P.D., KIKKAWA, J. & INGRAM, G.J. 1979. Habitat relations of vertebrates in subtropi- cal heathlands of coastal southeastern Queensland. Pp.280-299. In Specht, R.L. (ed.), ‘Ecosystems of the world. 9A. Heathland and related shrublands. Descriptive studies*. (El- sevier Scientific Publishing Co.; Amsterdam). 497 pp. KIKKAWA, J., INGRAM, G.J. & DWYER. P.D. 1979. The vertebrate fauna of Australian heath- lands - an evolutionary perspective. Pp. 23 1 -279. In Specht, R.L. (ed.), ‘Ecosystems of the world. 9A. Heathland and related shrublands. Descrip- tive studies*. (Elsevier Scientific Publishing Co.: Amsterdam). 497 pp. INGRAM, G.J. & RAVEN. R.J.eds 1991. ‘An atlas of Queensland’s frogs, reptiles, birds and mam- mals.’ (Queensland Museum: Brisbane). 391pp. KINGHORN, J.R. 1929. Two new snakes from Australia. Records of the Australian Museum 17: 190-192. STONEMAN, R. 1991, ‘Greek mythology: An en- cyclopedia of myth and legend.’ (Aquarian Press: London). 192pp. STORR, G.M. 1981. The genus Ramphotyphlops (Ser- pentes: Typhlopidae) in Western Australia. Records of the Western Australian Museum 9: 235-271. WAITE, E.R. 1918. Review of the Australian blind snakes (Family Typhlopidae). Records of the South Australian Museum 1: 1-34, pi. 1. WILSON, S.K. & KNOWLES, D.G. 1988. ‘Australia’s reptiles: a photographic reference to the terrestrial reptiles of Australia.’ (William Collins: Sydney). 447pp. NEW REPTILE RECORDS FROM RAINFORESTS OF SOUTH AND MIDEASTERN QUEENSLAND J. A. COVACEVICH, P.J. COUPER AND GJ. INGRAM Covacevich, J.A., Couper, PJ. & Ingrain, GJ. 1993 12 24: New reptile records from rainforests of south and mideastem Queensland. Memoirs of the Queensland Museum 34(1):185-187. Brisbane. ISSN 0079-8835. Recent surveys of 13 previously unsurveyed small rainforests of south and mideastem Queensland have refined knowledge of the distribution of many reptile species. Nangura spinosa gen. etsp.nov., Lygisaurus zuma sp. nov., Phyllurus isis, P. nepthysand P. ossa spp. nov. have been described largely as a result of this survey. Q Survey, reptiles, rainforest, Queensland, Australia. J.A. Covacevich, PJ. Couper & GJ. Ingram, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; 17 July, 1993. Following a decade of fairly intense research, moist, near-coastal rainforest reptiles are now probably better known than those of other habitats in Queensland. Many new species have been described from Queensland’s notophyll and mesophyll vine forests, and revisionary work on several rainforest genera has now been completed or is nearly so. Nowhere has the product of well- organized rainforest research been more evident than in the Wet Tropics, recently (1988) placed on the World Heritage Register. Since 1979, 13 of the 20 reptile species endemic to those rain- forests have been described (Covacevich, in press). From recent descriptions and revisions, from reference collections (notably of the Queensland and Australian Museums), and from the earlier literature, i I has been possible to review distributions of many rainforest species, at least to the extent of defining their north-south limits and commenting on aspects of their zoogeog- raphy and conservation (Covacevich & Mc- Donald, 1991; Covacevich, in press). Much is known as a result of collecting under- taken either randomly or in surveys in the large rainforests. Major surveys (e.g. Anonymous, 1976; Broadbent & Clark, 1976; Queensland Museum, 1977) have focussed on larger tracts of rainforest. In these studies mideastern Queensland rainforests have not generally received the same attention as those further north or south, so their reptile species are not so well known. In south and mideastem Queensland, there are many small, isolated rainforests about which nothing herpetological is known. Semievergreen vine thickets, particularly, are poorly known and have been reduced by clearing for agriculture (Forster, et al., 1991). In 1991, our proposal to the National Rainforest Conservation Programme to make small collec- tions of reptiles in previously unsurveyed rain- forests of south and mideastern Queensland was approved for funding. The object was to add to knowledge of the distribution of rainforest rep- tiles, particularly the less well-known species. Our study sites were selected from areas from which there were no museum records of reptiles. No attempt was made to ensure the sites were comparable in terms of size, soil type, aspect, season or survey time, because our interest in the reptile species was purely discovery. Replication was not considered necessary. Four-ten ‘man’ days were spent collecting reptile specimens at most sites, using conventional methods of rolling logs and rocks, and peeling bark and ‘working' sun patches. Spotlighting was undertaken at all sites. At some very small rainforests (e.g. War- riwillah, a remnant of the now-cleared Isis Scrub) only one day was spent. In the species list (Table 1) the number of ‘man’ days spent at each is indicated in brackets. These records are not, and are not intended to be complete. Rather, they are a small contribution to knowledge of the distribu- tions of some of the species of reptiles. For most species, voucher specimens have been lodged in the Queensland Museum. A species name not accompanied by a ‘J' number is based on sight record only, always by al least one of us. Collection sites are arranged north to south. NEW INFORMATION This work to improve the accuracy of distribu- tion data for reptiles in south and mideastem Queensland has made returns beyond our expec- 186 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1 . Some small, previously-unsurveyed rainforests of south and mideastem Queensland: reptile records from October, 1991 to April, 1993. MT OSSA 20°56’ 148°49’, MEQ (6) notophyll vine forest: Phyllurus ossa (J53443-7, J53507), Oedura ocellata (J53463), Carlia rhomboidalis (J53456-61), Eulamprus amplus (J53465, J53531-2), E. tenuis (J53466), Lampropholis adonis {J53452-4, J53462), Saproscincusspectabilis (J53508), Physignathus lesueurii (J53533), Morelia spilota, Boiga irregularis, Dendrelaphis punctulata (J53442), Demansia psammophis (J53440-1). COFFEE CREEK. MT JUKES 2r00’ J48°57’, MEQ (10) notophyll vine forest: Phyllurus isis (J53485-6, J53518), Anomalopus verreauxi (J53600), Carlia rhomboidalis (J53562-4, J53576-81), Eulamprus amplus (J53523), £ quoyii, Lampropholis adonis (J53565, J53570-5, 35360^-10), Saproscincus spectabilis (J53566-7, J53582'4, J5361 1-2), Physignathus lesueurii (J53595), Morelia spilota (J53593), Dendrelaphis punctulata (J53515), Cacophis squamulosus (J53516), Demansia psammophis (J53487, J53585). ST HELEN’S GAP. VIA MT CHARLTON 21 '’00 1 48M3’, MEQ (3) notophyll vine forest: Phyllurus ossa (J53426-8). Carlia rhomboidalis (J53429-3 1), Eulamprus ampltts (J5347 1-2), E. tenuis (J53473), Lampropholis adonis (J53432-4), Saproscincus spectabilis (J53435-9), Varanus varius, Boiga irregularis (J53425). MT CHARLTON (foothills) 2U0r 148°44’, MEQ (9) notophyll vine forest: Oedura cf. tryoni (J53412), Phyllurus ossa (J53389-90, J53414). Carlia rhomboidalis (J53402-7, J5341 5), Eulamprus amplus (J53399-400, J53828-30). E. tenuis (J53401, J53416) Lampropholis adonis (J5341 0- 1 1 , J5341 7-2 D.Sapro^cmcu.y spectabilis (J53408-9, J53422-4) Varanus varius, Morelia spilota, Demansia psammophis. MT BLACKWCXJD NATIONAL PARK 21 '’02’ 1 48'’56\ MEQ (9) notophyll vine forest: Oedura ocellata (J53479), Phyllurus isis (J53480, J5351 1-2, J53591, J53602-3). Carlia rhomboidalis (J53474-5, J53514, J535 59-60), Eulamprus amplus (J53485, J535 1 3), Lampropholis adonis (J53476-8, J53483, J53558), Saproscincus spectabilis (J53481), Morelia spilota 053606), Boiga irregularis (J535 10, J53522), Cacophis harriettae (J53526), Rhinoplocephalus nigrescens (J53607), ROCKY DAM CREEK via Crcditon Site I 2n8’ 148'’32’, MEQ (4) notophyll vine forest: Oedura ocellata (J53333), PhyUurivi nepthys (J53 330-2), Eulamprus tenuis (J53334-5), Lampropholis adonis (53336-50) Varanus varius. Site 2 21°19’, 148°34’ (3) notophyll vine forest: Phyllurus nepthys (J53359-62), Carlia rhomlwidalis (J53352-4, J53363-6), Eulamprus amplus (J53358, 353827). E. tenuis (J53356-7). Larnpropluflis adonis (J53355. J53367-78), Saproscincus spectabilis, Physig- nathus lesueurii, Varanus varius, Dendrelaphis punctulata (J5335 1), EAST FUNNEL CREEK 21^36’ 149'’I2’, MEQ (8) notophyll vine forest: Carlia rhomboidalis (J53489-9I, J53496, J53501-2), Eulamprus quoyii (J53534), Lampropholis adonis (J53492-3, J53498, J53500, J53503-6, J53613), Saproscincus spectabilis (J53497, J5360i, J53614). Physignathus lesueurii (J53494), Morelia spilota (J533 19), Boiga irregularis (J53509), Demansia psatnmophis (J53524). WEST HILL CREEK headwaters. Connors Range 22'’5r 149°18', MEQ (2) notophyll vine forest: Carlia rhomboidalis (J53537-8), Lampropitolis adonis (J53539-50), Boiga irregularis (J53530). KA KA MUNDFSCRUBB Y CREEK 24‘’50’ I47°25’, SCQ (6) semievergreen vine thicket; Gehyra catenata, Lialis burtonis, Anomalopus brevicoUis (J56037-41), A. leuckartii (J56042-3, J56061), Carlia pectoralis (J56035, J56046), Cryptoblepharus virgatus (J56034), Cterwtus taeniolatus (J56044, J56062) Eulamprus martini (J56065-6. J56083-4), Lerista fragilis (J56063), L. punctatovittata (J56090), Gemmaiophora nobbi (J56060), Morelia maculosa (J56071), Dendrelaphis punctulata, Rhinoplocephalus nigrescens (J56052). KA KA MUNDI, PACKS ADDLE SPRING 24'*50’ I47°27’, SCQ (1) semievergreen vine thicket: Gehyra catenata i)56057), G. dubia (J56t)58-9), Anomalopus leuckartii (J56055), Lerista fragilis (J56056). WARRIWILLAH, via Cordalba 25°06’ I52‘’13’,SEQ(1) notophyll vine forest (riverine): Calyptotis scutirostrum (J57242-3), Lygisaurusfoliorum (J57244), Ophioscincus Cfxdoolensis (J57240- 1) Physignathus lesueurii, Boiga irregularis, Tropidonophis mairii. NANGUR STATE FOREST 26 ”07’ 151 “58’. SEQ (9) semievergreen vine thicket: Anomalopus verreauxii (J55996), Calyptotis scutirostrum (J56001-8) Carlia vivax (J56009), Cienotus arcanus (J56024), Lampropholis amicula (J57245), Nangura spinosa (J56029, J56031, J57246, J57247), Morethia taeniopleura (J56028), Saiphos equalis (J55997-600, J56027, J56030), Gemmatophora nobbi (J 56025), Vit/ranw^ varius, Dendrelaphis punctulata, Rhinoplocephalus nigrescens (J56026). MISTAKE MOUNTAINS 27°55’ I52“20’, SEQ (2) notophyll vine forest: Cautula zia (J31320), Coeranoscincus reticulatus (J54646-50), Lampropholis delicata (J54436), Saproscincus challengeri (J32276-7), Physignathus lesueurii (J54438), Morelia spilota (J54437), Hemiaspis signata (J54439), Tropidechis carinatus (J54434). tations. New species have been collected; new data published on Coeranoscincus reticulatus and Morelia spilota; and the range of Eulamprus amplus has been extended. Most significant was the discovery of Nangupa spinosa, a distinct spiny lygosomid, by Cameron James and Mark Fletcher in Nangur State Forest, a remnant patch of semievergreen vine thicket (Covacevich et al., 1993). Most of the type specimens of Lygisaurus zuma Couper, 1993 were collected at Boulder Creek, near Mt Charlton, MEQ, in open forest during this survey; Phyllurus isis and P. ossa Couper et al., 1993 were described from specimens discovered in this survey, and P. nep- thys Couper et al., 1993 has been described fol- lowing re-examination of existing material and of new specimens found on this survey. Coeranoscincus reticulatus is a rare skink (Mc- Donald et al., 1991) occurring narrowly in southeastern Queensland and northeastern New i NEW REPTILE RECORDS FROM QUEENSLAND 187 South Wales rainforest As a result of work on the Mistake Mountains, Couper et at (1992) have described the nest, eggs and young of this species. Morelia spilota is a well-known predator of mammals and birds. In October, 1991 at East Funnel Creek, 21°37\ 149°12’, near Sarina MEQ, Covacevich & Couper ( 1 992) found a large male M. spilota dead, following ingestion of a large specimen of the Cane Toad, Bufo marinas. M. spilota does not prey upon B. marinas under normal conditions. This discovery confirms pre- vious anecdotal accounts. Eulampras amplas was described (Covacevich & McDonald, 1980) from two populations - Eun- gella and Conway National Parks, via Mackay and Proserpine, MEQ. It has been rated as ‘rare’ (McDonald et at, 1991). It is now known to occur in several intervening areas of rainforest, al- though its conservation status remains unchanged by the new records. ACKNOWLEDGEMENTS We thank officers of the Department of En- vironment and Heritage (Alan Beckman, Shamus Conway, Alan Horsup, Cameron James, Paul Lawless-Pine, Keith McDonald, Barry Nolan, Steve Pearson, Gary Rees and Peter Thompson); property owners who gave us access to their forests (Leo Baretta, Don Brownsey, Merv Colby, Mr A. Gardner, Mike Hull, Garth Ilett, Ian Jenkins, and Paul and Gloria McFazden); and Terry Beutel, Michael Cunningham, Barry Hein- rich, Glen Hodges, Duncan Limpus, Tim Low, Martin Schultz and Paul Webber. LITERATURE CITED ANONYMOUS, 1976. ‘Fauna of eastern Australian rainforest: preliminary report on sites surveyed by the Queensland Museum in mid-eastern and northeastern Queensland’. (Queensland Museum: Brisbane). 78pp. BROADBENT, J. & CLARK, S. 1976. ‘A faunal survey of east Australian rainforests: Interim Report’. (Australian Museum: Sydney). 132pp. COUPER, P.J. 1993. A new species of Lygisaurus de Vis (Reptilia: Scincidae) from mideastern Queensland. Memoirs of the Queensland Museum 33: 162-166. COUPER, P.J., WHITTIER, J., MASON, R.T & IN- GRAM, G.J. 1992. A nesting record for Coeranoscincus reticulatus. Memoirs of the Queensland Museum 32: 60. COVACEVICH, J. (in press). Rainforest reptiles of Australia’s World Heritage Wet Tropics: zoogeography and conservation issues. Journal of the International Herpetological Symposium. COVACEVICH, J.A. & COUPER, P.J. 1992. The Carpet Python, Morelia spilota (LacepMe), another unsuccessful predator of the Cane Toad Bufo marinas (Linnaeus), in Australia. Contribu- tions in Herpetology, Greater Cincinnati Her- petological Society: 57-9. COVACEVICH, J. & MCDONALD, K.R. 1980. Two new species of skinks from mid-eastern Queensland rainforest. Memoirs of the Queensland Museum 20: 95-101. COVACEVICH, J. & MCDONALD, K.R. 1991 . Frogs and reptiles of tropical and subtropical eastern Australian rainforests: distribution patterns and conservation. Pp.281-309. In Werren, G. & Ker- shaw P. (eds), ‘The rainforest legacy, Australian National Rainforests Study, Volume 2 - flora and fauna of the rainforests’. Australian Heritage Commission Special Australian Heritage Publi- cation Series Number 7(2). (Australian Govern- ment Publishing Service: Canberra). 414pp. COVACEVICH, J.A., COUPER, P.J. & JAMES, C. 1993. A new skink, Nangura spinosa gen. et sp. nov., from a dry rainforest of southeastern Queensland. Memoirs of the Queensland Museum 34(1); 159-167. FORSTER, P.L, BOSTOCK, P.D., BIRD, L.H. & BEAN, A.R. 1991. ‘Vineforest plant atlas for south-east Queensland. An assessment of conser- vation status’. (Queensland Government, Queensland Herbarium: Brisbane). 478pp. MCDONALD, K.R., COVACEVICH, J.A., IN- GRAM, G.J. & COUPER, P.J. 1991. The status of frogs and reptiles. Pp,338-345. In Ingram, G.J. & Raven, R.J. (eds), ‘An atlas of Queensland’s frogs, reptiles, birds and mammals’. (Board of Trustees, Queensland Museum: Brisbane). 39Ipp. QUEENSLAND MUSEUM, 1977. 'Fauna of eastern Australian rainforests II. Preliminary report on sites surveyed by the Queensland Museum in southeastern and far northeastern Queensland, with additional results from sites surveyed pre- viously in northeastern Queensland’. (Queensland Museum: Brisbane). 102pp. 188 MEMOIRS OF THE QUEENSLAND MUSEUM POSSUM ASSEMBLAGES IN RAINFOREST OF THE CARBINE UPLANDS, NEQ, WITH SPECIAL REFER- ENCE TO HEMBELWEVS LEMVROIDES. Memoirs of the Queensland Museum 34( / ):}88. 1993:- Four ringtail pos- sum (Pseudocheiridae) species are confined to the upland rainforests of northeastern Queensland’s Wet Tropics. These are the Lemuroid Possum (Hemibelideus lemuroides). Green Possum {Pseudochirops archeri), Daintree Ringtail {Pseudo- cheirus cinereus) and Herbert River Ringtail {P. herberten- sis). At least five other, wider-ranging species (the Long-tailed Pygmy Possum, Cercartetus caudatus\ Striped Possum, Dae- tylopsila trivirgala; Common Ringtail, P. peregrinus\ Sugar Glider, Petaurus breviceps and Coppery Brushtail, Trichosurus vulpecuta johnstonii) are also regularly en- countered. These high altitude rainforests are not uniformly distributed. Mountainous terrain divides the area into discrete blocks, montane subregions (sensu Winter el al., 1984; Winter, 1991). A major altitudinal discontinuity exists be- tween the Atherton Tableland and the Carbine Uplands. Geographic separation is evident for upland obligate fauna unable to negotiate the relatively low altitude Black Mountain rainforest corridor connecting them. With isolation, allopatric speciation has occurred. This is reflected in the separation of P. cinereus, of the Carbine Uplands (Murray et al., 1989), from P. herbenensis of the Atherton Tableland The former was known as the ashen-coloured race or subspecies of P.her- bertensis . Vehicular spotlight runs were conducted along the Ml Lewis Forestry Rd (approximately 1 1 0km NW Cairns, NEQ) during the summers of 1986/87 and 1987/88. Altitudes be- tween 500-1200 m were surveyed during 13 spotlight traver- ses, averaging approximately 6km and 2 hours duration. Most of this survey was undertaken at altitudes exceeding 900m ( 19.75 hours) with only 6.75 hours of survey carried out below this. Only when possums could be positively identified were they accepted and recorded. Altitudes were recorded by al- timeter. This method was employed to detect changes in possum density and community make-up with changes in altitude along the traverse. In addition, it was anticipated that a sufficient number of H. lemuroides sightings could be ob- tained to estimate the relative proportion of leucisl individuals within the population, and provide evidence to support or refute llie suggested lower altitudinal limit of 900m (Winter et al., 1984) of the Carbine population. Results of the survey are summarised in Table 1. While only four possum species were recorded during the period of the survey (Table 1), an additional two species are known to occur In rainforests of the Carbine Uplands. ’ITie species C. caudatus and P. breviceps are recorded only oc- casionally. The numerically dominant species within all al- titudinal bands between 501-1100 was P. cinereus. Above 1100m, more //. lemuroides were recorded. A general trend of increasing possum abundances with increasing altitude w as partially due to this sudden appearance of //. lemuroides at the higher elevations. The lowest elevation al which H. lemuroides was recorded was 1070m. If such a lower altitudinal limit is consistent for the entire northern population isolate of this species, it may indicate a restricted relictual distribution and lengthy .separation from the population on the Atherton Tableland. The incidence of leucism among this population was high, 31.56% of individuals recorded. TABLE 1. Possums on Mt Lewis Rd, Carbine Uplands. Altitude (m) P.cinereus H. lemuroides (dark/pale) P.archeri D.trivirgata Total per spoiliglit hour 1101-1200 65 98(66/32) 1 16.40 1001-1100 34 3(3/0) 2 1 121 901-1000 18 1 1 4.71 801-900 13 2 5.00 701-800 4 1 2.00 601-700 2 2.67 501-6002 4.00 Totals 138 101 (69/32) 7 2 9.36 The rainforest possum assemblage of the Carbine Uplands differs in several ways from that of the Atherton Tableland. Replacement of F. herbertensisv^ithP. d/ierewj, and absence of T. vulpecula jolmstonii from the Carbine Uplands are sig- nificant differences. Leucism in the Atherton population of H. lemuroides is also a rarity. One biologist noted only three pale individuals in thousands of records made over a period of 6 years of upland possum research on the Atherton Tableland (N. Goudberg, pers. comm.). Further, the sug- gested lower altitudinal limit of 480m (Winter et al., 1984) is substantially lower for Atherton Tableland H. lemuroides than both the suggested Carbine Uplands lower limit of 900m (Winter et al.,1984) and 1070m, which was the lowest record made in this survey. (Many hours of traverse prior and sub- sequent to those reported here have failed to reveal any H. lemuroides at lower elevations). These observations sug- gest investigation of the taxonomic status of animals from the northernmost population would be worthwhile. Acknowledgments Supported by the Rector of the University College, Univer- sity of New South Wales, Australian Defence Forces Academy, Prof. G. V. H. Wilson. Special thanks are due also to Mr John Meade. Literature Cited Murray,!. D., McKay, G. M., Winter, J.W.&lngelby,S. 1989. Cytogenehes of the Herbert River ringtail possum, Pseu- docheirus herbertensis (Diprotodonta: Pseudo- cheiridae): evidence for two species. Genoma, 32: 1119-1123. Winter, J. W. 1991. Mammals. P.43. In Nix, H. A. & Switzer, M. A. (eds), ’Rainforest animals:allas of vertebrates endemic to Australia’s WetTropics’, Kowari 1. (CRES- AN U/ANPWS : Canberra). Winter, J. W., Bell. F. C, Pahl, L. 1. & Atherton, R. G. 1984. ) The specific habitaisof selected northeastern Australian |l rainforest mammals’. Report to WWF (Australia). 135 pp. M. P. Trenerry, 4/14 Casella St., Earlville, Queensland, 4870, Australia.; G. L. Werren, cf- Douglas Shire Council, PO Box 357, Mossman, , Queensland, 4873, Australia. DISTRIBUTION AND CONSERVATION OF FROGS AND REPTILES OF QUEENSLAND RAINFORESTS J.A. COVACEVICH AND K.R. MCDONALD Covacevich, J.A & McDonald, K.R. 1993 12 24: Distribution and conservation of frogs and reptiles of Queensland rainforests. Memoirs of the Queensland Museum 34 ( 1 ): 1 89-199. Brisbane. ISSN 0079-8835. Forty-six species of frogs and 95 species of reptiles are now known from the wet and dry rainforests of Queensland. Close to 70% of frog species and 63% of reptile species are well protected; nearly 20% of frog and reptile species are still either poorly or not protected in reserves. Conservation programmes for Queensland’s rainforest frog and reptile species must address the following problems: unexplained declines in frog populations; survey of moist rainforest on Moa Is.; protection of moist rainforest at Somerset and the Mcllwraith Ra., FNEQ, and of dry rainforests; and the paucity of ecological data on most species. Many new species of frogs and reptiles have been described in the last decade, and the herpetofauna of Queensland’s rainforests is now well-known taxonomically. It is important that research on this group not consume an excessive proportion of the research resources, to the detriment of the futures of other taxa and habitats. □ Australia, rainforest, frogs, reptiles, declining frogs, conservation. Wet Tropics, distribution. J.A. Covacevich, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia; K.R. McDonald, Queensland Department of Environment and Heritage, PO Box 834, Atherton 4883, Australia; 5 September, 1992. A review of patterns of occurrence and conser- vation of frog and reptile species restricted to tropical and subtropical moist rainforests of east- ern Australia has been published recently (Covacevich & McDonald, 1991). That review was based essentially on museum specimens and references available in the early 1980s, and sub- mitted for publication in 1984. The period 1984- 1990 was a very active one for research on Queensland’s rainforest frogs and reptiles and, while some up-dating was possible while the paper was in press, much that was new could not be included. This work is both narrower and broader than the previous study. Here, we confine ourselves to Queensland rainforests, because most of what is I new regarding rainforest frog and reptile species has been discovered in Queensland. However, in addition to obligate rainforest species we include all species recorded in rainforests. Further, we have adopted a broader definition of rainforest, including both ‘wet rainforests’ (vine forests and fern forests of all types) and ‘dry rainforests’ (= deciduous and semi-deciduous vine thickets). As well as incorporating data on facultative rainforest species (along with the obligate rain- forest species), this review includes data of Greer et al., 1983 {Lerista vittata)\ Greer, 1985 {Sphen- omorphus cracens); Wells & Wellington, 1985 (many spp., including Cacophis churchilli and Saltuarius swaini)\ Greer & Cogger, 1985 {Anomalopus and Ophioscincus spp.); Zweifel, 1985 (microhylid frogs); Davies et al., 1986 (Litoria .xanthomera); Corben & Ingram, 1987 (Mixophyes fleayi); Ingram & Covacevich, 1988 (Lygisaurus spp.); Ingram & Covacevich, 1989 (Car//fl spp.); Ingram, 1991 (Lampropholis spp.); Sprackland, 1991 {Varanus teriae); McDonald, 1992 (conservation of northern rainforest frogs); Greer, 1992 {Eulamprus tenuis group); Couper et al., 1993 (Saltuarius spp. and Phyllurus spp.); Covacevich el al., 1993 (Nangura spinosa); and Sadlier et al., 1993 (Saproscincus challengeri species complex); Richards et al., 1993 (Litoria eucnemiSy L. genimaculata). Since the 1980s the results of frog/reptile surveys in previously un- known or poorly known rainforests have been published (e.g. Horsup et al., 1993; Covacevich et al, 1993). These are incorporated, along with additional records compiled incidentally by one of us (KMcD). New data has been combined with that known previously into a list of frog and reptile species recorded from Queensland’s rainforests sensu lato (Table 1). For each species are provided: ranges in both latitude and altitude; notes on habitat; conservation status; and level of habitat protection. 190 MEMOIRS OF THE QUEENSLAND MUSEUM DISTRIBUTION In assessing the species for inclusion in Table 1, three categories of reptiles and frogs occurred only occasionally in rainforest and are not con- sidered further: open vegetation species which were using patches of rainforest or gallery forest as dry season refugia (e.g. Bufo marinas, Um- nodynastes ornatus, Litoria bicolor, L. gracilen- ta, L. nigrofrenata, L. rothii, Sphenophryne gracilipes, Glaphyromorphus pumilus, Lygis- aurus macfarlani, L. sesbrauna); species occur- ring on rainforest margins or utilising natural openings such as rock faces and scree slopes (e.g. Heteronotia binoei, Oedura rhombifer, Dendrelaphis calligastra, D. punctulatus, Trop- idonophis mairii, Demansia torquata, Morelia maculosa); species entering rainforest which has been disturbed by roads, logging, mining or real estate (e.g. Bufo marinas, Litoria nasuta, Lim- nodynastes peronii, Litoria nigrofrenata, Lialis burtonis, Heteronotia binoei, Oedura rhombifer, Carlia longipes, C. rostralis, C. pectoralis, Tropidonophis rnairii, Morelia maculosa, Demansia psammophis, D. torquata). Conservation status was determined using the definitions of Thomas and McDonald, 1989. Level of habitat protection is defined: well protected (recorded from several conservation reserves, or all or most of the population in reser- ves); moderately protected (found in more than two reserves); poorly protected (only recorded from one or two reserves); unprotected (not on any reserves). Table 1. Frog and reptile species recorded from Queensland’s wet and dry rainforests, based largely on records of the Queensland Museum, the literature and observations by KMcD. Frogs Myobatrachidae Adelotus brevis (Gunther 1863): Eungella NP, MEQ; Byfield, MEQ -Grove Ck, NSW; 21° OO’S, 148° 35’E - 21° ir, 148'^35’;22°50M50°4r-33°50M48°23’;sea level - 1000+m; subtropical vine forest, also heaths, moist forests and moist sandstone gorges of sub-coastal upland areas; secure; well protected. Assa darlingtoni (Loveridge 1933): Conondale Ra., SEQ - Gibraltar Ra., NENSW; 26° 37’S, 152° 35’E - 29° 36’, 152° 13’ ; usually above 1000m, infrequently to 300m; subtropical vine forest; rare; moderately protected. Kyarranus kundagungan Ingram & Corbcn, 1975: Mistake Mtns - Mt Superbus, SEQ; 27° 55’S, 152° 20’E - 28^^ 12’, 152° 29*; 300 - 1000m; subtropical vine forest; rare; well protected. Kyaranus ioveridgei (Parker, 1940): McPherson Ra., SEQ - Ml Warning, NENSW; 28° 15’S. 153° 15’E - 28° 24’, 153° 16’; above 750m; subtropical vine forest; rare; well protected. Lechriodusfletcheri (Boulenger, 1890): Cunningham’s Gap, SEQ - near Gosford, NSW; 27°58’S, 152o 25’E - 33° 25’, 151° 20 ; above 750m; subtropical vine forest; secure; well protected. Mixophyes fasciolatus Gunther, 1864: Eungella NP, MEQ; 20° 50’S, 148° 28’E - 21° 1 1’, 148° 32’; Kroom.bit Tops, MEQ - Gosford, NSW; 24° 24’,151°00’ - 33° 25’, 151° 17’; sea level - 1 100m; subtropical vine forest, adjacent moist open forests; secure; well protected. Mixophyes fleayi Corben and Ingram, 1987: Conondale Ra., SEQ - Tooloom Scrub, NSW; 26° 30’S, 152° 37’E - 28° 40’, 152° 30’; 620m+; subtropical vine forest; insufficiently known; poorly protected. Mixophyes iteratus Stiaughan, 1968: Conondale Ra., SEQ -Narooma, NSW; 26° 37’S, I52°35’E-36° 13’, 150° 09’; 100 - 1000m; subtropical vine forest; insufficiently known; poorly protected. Mixophyes schevilU Loveridge, 1933: Big Tbind - Mt Halifax, NEQ; 15° 42’S, 145° 16’E - 19° 07’, 146° 22’; sea level - 1500in; monsoon vine forest; secure; well protected. Rheobatrachus silus Liem. 1973; Blackall and Conondale Ra., SEQ, Booloumba Ck - Kilcoy Ck, SEQ ; 26° 37’S, 152° 35’ - 26° 47’, 152° 38’; 400 - 800m; subtropical vine forest; endangered, ? extinct; poorly protected. Rheobatrachus vitellinus Mahony, Tyler & Davies, 1984: Eungella NP, Clarke Ra., MEQ; 20° 50’S, 148° 33’E - 21° 07’, 148° 33’; 400 - 1000m; subtropical vine forest; endangered, ? extinct; well protected. Taudactylus acutirostris (Andersson, 1916): Big Tbind - Mt Graham, NEQ; 15° 42’S, 145° 16’E - 18° 24’, 145° 52’; 300 - 1500m; monsoon vine fore.st; endangered; well protected. Taudactylus diurnus Straughan & Lee, 1%6: Blackall and Conondale Ras, Mt Glorious, SEQ; 26° 37’S, 152° 35 ’E - 27° 23’, 152° 47’; 400 - 800m; subtropical vine forest; endangered, ? extinct; poorly protected. Taudactylus eungellensis Liem & Hosmer, 1973: Cathu SF - Eungella NP, Clarke Ra., MEC); 20° 50’S, 148° 33'E-2I°07’, I48°33’;200- 1000m; subtropical vine forest; endangered; well protected. Taudactylus liemi Ingram, 1980: Cathu SF, Clarke Ra. - Crediton SF,MEQ;20°51’S, I48°34’E-2r 11’, 148° 35’; 250 - 1000m; subtropical vine forest; rare; well protected. Taudactylus pleione Czechura, 1986: Kroombil Tops SF, SEQ; 24° 24’S, 150° 59’E; 900m; subtropical vine forest; insufficiently known; poorly protected. Taudactylus rheophilus Liem & Hosmer, 1973 : four allopatric populations on Thornton Pk, Carbine Tbind, Lamb DISTRroUTION OF FROGS AND REPTILES 191 Ra.» Bellenden Ker Ra., NEQ; 16° lO’S, 145° 22’E - 17° 16\ 145° 22’; 9(X)m - 15(X)m; monsoon vine forest; endangered; well protected. Ranidae Rana daemeli (Steindachner, 1868): Cape York tip - Rollingstone, NEQ; extralimital in PNG; 10° 42’S. 142® 32’E - 19° 03\ 146° 23*; sea level - 6^m, mostly < 100m; monsoon vine forest, usually from open forests, entering rainforest along roads & clearings, Cape York populations from riparian rainforest; secure; well protected. Hylidae Utoria caerulea (White, 1 790): throughout Q, extralimital in WA, NT, SA, NSW; sea level - 800m. Predominant- ly associated with open vegetation, entering disturbed rainforest & dry vine thickets; secure; well protected. Utoria chloris (Boulenger, 1893): Conway Ra. NP - Eungella NP, MEQ; Bulburin SF, MEQ - Gosford, NSW; 20°24\ 148° 35’-21°08\ 148°30’;24° 15’, 151° 24’ - 33°26\ 151°20’;sea level- 1000m; subtropical vine forest; secure; well protected. Utoria eucnemis (Lonnberg, 1900): Ducie R. - Wenlock R.; Iron Ra. - Mcllwraith Ra., FNEQ; 12° 07’S, 142° 2rE - 12° 16*, 141° 59*; 12° 46*, 143° 16* - 13° 50*. 143° 17’; sea level - 20m; monsoon vine forest; secure; poorly protected. Utoria genimaculata (Horst, 1883): Big Tblnd - Paluma, NEQ; 15° 42’S, 145° 16*E - 19° 01*. 146° 12’; sea level - 1300m; monsoon vine forest; secure; well protected. Utoria infrafrenata (Gunther, 1867): Cape York tip, FNECJ - Mutarnee, NEQ; Extralimital in PNG; 10° 42’, 142° 32’ - 19° 00*, 146° 20’; sea level - 580m; monsoon vine forest, usually from open forests enters riparian rainforest & rainforest edges; secure; well protected. Utoria lesueurii (Dumeril & Bibron, 1841): Cooktown, NEQ - Victoria; 15° 25*S, 145° 05’E-V; sea level - 1200m; monsoon, dry, subtropical vine forests & moist open forests; secure; well protected. Zirona Tyler & Davies 1978: Leo Ck- Station Ck, FNEQ; 13® 42’S, 143° 18*E- 13°57’, 143° 19’; 400 - 820m; monsoon vine forest; rare; not protected. Utoria lorica Davies & McDonald, 1979: Alexandra Ck - Hilda Ck, Thornton Pk NEQ; 16° 07*, 145° 20’ - 16° 10’, 145° 23’; 6^ - l(X)0m; monsoon vine forests; vulnerable; well protected. Utoria nannotis (Andersson, 1916); Big Tb!nd-Mt. Halifax. NEQ; 15° 42’S. 145° 16’E- 19° ll’S 146°27’E; 80-1300m; monsoon vine forest; vulnerable; well protected. Utoria nyakaleftsis (Liem, 1974): McDowall Ra. - Douglas Ck, Kirrama Ra., NEQ; 16° 07 S, 148° 20 E - 18 13*, 145° 48’; 380 - 1020m; monsoon vine forest; endangered; well protected. Utoria pearsoniana (Copland, 1961): Kroombit Tops, SEQ - Lismore, NSW; 24° 24’S, 151® 01 E - 28° 50 , 153° or ; 200 - 1000 m; subtropical vine forest, sometimes along perennial, densely vegetated streams in open forest adjacent to rainforest; insufficiently known; well protected. Utoria revelata Ingram, Corben & Hosmer, 1982: three allopatric populations; Atherton Tblnd, NEQ; Clark Ra., MEQ; Mt Tamborine, SEQ - Smith’s Lake. NSW; 17° 23’S. 145° 42’E - 17° 36*. 145° 29’; 21° 01*. 148° 36’ -21°07’, I48°31’;27° 55*. 153° I0’-32°23’, 152° 29’; 400 - 900m; monsoon and subtropical vine forests, open forests, and pastures; secure; moderately protected. Utoria rheocola (Liem, 1974): Amos Bay - Broadwater Ck, NEQ; 15° 41 S, 145° 19 E - 18 23 , 145 57 , sea level - 1180m; monsoon vine forest; vulnerable; well protected. ^ Utoria xatithortxera Davies, McDonald & Adams, 1986: Big Tl>lnd -Bluewater Ra., NEQ; 15 42 S, 145 17 E - 19° 07’, 146° 23’; 20 - 1300m; monsoon vine forest; secure; well protected. Nyctimystes dayi (Gunther, 1897): Big Tblnd - Paluma, NEQ; 15° 42 S, 145° 16 E - 19 01 , 146 13 , sea - 1200 m; monsoon vine forest, along densely vegetated perennial streams adjacent to rainforest, vulnerable, well protected. ^T^^l^Tombiens Zweifel, 1986: Shipton’s Flat - Mt Lewis, NEQ; 15° 47’S, 145° 14’E - 16° 35’, 145° 16’; 200 - 1300m; monsoon vine forest; rare; well protected. „ iz:o^c» Cop/iixa/usconcmnu^Tyler, 1979:MtFinnigan-MtLew'is,NEQ; 15 49 S, 145 17E-16 35,145 16;560 - 1300m; monsoon vine forest; rare; well protected. -, 0 /^ Cophixalus crepim,is Zweifel, 1 985: Mcllwraith Ra., FNEQ; 13° 44 ’S, 143° 20’E, 380 - 54(^, rare; unprotecte^^^ Ccp/iaa/i«eA:/gttt«Zweifel&Parker,1969:BigTblnd-MtFinnigan,NEQ;15 42 8,145 16 E- 15 49,145 16’; 180 - 1 120 m; monsoon vine forest; rare; moderately prot^ted. Cophixalus hosmeri Zweifel, 1985: Mt Spurgeon - Mt Lewis, NEQ; 16 27 S, 145 12 E - 16 35 , 145 16 , 960 - 1 370m; monsoon vine forest; rare; moderately protected. Mnn i ? 1 dfV F 1 8° 74’ rn/ncems Zweifel, 1985: Crystal Cascades -Dalrymple Gap, NEQ, 16 58 8,145 40 E- 18 24, 146° 05’ ; 40 - 800m; monsoon vine forest; secure; well protect^. Cophixalus mcdonaldi Zweifel, 1985: Mt Elliot, Bowling Green Bay NP, NEQ, 19 30 8 , 146 58 E, 900+ m, monsoon vine forest; rare; well protected. r. i- NTr:r». no ia«:o/iQ’P 17 ° Cophixalus neglectus Zweifel, 1962: Mt Bellenden Ker and Mt Bartle Frere, NEQ, 17 16 8 , 145 49 E - 17 24’, 145° 51’; 900 - 1600m; monsoon vine forest; rare; well protected. 192 MEMOIRS OF THE QUEENSLAND MUSEUM Cophixalwi ornatus (Fry, 1912): Mt Spurgeon - Bluewater Ra., NEQ; 16° 27’S, 145° 23’E - 19° 12’, 146° 22’; sea level - 1520m; monsoon vine forest; secure; well protected. Cophixalus peninsularis Zweifel, 1985: Leo Ck, Mcllwraith Ra., FNEQ; 13° 44’S, 143° 20’E; 520-540m; monsoon vine forest; insufficiently known; unprotected. SphenophymefryiZyffciicU 1962: Big Tblnd- Lamb Ra. NEQ; 15°42’S, 145° 16’E- 17° 08’, 145° 36’; sea level - 1300m; monsoon vine forest; secure; well protected. Sphenophyme pluvialis Zweifel, 1965: Gap Ck, Cedar Bay NP - Mt Fox SF, NEQ; 15° 49’S, 145° 19’E - 18° 35*, 145° 49*; sea level - 1300m; monsoon vine forest; secure; well protected. Sphenophyme rohusta{¥ry, 1912): Lamb Ra. -Bluewater Ra., NEQ; 17®06’S, 145° 36*E- 19° 12’, 146° 23’; 360 - 1520m; monsoon vine forest; secure; well protected. Reptu.es Gekkonidae Carphodactylus laevis 1897): Big Tblnd- Douglas Ck,Kirrama, NEQ; 15°42’S, 146° 16’E- 18^12’, 145° 47* ; 300 - 1400m; monsoon vine forest; secure; well protected. Cyrtodactylus louisiadensis (de Vis, 1892): two disjunct populations: Mcllwraith Ra., FNEQ; Cape Flattery, NEQ-Chillagoe, NEQ; extralimilal in PNG; 12° 30’S, 143° 16’E; 15° 01’, 145° 13’ - 17° 16’, 144° 37’; sea level - 600m; monsoon vine forest, also occurs in moist open forests on Cape York; secure; moderately protected. Phyllurus caudiannulatus Covacevich, 1975: Bulburin SF; 24° 3 1* 1 5 1 ° 30’ ; 200 - 600m; subtropical vine forest; rare; unprotected. Phyllurus isis Couper, Covacevich & Moritz, 1993: Mt Blackwood -Ml Jukes, MEQ; 21 ° 02’S, 148° 56’E - 21° 02’, 148° 57’; 250 - 500m; monsoon vine forest; rare, well protected. F/iy//MrM.y Couper, Covacevich & Moritz, 1993:Clarke Ra., MEQ; 21° Ol’S, 148°36’E-21° 19’, 148° 34’; 250 - 1000+ m; subtropical vine forest; rare; well protected. Phyllurus ossa Couper, Covacevich & Moritz, 1993: Mt Dryander, Conway NP - Mt Charlton, MEQ; 20° 15’S, 148°33’E - 21° 01*, 148° 44'; 200 - 400m; monsoon vine forest; rare; well protected. Saltuarius cornutus (Ogilby, 1892): Big Tblnd - Paluma, NEQ; 15° 42’S, 146° 16’E - 19° 01’, 146° 14’; 100 - 1 100m, mostly above 600m; monsoon vine forest; secure; well protected. Saltuarius occultus Couper, Covacevich & Moritz, 1993: Peach Ck, McIlwTaith Ra., FNEQ; 13° 45’, 143° 19’ - 13° 46’, 143° 19’; 500 -800m; monsoon vine forest; insufficiently known; unprotected. Saltuarius salebrosus (Covacevich, 1975): Blackdown Tblnd, MEQ -Cracow, SEQ; 23° 46’S, 149° 06’E - 25° 18’, 150° 18*; sea level - IO(X)m; dry vine thickets, often associated with sandstone uplands & gorges, open forest; secure; moderately protected. Sa/ri^nW.y>va/A»(Wells44’E- 21°06’, 148° 29’; 150- 1100m; subtropical & monsoon vine forest; rare; well protected. Eulamprus brachysoma (Lonnberg & Andersson, 1915): Coen, NEQ - Lever’s Plateau, nr Lamington NP, SEQ; 13° 56’S, 143° 12’E - 28° 20’, 152° 52’; sea level - 1 100m; monsoon & subtropical vine forest; secure; well protected. Eulamprus frerei Greer, 1992: Mt Bartle Frere, NEQ; 17° 24’S, 145° 48’E; 1600m; monsoon vine forest; insufficiently known; well protected. Eulamprus luteilateralis (Covacevich & McDonald, 1980): Ml William & Mt Dalrymple area, Eungella NP, MEQ; 21° 08’S, 148° 30’E; above 900m; subtropical vine forests; rare; well protected. Eulamprus martini Wells & Wellington, 1985: Homevale, MEQ - Coffs Harbour, NSW; 21 23 S. 148 32 E - 30° 20’, 153° 07’; sea level - 1000m; dry vine thickets, usually from open forests; secure; well protected. ^ Eulamprus murrayi (Boulenger, 1887): Conondale Ra., SEQ - Barrington Tops, NSW; 26 37 S, 152 35 E - 32° 02’, 151° 24’; 300 - 1000+m; subtropical vine forest; rare; moderately protected. Eulamprus quoyii (Dumeril & Bibron, 1839): Dainlree, NEQ - Budawang, NSW; 16° 15’S, 145° 19’E - 30° 18’, 153° 08’; sea level -1000m; subtropical & monsoon vine forest; secure; well protected. Eulamprus sokosoma Greer, 1992: Taravale, NEQ - Injune, MEQ; 19° 07’S, 146° 04’E - 25° 51 , 148 34 ; 20 - 80(hn; dry vine thickets; rare; poorly protected. £«/.im/,n/./em.u(Gray,183I):Holboumcls.,MEQ-Bega,NSW:19°43’S,148°22’E-36°40’,149°55’;sea level - 750m; dry vine thickets, subtropical vine forest; secure; well protect^. ngrmM,y(De Vis 1888 ): Shipton’s Flat -Smoko Ck, NEQ; 15 48 S, 145 14E- 18 09,145 37; sea level - 1600 m; monsoon vine forest; secure; well protected. ,^ocn>c ^AAoz^^^:: Glaphyromorphus cracens {Grcex, 1985): Ml Mulligan - Wyandotte Ck, NEQ; 16 50 S, 144 51 E - 18 35 , 144° 33’; 400- lOOOm; dry vine thickets, predominantly from open fo^este; seciue; ^rly protec^^^^ G/np/i>Tomo;7?/ii4v/i4sdcWi5(Greer, 1979):BigTblnd-BluewaterRa.,NEQ;15 42 S, 145 16 E - 19 15, 146° 29’; 40 - 1 160m; monsoon vine forest; secure; well prot^ted. Glaphyromorphus mjobergi (Ldnnberg & Andersson, 1915): Mount Carbine TbInd - Vine Ck, NEQ, 16 34 S, 145° 15*E - 17° 36’ 145° 29’* above 650m; monsoon vine forest; rare; well protected. • (Peters, 1871); Kaban. NEQ - Mt Walsh NP SEQ; 17° 3 1 ’S, 145° 25’E - 25° 35’ , 152° 03’; sea levd - lOOOm; dry vine thicket, predominantly from open forests; secure, well protected. 194 MEMOIRS OF THE QUEENSLAND MUSEUM Gnypetoscincus queenslandiae (DeWis, 1890): Big Tblnd-Mt Thom, NEQ; 15°42’S, I45°16’E- 18° 16’, 145° 48’; 40 - 1200ni; monsoon vine forest; secure; well protected. Lamprophol is adonis lngr 2 im, 1991: Conway NP,MEQ-Maleny,SEQ; 20° 15’S, 148° 32’E-26°47’, 152° 49’; 20 - 9^m; monsoon & subtropical vine forest; secure; moderately protected. Lampropholis amicula Ingram & Rawlinson, 1981: Kroombit Tops, SEQ - Karuah SF, NSW; 24° 22’S, 150° 59’E - 32° 36’, 151° 56’; 20 - 900m; subtropical vine forest; secure; moderately protected. Lampropholis coggerilngxd^m, 1991:Big Tblnd - Mt Elliot, NEQ; 15°42’S, 145° 16’E -19° 29’, 146° 58’, sea level - 1500m; monsoon vine forest; secure; well protected. Lampropholis colossus Ingram, 1991: Bunya Mtns, SEQ; 26° 53’S, 151° 37’E; 1000 -1200m; subtropical vine forest; rare; well protected. Lampropholis couperi In^am, 1991: Byfield, MEQ - Mt Glorious, SEQ; 22° 50’S, 150° 4rE - IT 25’, 152° 50’; 40 - 700m; subtropical vine forest; insufficiently known; poorly protected. Lampropholis mirabilis Ingram & Rawlinson, 1981: Magnetic Is. - Mt Elliot, NEQ; 19° OS’S, 146° 50’E - 19° 29’, 146° 58’; sea level - 1234m; monsoon vine forest & dry vine thicket; rare; well protected. Lampropholis robertsi Ingram, 1991 : Thornton Pk - Mt Bartle Frerc, NEQ; 16° lO’S, 145° 23’E - 17° 23’ , 145° 48’; 600 - 1520m; monsoon vine forest; secure; well protected. Lerista colliveri Couper & Ingram, 1992: Battery Stn - Hughenden, NEQ; 19° 26’S, 145° 52’E - 20° 51’, 144° 12’; 300 - 400m; dry vine thickets, predominantly from open forests; secure; poorly protected. Lerista vittata Greer, McDonald & Lawrie, 1983: Mt Cooper Stn, NEQ; 20° 31’S, 146° 55’E; 250 - 300m; dry vine thickets; vulnerable; unprotected. Lygisaurus laevis Ingram & Covacevich, 1988: Amos Bay - Bramston Beach, NEQ; 15° 4rS, 145° I9’E - 17° 21’, 146° 01 ’; sea level - 950m; monsoon vine forest; secure; moderately protected. Lygisaurus tanneri Ingram & Covacevich, 1988; Mclvor R., Starcke Stn - Endeavour R., NEQ; 15° 03’S, 145° 08’E - 15° 27’, 145° 08’; sea level - lOOm; monsoon vine forest; insufficiently known; unprotected. Nangura spinosa Covacevich, Couper & James, 1993: Nangur SF, SEQ; 26° 07’S, 151° 58’E; 300m approx.; semievergreen vine thicket; rare; moderately protected. Ophioscincus cooloolensis Greer & Cogger, 1985; Aldoga, nr Gladstone - Cooloola, SEQ; 23° 48’S, 151°04’E - 25° 58’, 153° 07’; sea level - 900m; subtropical vine forest; rare; moderately protected. Ophioscincus ophioscincus (Boulenger, 1887); Bulburin SF - Brisbane, SEQ; 24° 3rS, 151° 33’E - IT 28’, 153°0'; sea level - 500m; subtropical vine forest, heathlands; rare; moderately protected. Ophioscincus truncatus(\>a^T^. 1876): Coolum, SEQ-Way Way SF, NSW;26°33’S, 153°05’E-30°47’, 152° 56’; sea level - 800m; subtropical vine forest; adjacent open forests; rare; moderately protected. Saiphos equalis (Gray. 1825): Kroombit Tops, SEQ - Wollongong, NSW; 24° 20’S, 15 1 ° Ol’E - 34° 35’, 150° 52’; sea level - llOOm; subtropical vine forest, coastal heathland and moist sclerophyll forests; secure; moderately protected. Saproscincus challengeri (Boulenger. 1887): McPherson Ra., area, SEQ & NENSW; 28° 03S, 152° 24’E - 28° 24’. 153° 04’; sea level -500m; rainforest, moist coastal lowland forest; secure; well protected. Saproscincus czechurai Ingram & Rawlinson, 1981: Big Tblnd - Walter Hill Ra., NEQ; 15° 42’- 145° 16’ - 17*^ 47’, 145° 49’; 500 - 1000m; monsoon vine forest; secure; well protected. Saproscincus galli Wells & Wellington, 1985: Mt Tamborine, SEQ - Sydney area, MENSW; 27° 58’S, 153° 1 1’E, 33° 53’, 151° 13’; sea level - lOOOm; rainforest, low closed remnant forest; secure; well protected. Saproscincus rosei Wells & Wellington, 1985: Gympie, SEQ - Barrington Tops, MENSW; 26° 11’, 152° 40’E - 32° 10’, 151° 26’; sea level - lOOOm; rainforest, moist riverine forest; secure; well protected. Saproscincus spectabilis (de Vis, 1888): Mt Webb - Mt Elliot, NEQ and Mt Dryander- East Funnel Ck, MEQ; 15° 04’S, 145° 08’E - 19° 29’, 146° 58’; 20° 15’, 148° 38’ - 21° 36’, 149° 12’; sea level - lOOOm; monsoon & subtropical vine forest; secure; well protected. Saproscincus tetradactyla Kluge & Greer, 1979: Mossman Gorge, Daintree NP - Paluma, NEQ; 16° 28’S, 145° 16’E- 19° 00’, 146° 12’; 50- lOOOm; monsoon vine forest; secure; well protected. Agamidae Hypsilurus boydii (Macleay, 1884): Big Tblnd - Paluma. NEQ; 15° 42’S. 145° 17’E - 19° 01’. 146° 14’; 20 - 1160m; monsoon vine forests; secure; well protected. Hypsilurus spinipes (Dumeril & Dumeril, 1851): Gallangowan, SEQ - Ourimbah, NSW; 26° 26’S, 152° 17’E - 33° 32’, 151° 22’; 50 - 680m; subtropical vine forests; secure; well protected. Physignathus lesueurii (Gray, 183 1): Big Tblnd - Victoria; 15° 42’S, 145° 16’E - V; sea level - 700m; subtropical & monsoon vine forest; secure; well protected. Varanidae Varanus prasinus (Schlegel, 1839): Moa Is.. Torres Strait; 10° 1 1’S, 142° 16’E; 180m; ‘closed mesic forest’; rare; unprotected. (Whittier & Moeller, 1993). Varanus scalarisMertcns, 1941; Heathlands, FNEQ - Paluma. NEQ; 11°50’S, 142° 39’E- 19° 01’, 146° 13’; sea level - 900m; monsoon vine forests, open forests; secure; well protected. DISTRIBUTION OF FROGS AND REPTILES 195 Vartmus teriae Sprackland, 1991: Buthen Buthen - Lankelly Ck, FNEQ; 13° 2rS, 143° 27’E - 13° 53’ 143° 17’; 100 - 500m; monsoon vine forest; rare; unprotected. Varanus vanW (White & Shaw, 1790): Shipton’s Flat, NEQ - Victoria; 15° 48’S, 145° I4’E -V- sea level - 700m; subtropical & monsoon vine forests, usually from open forests; secure- well protected Colubridae Boiga irregularis (Merrem, 1802): Badu Island, Torres Strait - NSW; 10° 07’S, 142° 07’E - NSW; sea level - 1 100m, dry vine thickets; monsoon & subtropical vine forests, also in open forests; secure; well protected Stegonotus cucullatus (Dumeril, Bibron & Dumeril, 1854): Badu Is, Torres Strait, FNEO - Rollingstone NEO extralimital in PNG; 10°42’S, 142°32’E- 19° 03’. 146° 23’; sea level -250m; monsoon vine forests; secure- well protected. Elapidae Cacophis churchilli WeUs & Wellington. 1985; Thornton Pk - Palm Is.; 16° lO’S, 145° 23’E - 18°40’, 146° 33’; sea level - 1000m; monsoon vine forest; secure; well protected. Cacophis kreffiii Gunther. 1863: Broken R. - Crediton. MEQ; Cooran Plateau, SEQ -Gosford, NSW; 21° lO’S 148 31 E - 21*^ 13 , 148° 30 ; 26° 20’, 152° 50’ - 32*^26’* 151° 21 ’; sea level - 720m; subtropical vine forests, moist & dry sclerophyll forests; secure; well protected. Cacophis squamulosus (Dumeril. Bibron & Dumeril, 1854): Mt William, Eungella NP, MEO - Wolloneong NSW;21®02’S, 148°32’E-34°26’, ]50°53’;sca level- 1 000m; subtropical vine forests; secure; well protected Hemiaspis signata (Jan, 1859): Thornton Pk - Mt Spec, NEQ; Miriam Vale. SEQ - SCNSW; 16° ll’S, 145° 20 E - 18° 57 . 146° 11’; 24° 00’, 151° 34’ - SNSW; sea level -1200m; subtropical vine forest, moist sclerophyll forest & heathlands; secure; well protected. Hoplocephalus stephensii Kreffl, 1869: Kroombit Tops, SEQ - Gosford, NSW; 24° 20’S, 151° Ol’E - 32° 26’, 151° 20’; 300 - lOOOm; subtropical vine forests, heathlands; secure; moderately protected. Notechis scutatus (Peters, 1861): Mt Moffat section, Carnarvon NP; Ocean L., Fraser Is - V; 24° 55’S, 148° 03’ - 24° 58’, 148° 08’; 24° 55’, 153° 16’ - V; sea level - 1000+m; subtropical vine forest, usually from open forests; secure; well protected. Pseudechisporphyriacus (Shaw, 1 794): Big Tblnd - Blue Water Ra., NEQ; Mt Dryander, Conway NP - Crediton, MEQ; Gladstone. SEQ - Victoria; 15°42’S, 145® 17’E - 19*’ 14’. 146° 24’; 20° 16’, 148° 35’ -21° 13’, 148° 32’; 23° 51’, 151° 16’ - V; sea level -1000+m; monsoon & subtropical vine forests; secure; well protected. Rhinoplocephalus nigrescens (Gunther, 1862): Big Tblnd - V; 15° 42’S, 145° 16’E - V; sea level - llOO+m; subtropical & monstwn vine forests and adjacent moist open vegetation; secure; well protected. Tropidechis carinatus (Krefft, 1863): Thornton Pk - Mt Spec, NEQ; Fraser Is., SEQ -Barrington Tops, NSW; 16° lO’S, 145° 23’E - 18° 57’, 146° 1 1’; 24° 48’. 153” 13’ -32° 02’, 151° 24’; sea level - 1374m; subtropical & monsoon vine forests, heaths and moist open forests; secure; well protected. Boidae Morelia amethistina (Schneider, 1801): Cape York P., FNEQ -Townsville, NEQ; 11°48’S, 142°2rE- 19° 20’, 146° 30’; sea level - 1 100m; monsoon vine forests and open forests, extralimital in NG and adjoining islands; secure; well protected. Morelia spilota (Lacepede, 1804): throughout Q; sea level - 1 100m; dry vine thickets, monsoon & subtropical vine forest & open forests; secure; well protected. Morelia viridis (Schlegel, 1872): Claudie R. - Rocky R., FNEQ; extralimital in PNG; 12° 46’S, 143° 17’E - 13° 49’, 143° 28’; sea level - 400m; monsoon vine forest; rare; poorly protected. Typhlopidae Ramphotyphlops nigrescens (Gray, 1845): Kroombit Tops, SEQ - Victoria; 24° 24’S, 150° 59’E - V; sea level - 800m; subtropical vine & moist open forests; secure; n^eralely protected. Ramphotyphlops polygrammicus (Schlegel, 1839): islands of Torres Strait;Cockaloo Ck, Heathlands, FNEQ -Eungella NP, MEQ; 10°23’s ]42°02’E;1 1° 39’. 142° 37’ -21° 10’, 148° 30’; sea level -900m; monsoon vine fnrp^f' ^ippiirp' rnrvHprAfplv nrotp/'lpH Ramphotyphlops silvia In^am & Covacevich, 1993: Fraser Is. NP -Rainbow Beach area, SEQ; 24° 52’S, 153° 16’E - 26° 10’, 152° 55’; sea level - 100m; subtropical vine forest; rare; well protected. CONSERVATION Most of Australia’s rainforests are in Queensland and, in general terms, are now well- protected. They present one of the success stories in the history of the Australian conservation movement. The Rainforest Conservation Society Inc., formed in 1982, played the major role in heightening public awareness of rainforests, con- verting this to political will to increase protection of rainforests and to provide finance to increase research to understand them better. Rainforests of the Wet Tropics, which have the highest frog-rep- tile species endemism known for any part of Australia, were placed on the World Heritage register in 1988; much of the moist rainforest of mideastem Queensland is now protected in Na- tional Parks; moist rainforests of most of southeastern Queensland have been nominated for inclusion on the World Heritage List (Govern- 196 MEMOIRS OF THE QUEENSLAND MUSEUM ment of Australia, 1992); remnants of the dry rainforests of the southern part of Queensland, too, are now protected from further clearance by the Forest Service (Forster et al., 1991), although most are not declared national parks or other specially protected reserves, and there is thus no legislative protection for them. Not protected or not well-protected are major moist rainforests in the far north of the state (on Moa Is., at Somerset and in the Mcllwraith Ra.); dry rainforests north of the Carnarvon Ra., MEQ; and small, isolated patches of dry rainforest on the western side of Cape York Peninsula. Gazettal as National Park, the highest level of protection possible for a forest in Queensland, cannot guarantee protection. The Forty Mile Scrub near Mt Garnet, NEQ, has been National Park since 1970. This is a semievergreen vine thicket. Here, following disturbance of the ground by feral pigs, there has been an invasion by Lantana cainara, which has created a rich understorey. This, in turn, has created a fire hazard in a forest which was previously highly fire resistant; and fire has destroyed substantial portions of this forest (R. Fensham, pers. comm.). Moist rainforest on Moa Is., although not safe in terms of special protective status by legislation, is in excellent condition and, at least for the foreseeable future, safe by default. Moa Islanders are sea-farers, and have thus made virtually no incursions into the rainforest on the high part of their island (J. Whittier, pers. comm.). Dry rain- forest (= vine forest and vine thicket) on the western side of Cape York Peninsula, near Weipa, NWQ, has recently been surveyed (Cameron & Cogger, 1992). They note these habitats are small and fragmented and that their frog-reptile fauna is depauperate, a feature they share with other rainforests on the Cape. (Species paucity in northern Cape York Peninsula rain- forests vs extraordinarily high species diversity in southern Peninsula rainforests has been discussed many times, most recently by Covacevich, in press. Those of the north have had ‘... a discon- tinuous history because of climatic fluctuations during the Quaternary.’..., Cameron & Cogger, 1992). Protection of habitat is fundamental to protect- ing frog and reptile (and other) species, but it does not guarantee their survival . The alarming case of unexplained declines in populations of numerous species of rainforest frogs attests to this. First signs of declines in rainforest frog populations were reported in the late 1970s. (Winter & Mc- Donald, 1986) who noted possible problems with Rheobatrachus vitellinus and Taudactylus eun- gellensis at low altitudes in the Eungella NP, MEQ. McDonald (1990) undertook special sur- veys of R. vitellinus, T. eungellensis and T. liemi in the area. His work showed that R. vitellinus disappeared in March, 1985; that T. eungellensis had disappeared in June, 1986; and that T. liemi appeared safe. T. eungellensis has been found once since then (Couper, 1992) giving a glimmer of hope for this species. In southeastern Queensland the congeners of these highly specialised frogs have not been seen, despite in- tensive searching, since 1979 (Czechura & In- gram, 1990). The depressing state of affairs with populations of several tropical rainforest frog species in various degrees of decline has been reviewed recently (Richards et al., 1993). Two species (Litoria nyakalensis and T. rheophilus) were not seen in 1991 -2 surveys; the former being seen last in November, 1990, the latter in January, 1990. For other species (L. nannotis, L. rheocola, L. lorica, Nyctimystes dayi, T. acutirostris) grave fears are held. Populations of some of these in some well-documented sites have disappeared; others are clearly in decline across their whole range. Although natural population fluctuations are well known for frog species (e.g. Peachmann et al., 1991), these can be difficult to distinguish from true declines. Richards et al. (1993) con- clude that the declines they report are ‘real ’ ; those reported by Czechura and Ingram (1990) certain- ly appear ‘real’, if a 14-year absence has any meaning. Several explanations have been ad- vanced to explain declines in frog populations elsewhere. These are summarized and excluded as relevant to the problem in the Wet Tropics frogs by Richards et al. (1993). Factors which do not appear to be relevant to these declines in the Wet Tropics and southeastern Quieensland are: destruction of habitat; historic uses of sites for timber-gathering or mining; other direct human factors (collecting, road mortality); acid-rain; heavy metal, pesticide or other water pollution; drought-induced lowering of the water table; and flooding. Damage to stream habitats by feral pigs had increased at one study site and could be neither discounted nor verified as a causal factor. Similarly, the role of climatic change could not be fully assessed and was neither blamed nor discounted. By any standard the habitats of all these frog species are extremely well-protected and have been for considerable time. Declines in other taxa DISTRIBUTION OF FROGS AND REPTILES 197 in these habitats have not been observed, but that does not, of course, mean they have not taken place. Nor have they been reported in frog species from habitats other than rainforest, with one ex- ception. (The exception is Litoria aurea, a southern species from open forests, generally at low altitudes, G. Ingram, pers. comm.). There is at present no clear reason to account for the declines or disappearances documented and, ob- viously, no clear course to follow to attempt to save the frogs. The words of Czechura and In- gram (1990) are as accurate and worrying now as they were when written ‘... it is very difficult to decide whether or not the missing frogs are ex- tinct. Extinction is not easy to prove. If the miss- ing frogs are extinct, their passing was not a slow process. Rather it appears to have been a catastrophic event that could not have been an- ticipated or prevented.’ Tlie fauna authority in Queensland (Depart- ment of Environment & Heritage), Australian Nature Conservation Agency, James Cook University, the Wet Tropics Management Agen- cy, the University of Queensland and the Univer- sity of Newcastle are supporting the following projects under an umbrella study of declining frogs of Queensland and New South Wales (Mc- Donald): survey of the distribution and status of Litoria longirostris (McDonald); monitoring declining frog populations of the Wet Tropics Biogeographic Region (McDonald); genetic study of several threatened frog species of the Wet Tropics (Moritz & Cunningham); intensive survey of tadpole populations of streams in dif- ferent catchments of the Wet Tropics (Trenerry); study of mortality rates of semi-captive popula- tions of T. acutirostris and T. rheophilus (Mahoney & Dennis); captive breeding of two species of threatened stream-dwelling frogs in the Wet Tropics (Richards & Alford); preliminary survey of diseases of Wet Tropics frogs (Spearc); community ecology of Eungella stream-dwelling frogs (Hero, McDonald, Alford); survey of dis- tribution and abundance of rare or threatened stream-dwelling frogs, SEQ (McDonald & In- gram); survey to assess the status of T. pleione (McDonald & Hero). In addition, the Wet Tropics Management Agency will fund a 3-year study of the ecology and conservation of declining frogs (Wet Tropics Management Agency, in litt.) and the Endangered Species Program will fund a survey of declining frogs in New South Wales. From these studies it is hoped that it will be possible to devise remedial strategies. Since January, 1 984, 8 new species of frogs and 24 new species of reptiles have been described from Queensland’s rainforests. Increased public interest in rainforests has spearheaded a dramatic increase in research on rainforest fauna in recent years. This has led to a rate of discovery and description of new taxa that equals or surpasses the previous great eras of discovery of frog and reptile species between l%0-80, and pre-19()0. For many rainforest taxa, especially inver- tebrates, only a small proportion of the species are described. For the frogs and reptiles, however, the stage has been reached where the species and their broad limits of occurrence are fairly well-known. There are notable exceptions to this suggestion. Cryptic species are known within Mixophyes scfievilli (M. Mahoney, pers. comm.)and Gny- petoscincus queenslandiae (C. Moritz, pers. comm.) as they are presently defined; remote, as yet unexplored, specialized habitats may yet yield new species; and, amongst the burrowing forms, there are probably undescribed taxa. Notwithstanding the facts that much is now known about rainforest frog and reptile species and that many are now secure and moderately to well protect^ (Table 2), much remains to be learnt. Several species remain endangered or vul- nerable, and either poorly protected or un- protected (Table 2). Several species from the Mcllwraith Ra., FNEQ urgently need protection in reserves (e.g. L. eucnemis, L longirostris, C. crepitans, C. peninsularis and S. occulius^ all of which are either rare or insufficiently known and poorly or not protected). Precise details of their distribution are lacking, as are any data on breeding or other habits for most species. Further, there are some rainforests in which no herpelological work has been undertaken. One example is Nangur SF, only 250km NW of Brisbane. In late 1992, this semievergrecn vine thicket was investigated hcr- petologically for the first time. N. spinosa, a dis- tinct member of the Sphenomorphus group has since been described from there (Covacevich et al., 1993). No survey work has been undertaken in the Moa Is. moist rainforest and in many other semievergreen vine ihickeLs of both southern and northern Queensland. In Queensland much of the research money spent in recent years on vertebrates has been used on either rare and threatened species or on those from rainforests. From this, valuable data have been published, and there are now reliable data for some species. Reasonable decisions regarding 198 MEMOIRS OF THE QUEENSLAND MUSEUM Table 2. Conservation status and levels of protection of Queensland’s rainforest frog and reptile species. LEVEL OF PROTECTION CONSERVATION STATUS TOTAL E v R S IK Well 5(0) 4(1) 7(10) 16(43) 1(2) 33(56) Moderately 0(0) 0(0) 3(5) 1(17) 0(0) 4(21) Poor 2(0) 0(0) 0(3) 1(5) 3(2) 6(11) Unprotected 0(0) 0(1) 2(5) 0(0) 1(2) 3(8) E (endangered); V (vulnerable); R (rare); S (secure); IK (insufficient- ly known); unbracketed - frogs; bracketed = reptiles. plans for future conservation of rainforest frogs and reptiles can now be made. With finite resources it is important to remem- ber that rainforest is not the only important habitat in Queensland and that rainforest frogs and rep- tiles are, generally, reasonably well-researched taxonomically and well-protected. Few ecologi- cal data are available, however, for most species, and lack of data is a major problem. Efforts must continue towards solving the disappearing frogs’ problem and others that may occur, and building a store of knowledge about the ecology of rain- forest species. There is now, however, a reasonable case to be made for some shift of emphasis to other species in other habitats more vulnerable and less well-known (e.g. deserts, black soil plains, heaths and grasslands). ACKNOWLEDGMENTS Ross Sadlier, Elizabeth Cameron, Ross Alford, Steven Richards, Patrick Couper, Glen Ingram, Greg Czechura and staff of the Queensland Department of Environment and Heritage have generously assisted us with this paper. The Na- tional Rainforest Conservation Programme, The Wet Tropics Management Agency and the Australian Nature Conservation Agency provided funds which contributed to compiling these data. LITERATURE CITED CAMERON, E.E. & COGGER, H.G. 1992. The her- petofauna of the Weipa Region, Cape York Peninsula. Technical Reports of the Australian Museum. No. 7:1-200. CORBEN, C.J. & INGRAM, G.J. 1987. A new barred river frog (Myobatrachidae: Mixophyes). Memoirs of the Queensland Museum 25( 1 ): 233- 37. COUPER, P.J. 1992. Hope for our missing frogs. Wildlife Australia. 29(4): 11-12. COUPER, P.J., COVACEVICH, J.A. & MORITZ, C. 1993. A review of the Leaf-tailed Geckos en- demic to eastern Australia: a new genus, four new species, and other new data. Memoirs of the Queensland Museum 34(1): 95-124. COVACEVICH, J. in press. Rainforest reptiles of Australia’s World Heritage Wet Tropics: zoogeography and conservation issues. Journal of the International Herpetological Symposium. COVACEVICH, J. & MCDONALD, K.R. 1991. Frogs and reptiles of tropical and subtropical eastern Australian rainforests: distribution pat- terns and conservation. Pp.281-3(>9. In Werien, G. & Kershaw P. (eds), The rainforest legacy, Australian National Rainforests Study, Volume 2 - flora and fauna of the rainforests’. Australian Heritage Commission Special Australian Heritage Publication Series Number 7(2). (Australian Government Publishing Service: Canberra). 414pp. COVACEVICH, J.A., COUPER, P.J. & JAMES, C. 1993. A new skink, Nangura spinosa gen. et sp. nov., from a dry rainforest of southeastern Queensland. Memoirs of the Queensland Museum 34(1): 159-168. CZECHURA, G.V. & INGRAM, G.J. 1990. Taudac- tylus diurnus and the case of the disappearing frogs. Memoirs of the Queensland Museum. 29(2):261-5. DAVIES, M., McDonald, k.r. & adams, m. 1986. A new species of green tree frog from Queensland Australia. (Anura: Hylidae). Proceedings of the Royal Society of Victoria 98:63-71. FORSTER, P.I., BOSTOCK, PD.D., BIRD, L.H. & BEAN, A.R. 1991. ‘Vineforest plant atlas of south-east Queensland. An assessment of conser- vation status’. (Queensland Government, Queensland Herbarium: Brisbane). 466pp. GOVERNMENT OF AUSTRALIA. 1992. ‘Nomina- tion of central eastern rainforests of Australia for inclusion on the World Heritage List’. (Depart- ment of the Arts, Spoit, the Environment and Territories: Canberra). 270pp. GREER, A.E. 1 985. A new species of Sphenomorphus from northeastern Queensland. Journal of Her- petology. l9(4):469-473. 1992. Revision of species previously associated with the Australian scincoid lizard Eulamprus tenuis. Records of the Australian Museum. 44:7- 19. GREER, A.E. & COGGER, H.G. 1985. Systematics of the reduce-Iimbed and limbless skinks currently assigned to the genus Anomalopus (Lacertilia: DISTRIBUTION OF FROGS AND REPTILES 199 Scincidae). Records of the Australian Museum. 37(1): 11-54. GREER, A.E., McDONALD, K.R. & LAWRIE, B.C. 1983. Three new species of Lerista (Scincidae) from northern Queensland with a diagnosis of the wilkinsi group. Journal of Herpetology. 17(3):247-255. HORSUP, A. JAMES, C. & PORTER G. 1993. Sur- veys of dry rainforest vertebrates, south and mideastern Queensland. Memoirs of the Queensland Museum. 34(1): 215-228. INGRAM, G.J. 1991. Five new skinks from Queensland rainforests. Memoirs of the Queensland Museum. 30(3):443-453. INGRAM, G.J. & COVACEVICH, J. 1988. Revision of the genus Lygisaurm de Vis (Scincidae: Rep- tilia) in Australia. Memoirs of the Queensland Museum. 25(2):335-354. 1989. Revision of the genus Carlia (Reptilia, Scin- cidae) in Australia with comments on Carlia bicarinata of New Guinea. Memoirs of the Queensland Museum. 27(2):443-490. 1993. Two new species of striped blind snake. Memoirs of the Queensland Museum 34( 1 ): 1 8 1 - 184. McDonald, K.R. 1990. Rheobatrachus Liem and Taudactylus Straughan & Lee (Anura: Leptodac- tylidae) in Eungella National Park, Queensland: distribution and decline. Transactions of the Royal Society of South Australia. 114(4):187- 194. 1992. Distribution patterns and conservation status of north Queensland rainforest frogs. Conserva- tion Technical Report No. 1 , Queensland Depart- ment of Environment and Heritage, 51pp. PECHMAN, J.F.K., SCOTT, D.E., SEMLITSCH, R.D., CALDWELL, J.P., VITT, L.J. & GIB- BONS, J.W. 1991. Declining amphibian popula- tions: the problem of separating human impacts from natural fluctuations. Science 253: 892-95. RICHARDS, SJ. McDonald, k.r. & alford, R.A. 1993. Declines in populations of Australia’s endemic tropical rainforest frogs. Pacific Conser- vation Biology. 1:66-77. RICHARDS, S.J., McDonald, k.r. & Ingram, G.J. 1993. Recognition of Litoria eucnemis (Lonnberg, 1900) in Australia. Memoirs of the Queensland Museum 34 (1);94. SADLIER, R.A., COLGAN, D.J. & SHEA, G.M. 1993. Taxonomy and distribution of the scincoid lizard Saproscincus challengeri and related species in southeastern Australia. Memoirs of the Queensland Museum 34(1): 139-158. SPRACKLAND, R.G. 1991. Taxonomic review of the Varanus prasinus group with description of two new species. Memoirs of the Queensland Museum 30(3):56 1 -576. THOMAS, M.B.& McDonald, W.J.F. 1989. ‘Rare and threatened plants of Queensland’. 2nd ed. (Department of Primary Industries: Brisbane). WELLS, R.W. & WELLINGTON, R.C. 1985. A clas- sification of Reptilia of Australia. Australian Journal of Herpetology, Supplementary Series No. 1: 1-61. (Australian Biological Services: Sydney). WHITTIER, J.M. & MOELLER, D. 1993. Varanus prasinus (the Emerald Goanna) on Moa Island, Torres Strait, Australia. Memoirs of the Queensland Museum. 34(1): 130. WINTER, J. & McDonald, K. 1986. Eungella, the land of cloud. Australian Natural History. 22(l):39-43. ZWEIFEL, R.G. 1985. Australian frogs of the family microhylidae. Bulletin of the American Museum of Natural History. 182:265-388. 200 MEMOIRS OF THE QUEENSLAND MUSEUM A NEW REPRODUCTIVE MODE FOR AN AUSTRALIAN HYLID FROG. Memoirs of the Queensland Museum 34(1)200. 7995 > The rare frog, LiYona longirostris. was described (Tyler & Davies, 1977) from six specimens collected from the headwaters of the Rocky R., Mcllwraith Ra., Cape York Peninsula, FNEQ. Collectors of the type series observed a clump of ‘bluish-gray eggs* 30 cm above water level on a tree buttress, but did not know whether or not they were eggs of L longirostris. As part of a survey to determine the status of declining and possibly declining frogs, we visited the Mcllwraith Ra., from 14- ISSeptember. 1993. On the banks ofPeachCk(13°44’S, 143” 20’E) and Leo Ck (13°22’S, 143“ 22'E). in a notophyll vine forest, we found nests, eggs and embryos of L lon- girostris. Four nests were guarded by L. longirostris which straddled the nests. Eggs and embryos were lime green. Twenty three egg masses were found on dry substrate above the water, on trunks of Water Gum (Tristaniopsis exciliflora) (7); Rainforest Satinash {Syzigium sp.) (2); on granite rocks in Leo Ck (8); under leaves of rainforest shrubs (2), a living Licuala ramsayi palm frond (I); dead L. ramsayi fronds (2); and on the upper surface of a frond of Cyathea rebeccae Eggs had been laid from 4.5 - U 7.5 cm above the water (mean 26.36, SD 26.54, n 22) in areas where water depth immedi- ately below egg masses varied from 2.0 - 71.0 cm (mean 22.26, SD 18.69, n 20). All but one were associated with still water pools. Pool depths ranged from 15.5 - 120.0 cm (mean 49.74, SD 36.06. n 17). The exception (on the C rebeccae frond) hung over gently flowing water. Size of pools ranged from 0.5 m adjacent to fast flowing streams, to approximately 10 m in streams with gentle (- north-eastern Queensland, Individuals SM 20, 21 NEQ b - north-eastern Queensland, individuals SM 15, 05, 111 MQ - middle eastern Queensland, individuals SM 75, C506, C507 SE a - south-eastern Queensland, individuals SM 62, 71 SE b - south-eastern Queensland, individual SM 59 lie Cys Leu Met Thr Gin 11# lie Thr Gly Leu Leu Leu Ala Met HlA Tyr Thr NEQ s NEQ b ATC TUC CTA ATA ACT CAA ATC ATC ACA GGC CTC CTG CTA GCC ATA CAC TAC ACG MQ . .T SE a . .T SE b . .T Ala Aap Thr Thr Leu Ala Phe 5er Ser Val Ala Hla Met Cya Arg Aan Val Gin Phe Gly Trp MEQ a GCA GAC ACT ACA CTA GCC TTC TCC TCT GTC GCT CAT ATA TGC CGA AAC GTC CAA TTC GGC TGA NEQ b MQ . .G SE a . .G . .C . .C . .T . .C . .G SE b . .G NEQ a NEQ b MQ SE a Leu He Arg Aan Leu Hia Ala Aan Gly Ala Ser Phe Phe Phe He Cys He Tyr Leu Hla He CTC ATC CGC AAC CTC CAT GCA AAC GGG GCT TCA TTC TTC TTT ATC TGC ATC TAC CTT CAC ATC SE b NEQ a Gly Arg Gly Phe Tyr Tyr Arg Ser Tyr Leu Aan Lya Glu Thr Trp Aan Val Gly Val He Leu GGA CGA GGA TTC TAC TAC GGC TCC TAC TTG AAC AAA GAA ACC TGA AAC GTC GGA GTT ATC CTC NEQ b MQ A. . . .G SE a A. . . .G SE b Leu Leu Ala Leu Met Ala Thr Ala Phe Val Pro Met NEQ a CTC CTG GCC CTT ATA GCA ACT GCT TTC GTA CCT ATG NEQ b MQ . .A A.T SE a . .A A.T SE b . .A A.T FIG. 3. Sequence of part of the cytochrome b gene in the Large-billed Scrubwren in individuals from NEQ, MQ and SE. In this and Figs 4 and 7, the amino acid sequence is included for the purposes of a) verifying that true mtDNA sequence and not non-functional nuclear copies have been obtained and b) making the amino acid sequence itself available for any subsequent phylogenetic analyses at that level. MITOCHONDRIAL DNA OF QUEENSLAND BIRDS 207 White browed Scrubwren Sericornis frontslis cytochro/re h sequence starting from position 14855 relative to the complete human mtDNA sequence SF Leu Gly He Cys Leu Met Thr Gin He lie Thr Arg Leu Leu Leu Ala Met His Tyr Thr 3 1 NEQ A CTA GGT ATC TGC CTA ATA ACT CAA ATC ATC ACA GGC CTT CTA CTG GCC ATA CAC TAC ACA SF 4 1 NEQ SF 5 1 NEQ SF 32 MQ SF 33 MQ SF 35 MQ SF 37 MQ SF 2 SE SF 30 SE Ala Asp Thr Thr Leu Ala Phe Ser Cys Val Ala His Met Cys Arg Asn Val Gin Phe Gly Trp SF 3 : NEQ GCA GAC ACC ACC CTA GCT TTC TCT TCC GTT GCC CAC ATA TGC CGA AAC GTC CAA TTT GGC TGA SF 4 NEQ SF 5 NEQ SF 32 MQ SF 33 MQ SF 35 MQ SF 37 MQ SF 2 SE SF 30 SE Leu He Arg Asn Leu His Ala Asn Gly Ala Ser Phe Phe Phe He Cys He Tyr Leu His He SF 3 NEQ CTT ATC CGC AAC CTT CAC GCA AAC GGA GCC TCA TTC TTT TTT ATC TGC ATC TAC CTC CAC ATC SF 4 NEQ SF 5 NEQ SF 32 MQ SF 33 MQ SF 35 MQ SF 37 MQ SF 2 SE SF 30 SE Gly Arg Gly Phe Tyr Tyr Gly Ser Tyr Leu Asn Lys Glu Thr Trp Asn Val Gly Val He Leu SF 3 NEQ GGA CGG GGC TTT TAC TAC GGC TCC TAC CTA AAC AAA GAA ACC TGA AAC GTC GGA GTA ATC CTC SF 4 NEQ . .G SF 5 NEQ SF 32 MQ SF 33 MQ SF 37 MQ . .G SF 2 SE SF 30 ' SE Leu Ser Ser Pro His Ser Asn Cys Leu SF 3 NEQ CTC TCT AGC CCT CAT AGC AAC TGC CTT SF 4 NEQ SF 5 NEQ SF 33 MQ SF 2 SE SF 30 SE FIG. 4. Sequence of part of the cytochrome b gene in the White-browed Scmbwren in individuals from NEQ, MQ and SE. parisons within and among populations reaching 0.01 only within NEQ and within SE. As was the case with RFLP analysis, phylogenetic analysis of the small number of informative sites in the sequence data showed no consistent pattern of relationships among the three areas SE, MQ and NEQ (Fig. 5). Eastern Whipbird In the RFLP analysis, between 46 and 5 1 sites unambiguously were inferred after digesting the mtDNA genomes with the same 17 restriction endonucleases as used for Sericornis (Appendix 2, Table 4). The MQ sample was identical with one from southeastern Queensland, both of which in turn were very similar to the individuals from New South Wales (sequence divergences 0.2 ± 0.2% to 1.1 ± 0.4%). In contrast, in comparisons between MQ/SE and the individual from NEQ divergences reached between 2.5 ± 0.7% and 2.8 ± 0.8% (Table 5). Nucleotide diversities sug- gested a similar pattern, being 0.007 within SE (only single individuals were available from MQ and NEQ) and 0.009 between MQ and SE but 0.028 in comparisons between MQ and NEQ and 0.009 between SE and NEQ (Table 3). Structur- ing of mtDNA diversity in the limited sample of this species was most strongly suggested between NEQ and MQ/SE combined. No RFLP data were available for use as an outgroup in phylogenetic analysis for this species so an unrooted phylogenetic network linking all haplotypes was constructed using only unambiguous characters (Fig. 6a). In a tree 21 steps in length the single NEQ individual was on a branch separate from all other samples while those from south-eastern and middle eastern Queensland, which were the same haplotype, were aligned with one of the New 208 MEMOIRS OF THE QUEENSLAND MUSEUM White-browed Scrubwren Sericornis frontalis -shortest tree {17 steps) with branch lengths above and bootstrap values (500 replicates) below; MQ individuals italicized ’ |SF3 NEQ |SF2 SE 58 SF37 'I SF32 \ 2 SF35 J 79 SF5 NEQ 1 SF30 SE - SF4 NEQ 12 FIG. 5. Tree produced from analysis of sequence data in the White-browed Scrubwren showing bootstrap values and branch lengths. South Wales samples. Placing MQ with NEQ produced a tree one step longer that better ex- plained only one character compared with two better explained in the tree aligning MQ with SE. In sequence analyses, up to 264 bp were se- quenced in six individuals with a minimum of 228 bp (Fig. 7). Seven nucleotide sites varied across NEQ, MQ and SE. Among all the MQ and SE individuals, divergence ranged from zero (be- tween C529 and Ps 1 from middle eastern and south-eastern Queensland, respectively) to 0.9% and 1.1% in comparisons among SE individuals. Between SE/MQ and NEQ individuals divergen- ces ranged from 1% to 3% while divergence between the two NEQ individuals was 0.9%. Nucleotide diversities weakly suggested structur- ing to the diversity with between population values being generally lower than those within populations but with the highest value being in the NEQ-MQ comparison (Table 3). Though few phylogenetically informative characters were available, phylogenetic analysis was done using sequence from the Grey-crowned Babbler (Ed- wards et al., 1991) as an outgroup. A shortest tree 33 steps in length was produced that grouped the two NEQ individuals apart from all SE and MQ individuals (Fig. 6b). Placing the single MQ in- dividual with NEQ did not better explain any characters and resulted in a tree two steps longer. DISCUSSION mtDNA Phylogeography To shed light on the historical biogeography of MQ rainforests we have looked at the phylogeographic structure of mtDNA diversity in three bird species common to MQ, SE and NEQ rainforests and asked whether the structures are consistent with isolation by distance or vicariance models. The patterns revealed by RFLP and se- quence analyses in the three species in general were concordant in that where geographic struc- turing was present, the patterns were the same. In this regard, the identity between MQ and SE individuals at the sequence level in the Large- billed Scrubwren was striking, although RFLP assays did reveal some variation in other parts of the mtDNA genome. These patterns strongly sug- gest that in the Large-billed Scrubwren and, to a lesser extent the Eastern Whipbird, mtDNA diversity is apportioned between SE and MQ populations on one hand and NEQ on the other. For these two species, we can favour the hypothesis of a vicariance event about the Bur- dekin Gap having isolated MQ and SE popula- tions from NEQ ones. If isolation by distance has operated in these species, then its effects are minor compared with those of vicariance induced by the Burdekin Gap. This in turn implies that the Burdekin Gap has been a long-standing biogeographical barrier, a conclusion consistent with geographical patterns of morphological variation as expressed in existing subspecific taxonomy in these two species. In the White-browed Scrubwren, however, there was no clear geographic structure despite substantial variation in RFLP analyses and the phylogenetic relationships among the mtDNA alleles seem poorly resolved; the greatest diver- gence was between some SE haplotypes. The situation in this species is consistent with neither the isolation by distance nor the vicariance model and we suggest that this species has not evolved a stable phylogeographic structure in mtDNA diversity among its Queensland populations; the situation is at present less clear with regard to morphological diversity (Ford, 1985). This may be attributable to the species’ far greater habitat diversity relative to the other two species studied, as this probably facilitates rapid and widespread dispersal. Christidis & Schtxide (1991) argued from allozyme data that a zone of secondary intergradation in the White-browed Scrubwren occurs about the latitude of Rockhampton, which is south of the MQ rainforests. Unfortunately, MITOCHONDRIAL DNA OF QUEENSLAND BIRDS 209 Ava 1 Bam Bel / HI Bgl II Dra 1 Eco 0109 Eco RI Eco RV Hind III Nco 1 Nhe I Pvu II Spel Hinp 1 Hirrf 1 Msp I New South Wales D750 AAA A A A A A A A A A A A A A L226 ? ? L255 B B840 B B B B B C291 B B B B B South-East Queensland PsI . B B B B C B A? A? A? Middle Eastern Queensland C529 . B B B B C B - A? 7 North-East Queensland C564 B C B C C B C D B C C B A? C U/iO: Nowra, NSW L226: Kiola, NSW L255: Kangaroo Valley, NSW B840: Cambridge Plateau, NSW C29I: Mebbin State Forest, NSW Ps 1: Conondale Ranges, South-East Queensland C529: Cathu State Forest, Middle Eastern Queensland C564: Mission Beach, North-East Queensland TABLE 4. Restriction fragment profiles in the Eastern Whipbird. Question marks indicate uncertain typings. they were only able to analyse NEQ populations for one locus {AID) and further analysis of al- lozymes in northeastern Queensland populations would be desirable. The mtDNA data, then, provide a basis for suggesting that isolation of the MQ and SE rain- forests from each other is very recent compared with that of the NEQ rainforests from those of SE and MQ. Further, we may conclude that vicariance induced by the Burdekin Gap has been a major force in shaping patterns of genetic diver- sity within rainforest birds such as the Large- billed Scrubwren and Eastern Whipbird as well as in shaping species distributions. The White- browed Scrubwren, however, is a species for which mtDNA diversity and geographical range appear not to have been so affected; its broader habitat range and likely greater dispersal abilities may act to override the evolution of any phylogeographic structure. Speculations on MQ Biogeography The mtDNA evidence indicates recent connec- tions between SE and MQ rainforests. These con- nections could have been either of two types: (i) continuous tracts of rainforest only recently broken into present disjunct isolates or, (ii)scat- tered patches of a habitat such as vine thickets or dry rainforest that could have existed for much of the Pleistocene. Either interpretation would be consistent with the patterns of genetic diversity we have reported here, but the second is more consistent with both the absence in MQ of several highly specialized rainforest species of SE and NEQ rainforests, and the presence of less special- ized species such as we have studied here in each of SE, MQ and NEQ rainforests. Further, the lack of substantial rainforest connections could have promoted the evolution of MQ rainforest en- demics (Table 1). What remains to be explained is the initial absence from MQ of several rain- forest species found in both NEQ and SE rain- forests. As suggested by Winter ( 1 988) to explain the dearth of rainforest specialists among the mammals of SE rainforests, it could be that MQ rainforests contracted to an area too small to support obligate rainforest species during the driest period of the Pleistocene glacials. How- ever, the absence of the Yellow-throated Scrubwren 5. citreogidaris, a species now capable of occupying wet sclerophyll as well as rainforest habitats, remains difficult to under- stand. The mtDNA divergence between the SE and NEQ populations is at least as great as that between those of the Large-billed Scrubwren (Joseph & Moritz, 1993b) so they might be ex- pected to have reacted similarly to habitat chan- ges and vicariance events. A prediction from the second suggestion is that the endemic species of MQ rainforests are des- 210 MEMOIRS OF THE QUEENSLAND MUSEUM D 750, L 226 (NSW) L 255 (NSW) D 750, L 226 (NSW) 49 48 L 255 (NSW) 0.2 ± 0.2 48 B 840, C 291 (NSW) 0.9 ± 0.4 1.1 ± 0.4 Ps 1 (SE Qld), C 529 (MQ) 1.0 ± 0.4 1.2 ± 0.4 C 564 (NEQ) 2.5 ± 0.7 2.7 ± 0.8 B 840. C 291 (NSW) Ps 1 (SE Qld), C 529 (MQ) C 564 (NEQ) 46 47 41 45 46 40 48 47 40 0.9 ± 0.4 51 41 2.7 ± 0.8 2.8 ± 0.8 46 TABLE 5. Matrix of sequence divergence values between different haplotypes in the Eastern Whipbird. Numbers of sites inferred in each haplotype are on the leading diagonal, while numbers of shared sites and sequence divergence values and their standard deviations are above and below the diagonal, respectively. cendants of lineages that were either capable of surviving in small rainforest patches or that were non-rainforest forms. This could be tested by placing habitat preference of MQ endemics, espe- cially the most narrowly distributed ones with rainforest specialization, and their close relatives in a framework of the phylogenetic affinities among the species themselves. One could then argue whether habitat shifts from eucalypt to rainforest have occurred and, if so, whether MQ rainforest endemics are likely derived from non- rainforest inhabiting ancestral lineages. The leaf- Eastem Whipbifd Psophodes oinfaceus a) unrooted phytogenelic netwof1< from RFLP data showing branch lengths SEa j SEb 2 2 3 1 SEQW 3 1 MQ 70 b) shortest tree from sequerx^ data with branch lengths -^291 - L226 J JC529 MQ IpSI SEQkJ 3 ,1 C564 'i 11 ^ tips? J 25 — Grey-crowned Babbler Pomslostoenus Icmptyalis FIG. 6. Trees produced from analysis of RFLP (a) and sequence data (b) in the Eastern Whipbird to show bootstrap values and branch lengths. The letters a to c in (a) correspond to the haplotypes shown in Ap- pendix 2. tailed geckoes Phyllurus spp., which include nar- rowly distributed rainforest endemics in MQ and other species showing varying degrees of depend- ence on rainforest, arc an example of such a group (Couper et al., 1993). The Eungella Honeyeater of MQ rainforests (Table 1) would provide a further useful test as its closest relative is either the Yellow-faced Honeyeater M. chrysops of eastern Australian eucalypt forests or the Bridled Honeyeater M. frenata endemic to NEQ rain- forests (Longmore & Boles, 1983), Habitat shifts from rainforest to eucalypt habitats have long been considered to have been involved in the evolution of the Australian biota but the reverse has rarely, if ever, been proposed. MQ rainforests provide an opportunity to test for such shifts. The existence between MQ and SE of a number of very narrowly distributed species endemic to small, isolated patches of rainforest and wetter forest types would be consistent with long-term habitat connections between MQ and SE rain- forests having been through scattered habitat patches rather than continuous rainforest. Ex- amples are especially prominent in the Kroombit Tops region from where the day frog Taudactylus pleione Czechura, 1 986 and a number of charopid land snails (Stanisic, 1990) are known. The recently discovered and most distinctive scincid genus Nangura from near Murgon (Covacevich et al., 1993) is a further example. In conclusion, we submit that mtDNA in- traspecific phylogeography offers increased resolution over that available so far in studies of the historical biogeography of middle eastern Queensland rainforests. However, more insight may be forthcoming from the other species listed in Table 1 . Phylogenetic analyses of groups with narrowly distributed MQ endemics (e.g day frogs mit(x:hondrial dna of Queensland birds 211 Eastern Whipbird Psopbodes olivaceus cytochrome b sequence starting from position 14868 relative to the human sequence NEQ a - north-eastern Queensland, individual C 564 NEQ b - north-eastern Queensland, individual Ps 7 MQ - middle eastern Queensland, individual C 529 SE a - south-eastern Queensland, individual Ps 1 SE b - eastern New South Wales, individual C 291 SE c - eastern New South Wales, individual L 226 NEQ a Leu He Thr Gin He He Thr Gly Leu Leu Leu Ala Met His Tyr Thr C ACA CAA ATC ATC ACA GGC CTC CTC CTA GCC ATA CAT TAC ACA NEQ b CTA GT. MQ SE a SE b SE c Ala Asp Thr Asn Leu Ala Phe Ser Ser Val Ala His Met Cys Arg Asn Val Gin Phe Gly Trp NEQ a GCA GAC ACC AAC CTA GCC TTC TCC TCT GTA GCC CAC ATA TGC CGA AAT GTA CAA TTC GGA TGA NEQ b MQ SE a SE b SE c NEQ a NEQ b MQ SE a SE b SE c NEQ a NEQ b MQ SE a SE b SE c Leu lie Arg Asn Leu His Ala Asn Gly Ala Ser Phe Phe Phe lie Cys He Tyr Phe His He CTA ATC CGA AAC CTA CAT GCA AAC GGA GCT TCA TTC TTC TTC ATT TGT ATC TAC TTC CAC ATT Glv Arq Gly He Tyr Tyr Gly Ser Tyr Leu Asn Lys Glu Thr Trp Asn Val Gly Val He Leu GGA CGA GGA ATC TAC TAC GGA TCA TAC CTA AAC AAA GAA ACC TGA AAC GTC GGA GTC AT ■ ■ y ' ' " ] ' * ] ’ ] * ’ * [1 C CTC MQ SE a SE b SE c Leu Leu Thr Leu Met Ala Thr Ala Ser CTA CTA ACC CTC ATG GCA ACT GCT TCG G FIG 7. Sequence of part of the cytochrome b gene in the Eastern Whipbird in individuals from NEQ, MQ and SE. Taudactylus spp. and leaf-tailed geckos Phyl- lurus spp.) or those with representatives in the rainforests of all major eastern Australian rain- forest blocks (e.g. the White-throated Tree- creeper and Brown Thombill complexes) are likely to be especially fruitful. ACKNOWLEDGEMENTS CSIRO Postgraduate Scholarship to LJ and an Australian Research Council grant to CM. T. Gorringe took photographs of the figures and tables. Some of the ideas examined in this paper arose in discussions at the Second Broken Head Symposium on Evolutionary Genetics and we thank the participants of that forum for their input. The field work for this project was done with the able assistance of E.C. Edwards, A. Heideman, M. Cunningham, M. Pavelka and E.O. Willis. M. Cunningham advised on distribution and nomenclature of reptiles and S. Degnan and two referees made many helpful comment on a draft. Drs L. Christidis (Museum of Victoria) and R. Schodde (CSIRO) kindly donated some specimens of the Large-billed Scrubwren and Eastern Whipbird. We collected specimens under permits from the Queensland Department of En- vironment and Heritage and the Queensland Forest Service. The project was funded by a LITERATURE CITED AVISE, J.C. 1992. Molecular population structure and the biogeographic history of a regional fauna: a case history with lessons for conservation biol- ogy. Oikos 63: 62-76. AVISE, J.C., ARNOLD, J., BALL, R.M., BERMIN- GHAM, E., LAMB, T, NEIGEL, J.E., REEB, C.A. & SAUNDERS, N.C. 1987. Intraspecific phylogeography: The mitochondrial DNA bridge between population genetics and systematics. Annual Reviews of Ecology and Systematics 18: 489-522. 212 MEMOIRS OF THE QUEENSLAND MUSEUM BERMDMGHAM. E., ROHWER, S.» FREEMAN, S. & WOOD, C. 1992. Vicariance biogeography in the Pleistocene and speciation in North American wood warblers: a test of MengeFs model. Proceedings of the National Academy of Scien- ces, USA 89: 6624-6628. 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Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pul- monata) from subtropical rainforests. Memoirs of the Queensland Museum 30; 1-241. STORR, G.M. 1984. Revised list of Queensland birds. Records of the Western Australian Museum. Supplement Number 19. S WOFFORD, D. 1991. ‘Phylogenetic Analysis Using Parsimony. Version 3.0s’. (Illinois Natural His- tory Survey: Chicago). TRUSWELL, E.M. 1990. Australian rainforests: the 1(X) million year record. Pp 7-22. In Webb, L.J. & Kikkawa, J. (eds), ‘Australian tropical rain- forests - science, values, meaning’. (CSIRO: Melbourne). 185 pp. WALKER, D. 1990, Directions and rates of tropical rainforest processes. Pp 23-32. In Webb, L.J. & Kikkawa, J. (ed.s), ‘Australian tropical rainforests - science, values, meaning’. (CSIRO: Mel- bourne). 185 pp. WEBB, L. & TRACEY, J. 1981. Australian rain- forests: pattern and change. Pp 605-694. In Keast, J.A. (ed.). ‘Ecological biogeography of Australia’. (W. Junk: The Hague). 805 pp. WINTER, J. 1984. Conservation studies of tropical rainforest possums. Pp 469-481. In Smith, A.P. and Hume, I.D. (eds). ‘Possums and gliders’. 214 MEMOIRS OF THE QUEENSLAND MUSEUM (Australian Mammal Society: Sydney). 598pp. WINTER, J. 1988. Ecological specialization of mam- mals in Australian tropical and sub-tropical rain- forest: refugial or ecological determinism? Proceedings of the Ecological Society of Australia 15: 127-138. APPENDIX 1. Restriction site matrix for the Large-billed Scrubwren. Appendix 1. Matrix of restriction sites fronx RFLP data in the Large-billed Scrubwren Sericornis magnirostris. a - haplotype 1 - individuals SM2, 59, 60, 68; b - SM 71; c - SMI; d SM 66; e - SM75; f - SM 79; g - SM105; SF32 - Sericornis frontalis a - dare from south-eastern Queensland; e is from middle eastern Queensland; f and g are from norOi-oastem Queensland. Ava I Bam HI Bcf I Bgl 11 Dra I Eco R1 Eco RV Nhe I Pvu II Sac II Hind III a imno 111010000 111100 1100 1111 no 1 1100 1011000 11 ninnoo b 1111110 111000000 111100 1100 nil 111 1 1100 1111100 11 innnoo c iiniio 111010000 inioo 1100 nil 111 1 1100 1111100 11 111111100 d 1111110 111000000 niioo 1100 nil no 1 1100 1011000 11 ninnoo e 1111110 111011000 inioo 1100 nil 111 1 1100 ninoo 11 innnoo f 1111100 111101000 101100 1000 nil no 1 1110 lonooo n ninnoo g 1101100 111101000 101100 1000 nil no 1 1110 1011000 11 111111101 SF32 1111101 110000111 000011 0011 1110 111 0 1001 0000011 11 111100010 APPENDIX 2. Restriction site matrix for the Eastern Whipbird. Appendix 2. Matrix of restriction sites from RFLP data in the Eastern Whipbird Psophodes oHvaceus. a - haplotype 1 field collection numbers D750, L226 (NSW); b - L255 (NSW); c - B840, C291 (NSW); d - Psl (SE Qld) and C529 (MQ); e- C564 (NE Qld) Aval Bam HI Bel 1 a 11111 110111 101111 b 11111 110111 101111 c 11111 110111 101111 d 11111 111111 111111 e 11111 111111 101100 BglM Dra I Eap0109 EcoRI 101 inn 10101 1110111 101 11111 10101 1110111 101 10101 10111 1110111 101 10101 10111 1110111 111 10111 11101 1111111 EcoRV Hindm Ncol Nhe\ PvuU 11 111111 10101 in nil 11 101111 10101 111 nil 11 111111 11101 111 1011 11 111111 10111 111 nil 11 111011 01101 101 0110 VERTEBRATES OF DRY RAINFOREST OF SOUTH AND MIDEASTERN QUEENSLAND ALAN HORSUP, CAMERON JAMES AND GARY PORTER Horsup, A., James, C. & Porter, G. 1993 12 24; Vertebrates of dry rainforest of south and mideastem Queensland. Memoirs of the Queensland Museum 34(l):215-228. Brisbane. ISSN 0079-8835. In 1992, the Queensland Department of Environment and Heritage undertook a series of surveys of vertebrates of dry rainforests of south and mideastem Queensland. Eighteen sites were surveyed between Rockhampton and Gympie, west to Monto and Biloela. Sites were surveyed twice, six monthly, to examine seasonal variations. Generally, faunal diversity was low. Diversity was affected by the proximity of other rainforest or undisturbed forests; the presence of permanent water; size, type, and degree of disturbance from logging, cattle grazing or weed infestation. Seasonal variation at some sites was considerable. The new skink, Nangura spinosa (Covaccvich et al., 1993) at Nangur SF, near Murgon, SEQ was the most important discovery. □ Dry rainforest, surveys, frogs, Bufo marinus, reptiles, birds, mammals, conservation. A. Horsup, C. James & G. Porter, Queensland Department of Environment and Heritage, PO Box 3130, Rockhampton Shopping Fair, Rockhampton, Queensland 4701, Australia, 27 August 1993. With few exceptions, vertebrates of dry rain- forests of south and mideastem Queensland, be- tween Rockhampton and Gympie, are poorly known. Although this is the most common rain- forest there, little is protected. Much has been cleared, and grazing is common in remaining patches. These rainforests are recognised as im- portant refuges for populations of several temperate and tropical vertebrate species. In the early 1970's, Potorous tridactylus was found al Granite Ck SF, a 200km northern extension of its range (J. Toop, pers. comm.), and Trichosurus caninus and Thylogale thetis have been located at Kroombit Tops, 200km and 300km north of their previous known distributions (Woodall, 1986). METHODS All available information on the vertebrate fauna of the study area was collated by searching the literature (e.g. Ingram & Raven, 1991) and accessing the Queensland Museum and RAOU Bird Atlas Scheme databases. Rainforests were selected to cover a range of vegetation types and if they were of more than 50 ha, of less than secure tenure, of interest for acquisitional purposes by DEH, or linked with other areas of significant rainforest; or threatened by clearing, grazing or fire. Most sites were surveyed twice to lake into account seasonal variations. Each was surveyed over a four day period by the rapid fauna survey method (Denny, 1984). Sites were sampled by the gradsect method of Gillison & Brewer (1985). Two transects were sampled per site. The main axis of each transect was laid out at right angles to the prevailing slope or moisture regime. Over 4 days and nights, fauna records were collected by trapping (200 Elliott, 28 cage, 40 pit-trap, and 40 break-back trap nights), dawn and dusk observations, intensive searches (log-rolling, leaf litter searching), spotlighting, mist-netting, recording bat calls using an electronic detector, and identifying tracks and traces. Voucher specimens of species were lodged in the Queensland Museum wherever possible. RESULTS Study Sites The sites (Table 1, Fig. 1) ranged from 50 to 3000 ha and included six vegetation types (Table 1). Sites included freehold and crown land (NP, SF and TR). Bania SF and Granite Ck SF, contain large tracts of wet rainforest. Bania SF is situated at the head of the Burnett R., S Burnett Ra., and contains the largest stand outside a NP in southeastern Queensland. Smaller sites were selected to cover many rainforest types. Two were surveyed at Granite Ck SF to repeat the Australian-Queensland Museum’s study (Broadbent & Clark, 1976). Little information was available previously on the vertebrate species 216 MEMOIRS OF THE QUEENSLAND MUSEUM of the sites chosen. There are species lists for Granite Ck SF (Broadbent & Clark, 1976), Dan Dan SF (Queensland Forest Service), Goodnight Scrub (Robinson, 1977), and Rundle Ra. (Stock et al., 1988). Amphibians Nineteen species of native frogs (myobatrachids, hylids), and the introduced Cane Toad, Bufo marinus, were recorded (Appendix 1). All occur widely elsewhere in Australia. None can be considered endangered or threatened. The amphibians encountered in this study are typical- ly northern tropical (8 species) and eastern regional (6), with a minor inland (3) and southern (2) element. Generally, frog species diversity was low (mean, 3.8 species per site). No native frogs were found at Coominglah SF and, at three sites, only one native species was found. The highest num- ber of frog species was recorded at Boogooram- unya (9). The most commonly encountered and widespread frogs were Litoria caerulea ( 1 1 of 1 8 sites), Limnodynastes ornatus, L latopaimata, and L. lesueuri (all recorded at 7 of 18 sites). B. marinus was widespread, and recorded at all sites except Coongara Rock. As expected, more species and more in- dividuals were recorded in the wet season surveys than in those of the dry season. Twelve of the 19 native frog species were found in both seasons. The other seven species were found only in the wet seasons, and all were recorded at only one site. For example, at Oakview SF, seven native species were recorded in the wet season, none in the dry season. Oakview is a small rainforest (150 ha), with permanent water only 200m from its edge. In contrast, at Bania SF, a large patch of rainforest (>3000 ha), five species were recorded in the wet season, four in the dry season. Thirteen frog species were recorded at the four AMVF sites (mean, 5.5 per site), 10 species at the seven SEVT sites (mean, 2.4 per site), and eight species at the three CNVF (mean, 3.0 per site). Tliese differences are partly due to the SEVT sites being situated generally in the drier northern and western areas, while AMVF sites were more on the wetter coastal ranges (Fig.l). L. latopalmata and L. rubella were the most widespread native species, occurring in five of the six vegetation types. Reptiles Forty-nine species of reptile (agamids, gek- konids, pygopodids, scincids, varanids, boids, colubrids and elapids) were recorded (Appendix 1). All are native to Australia and, except for Nangura spinosa, occur widely elsewhere. Nineteen reptile species are eastern with pre- dominantly coastal distributions from NNSW to NEQ. Nine are endemic to coastal areas of MEQ and SEQ. Of these, Phyllurus caudiannulafus, Ophioscincus ophioscincus^ Calyptotis lepidorostrum, Lampropholis adonis, and L. couperi, are restricted to rainforest and wet sclerophyll forests, and V. spinosa is restricted to one patch of semievergreen vine thicket (Covacevich et al., 1993). Three species endemic Table 1. Survey sites, tenure (NP=National Park, TR=Timber Reserve, SF=State Forest, F=Freehold), site number (after Forster et al., 1991) vegetation, size, and time(l 992). Mt Etna Caves NP, (1), SEVT, semievergreen vine thicket, 300 ha. Mar. Rundle Range NP (10), semievergreen vine thicket, 500 ha, Apr/Oct. Mt Larcom F, (14), semievergreen vine thicket, 250 ha, Apr. Spring Creek F. (17), semievergreen vine thicket, 250 ha, Aug/Dec. Dan Dan SF 53, 08), semievergreen vine thicket, 835 ha, Mar/Aug. Coominglah SF 28, (38), semievergreen vine thicket, 1674 ha, Mar/Aug. Nangur SF 74, (92), semievergreen vine thicket, 500 ha, Jun/Nov, Deep Creek TR 581, (61), SEVT/AMVF, semievergreen vine thicket/Araucarian microphyll vine forest, 180 ha, May/Nov. Coongara Rock SF 1344, (69), semievergrecn vine thicket/Araucarian microphyll vine forest, 50 ha, Feb/Jul. Mt Coulston TR 471, (20), AMVF, Araucarian microphyll vine forest, 700 ha, May/Nov. Goodnight Scrub SF 169, (46), Araucarian microphyll vine forest, 250 ha, May/Oct. Boogooramunya SF 465, (77), Araucarian microphyll vine forest, 180 ha, Jun/Nov. Wrattens SF 546, (107), Araucarian microphyll vine forest, 210 ha, Mar/Sep. Bania SF 54, (39X AMVF/ANVF, Araucarian microphyll vine forest/Araucarian notophyll vine forest, 3000 ha, Apr/Nov. Oakview SF 220, (99), ANVF, Araucarian notophyll vine forest, 150 ha, Mar/Sep. Eurimbulah NP, (BB), CNVF, complex notophyll vine forest, KX) ha, Apr/Aug. Boyne Logging Area SF 391, (30), complex notophyll vine forest, 3000 ha, J un/Dec. Dawes Range SF 391, (33), complex notophyll vine forest, 3000 ha, Jun/Dec. SURVEY OF DRY RAINFOREST VERTEBRATES 217 f to MEQ and SEQ are not restricted to wetter forests: Glamphyromorphus punctulata (tall woodland and seasonally dry forests), P. salebrosus (dry rocky scrublands to moister closed forest), and L. amicula (open forest and heath). Reptiles were recorded during the study with equal probability in either season: 39 in both wetter and drier surveys, I By far the highest reptile diversity was recorded I at the AMVF and SEVT sites, with 42 and 36 species, respectively, while only 15 species were recorded at CNVF sites. Overall, AMVF-ANVF had the highest mean number of species per site (16), mainly because this rainforest type was present at only one very large site, Bania SF (>3000 ha). There was little difference in the mean number of species per site for the other rainforest types: AMVF (11.8), SEVT (10.6), SEVT- AMVF (12.5), CNVF (7.3). Birds A total of 1 69 bird species were recorded during the study (Appendix I). This represents ap- proximately 25% of the total number of birds known to breed in or regularly migrate to Australia. Of these, 79 were commonly recorded in rainforest, 75 were recorded more often in adjacent drier open forests (but were also found along the ecotone between rainforest and open forest), and 15 are shorebirds or waterbirds. Rainforest bird species which were expected, but were not sighted are: Double-eyed Fig Parrot (race coxeni). Striated Thombill, Little Bronze Cuckoo, Rufous Bronze Cuckoo, White-browed Robin, Southern Log Runner, Large-billed Warbler, Bell Miner, and Black Butcherbird. The three species listed first have been recorded pre- viously at Granite Ck SF in the late 1970's (E. Zillman, pers. comm,). Fifty-seven families were recorded. Dominant were the Muscicapidae (19 species), Meliphagidae (14 species), Columbidae, Acan- thizidae, Accipitridae and Falconidae (all 1 1 species). Most species were recorded at sites in both surveys. The exceptions include the Dollar- bird, Superb Fruit-dove, cuckoos, monarchs, flycatchers, and whistlers. These species move seasonally with food availability and breeding. All bird species recorded occur naturally in Australia and none arc restricted to the study area. Fifty-eight species are primarily northern in dis- tribution (e.g. Brown Cuckoo-dove, Spectacled Monarch); 42 are southern (e.g. Eastern Yellow Robin, White-browed Scrubwren); 21 have a regional coastal affinity (e.g. Satin Bowerbird, White-headed Pigeon); 42 are widespread across Australia (e.g. Brown Goshawk, Grey Shrike- Thrush); and 6 have an inland affinity (e.g. Emu, Galah). The ranges of 25 of these birds species terminate in or near the boundaries of the study area. Southern species whose ranges terminate in the study area include the Green Catbird, Paradise Riflebird and Eastern Whipbird (nominate race); northern species whose ranges terminate in the study area include the Faiiy Gerygone and Bar- breasted Honeycater. Of the 79 bird species recorded primarily in rainforests, most were recorded at the AMVF sites (71 species), followed by the SEVT and CNVF sites (62 and 60 species). When the mean number of species per site was compared, the SEVT sites were lower (32.2 species/site) than the other two rainforest types (AMVF 48.3, CNVF 45.3). This may be explained partly by the northern location of five of the .seven SEVT sites which places them north of the distribution of several bird species (e.g. the Green Catbird, Paradise Riflebird, and Palc-yclIow Robin). Birds which were recorded at AMVF but not SEVT sites include: Green Catbird, Paradise Riflebird, Black-breasted Button-quail, Crimson Rosella, and Yellow-tailed Black-cockatoo. Conversely, several northern species recorded at SEVT sites were not recorded at AMVF sites because their distribution terminates within the study area: Satin Flycatcher, Large-tailed Nightjar, and Oriental Cuckoo. Relative to the SEVT sites, CNVF and AMVF sites generally contained more fruit-eating birds (F. Columbidae), more insect- eating birds (F. Acanthizidae and Muscicapidae), and more birds of prey (F. Accipitridae). The rainforest species most frequently recorded were Lewiif s Honeyeater and Torresian Crow (all 18 sites), Australian Brush-turkey, Southern Boobook, Australian King-parrot, Rainbow Lorikeet, Emerald Dove, Varied Triller, Rufous Fantail, While-browed Scrub-wren (17 sites), and Australian Owlet-nightjar, Fan-lailed Cuckoo, Little Shrike-thrush, Grey Fantail, Spangled Drongo, Pied Currawong (16 sites). Mammals Twenty-three species of bats were recorded during the study (Appendix I ). Six were recorded only electronically. One was very distinct (Mor- mopterus sp.), and is probably undescribed (M.C. de Oliveira, pers. comm.). The 23 species com- prise 35% of the known Australian bat fauna. Species encountered most frequently were Mini- 218 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1 . Location of significant areas of rainforest in the study area, and of major cities, towns and roads. Numbers for study sites correspond with those in Table 1 . SURVEY OF DRY RAINFOREST VERTEBRATES 219 opteris schreibersii (9 sites), M australis (8 sites), and Chalinolobus gouldii (7 sites). Thirty-three species of terrestrial and arboreal mammal were recorded (Appendix 1), Thirty-one species were recorded in the dry seasons, 23 in the wet seasons. Nineteen native species were recorded in rainforests. Five of these species are either endemic to, or closely associated with, rainforests; Melomys cervinipes, Rattusfuscipes, Antechinus flavipes, Thylogale thetis, and Wal- labia bicolor. The geographic affinities of the native ter- restrial and arboreal mammal fauna area are primarily eastern (15 species) and northern Australian (6). The occurrence of Trichosurus caninus at Dan Dan SF was a 20km extension of the northern limit of its known range (Kroombit Tops, SEQ, Woodall, 1986). Another species, T. thetis, is also near the northern limit of its dis- tribution, and Petrogale inornata reaches its southern limit in the study area. When the nine non-rainforest species are ex- cluded, the AMVF, SEVT, and CNVF sites had by far the most diverse mammal faunas, with 24, 21, and 17 species, respectively. The remaining three forest types supported only 7 terrestrial and arboreal mammal species. The patterns become clearer when these rainforest species are divided into groups based on size and taxonomy. Bats: Twelve species recorded during the study have generally northern distributions; four are eastern regionals; three are widespread; two have mainly southern distributions; one has a mainly inland distribution; and the unidentified Mor~ mopterus sp. is probably also northern, given that the calls of southern species are generally well known. The number of species recorded was ap- proximately related to the sampling intensity and the number of sites sampled in each vegetation type: 16 species at seven SEVT sites, 12 at four AMVF sites, nine at two SEVT-AMVF sites, seven at one AMVF-ANVF site, and five each at three CNVF sites and one ANVF site. Most of the bat species recorded, especially the insectivorous bats, are habitat generalists. None are rainforest endemics, although several species roost in closed forests. Nyctimene robinsoni was captured at dusk in rainforest at Dan Dan SF, and was probably emerging to feed on Eucalyptus species which were flowering at the rainforest edge. The sheathtail bats (F. Emballonuridae) were represented by only one species, Sac- colaimus flaviventris, which was recorded at Mt Coulston and Bania. This species is regarded as rare, possibly because it flies high and fast and is difficult to capture, and because it may be restricted to roosting in hollows of aged emer- gents. Myotis adversus, recorded at Eurimbulah, is comparatively rare throughout its range be- cause it is found usually only close to water, where it skims insects from the water surface. Rodents, dasyurids and bandicoots: Ten small mammal species were recorded during the study. The most frequently encountered species was M. cervinipes (10 of 1 8 sites), which was recorded at all AMVF and CNVF sites, but at only two of the seven SEVT sites. It was typical of structurally complex rainforests where there was a ground cover of leaf litter and logs. The next most fre- quently encountered small mammal was R. fus- cipes (5 sites), which was recorded at all CNVF sites, one AMVF site, and none of the SEVT sites. It was found where shelter was available in the form of fallen trees, or a good undergrowth of shrubs and ferns. A.Jlavipes, a habitat generalist, was recorded at only three sites in three vegeta- tion types; AMVF, CNVF, and ANVF. It was found only in sites where epiphytes and logs provided shelter and moss and lichens indicated a moist environment. There was a paucity of small mammals at the SEVT sites. Po.y5w/n.?;Three species were recorded with lit- tle differenee between rainforest type in the num- ber or composition of species. The most frequently encountered species were Pseudo- cheirus peregrinus and Trichosurus caninus (12 and 1 1 sites, respectively). The former species prefers a variety of vegetation types where shrubs form dense tangled foliage, and was recorded in all CNVF sites and 3 of 4 AMVF sites. T. caninus is at the northern limit of its range in the study area and prefers tall open to closed forest. It was recorded in 2 of 3 CNVF sites and 3 of 4 AMVF sites. Both species were recorded in less than half of the seven SEVT sites, although the five north- ern and coastal SEVT sites are outside the known range of T. caninus. T. vulpecula, a habitat generalist, was recorded at 8 sites in all vegetation types. Macropods: Eight species were recorded. Those most frequently encountered in closed forests were W. bicolor and Macropus dorsalis, which were both recorded at 10 sites evenly across all vegetation types. T. thetis, which prefers rainforest and den.ser vegetation, was recorded at only one site, Dawes Range. Two rock-wallaby species were recorded in rocky habitats in closed forests during the study: Petrogale inornata, Mt Etna, and P. herberti, Coominglah and Deep Ck. 220 MEMOIRS OF THE QUEENSLAND MUSEUM Other species: Both monotreme species were recorded. Tachyglossus aculeatus was identified at Bundle Range (SEVT) from bones. At Mt Coulston an echidna was observed foraging on the edge of the AMVF, but took refuge inside it when disturbed. Ornithorhynchus anatinus was observed at Dan Dan (SEVT), in Putter Ck on the forest edge. Dingos were recorded at nine sites. DISCUSSION This study has added to knowledge of the dis- tribution of many vertebrate species, and can easily be repeated to examine future patterns of occurrence. The most significant discovery was N. spinosa, a new skink, at Nangur SF (Covace- vich et al., 1993). This is remarkable in SEQ, which is relatively well known herpetologically. The study also produced some anomalous results which probably relate more to the weather preceding each survey, and to brevity of survey, than to real faunal distribution patterns. It is generally recognised that to sample small mam- mal species, a minimum of four nights trapping is required (Denny, 1984), and to adequately sample the amphibian fauna, sampling must be carried out following significant rains, and preferably near water. Reptiles are easier to find in warmer weather. Birds also have seasonal pat- terns of distribution and abundance, with some species (e.g. cuckoos), migrating north in winter. Because of prolonged drought in much of the study area, some sites (e.g. Spring Ck, where no rain fell between both surveys) were effectively sampled in two dry seasons. The vertebrates of the rainforests of the study area are only moderately diverse. When com- pared with adjacent open woodlands, however, they support an unique and highly diverse fauna. Removal of the rainforests would result in the disappearance of many species, notably fruit- eating birds, and small mammals such as Af. cervinipes and R. fiiscipes. The relationship between rainforest type and species diversity is complicated by many factors including size, proximity of other rainforests, climate, soils, site moisture regime, and degree of disturbance. Nevertheless, some general patterns are evident from the records. The fauna of SEVT was consistently less diverse than of the other two major rainforest types, AMVF and CNVF. AMVF and CNVF sites supported on average at least 50% more bird and small mammal species than SEVT sites. AMVF also supported around twice as many amphibian species as SEVT and CNVF sites. The 19 native amphibian species recorded during the survey represent only 10% of the na- tive amphibian fauna of Australia. No permanent or temporary water was present at some sites and, with prolonged drought, this result is not surpris- ing. The Australian Museum expeditions to Granite Ck SF in 1975 recorded 5 other amphi- bians, Adelotus brevis, Limnodynastes ornatus, Litoria caerufea, L, nasuta, and L. chloris (Broadbent & Clark, 1976), all relatively wide- spread species. In rainforest at KroombitTops, 50 km from Granite Creek SF, 21 species of frog have been recorded in a variety of habitats from wet rainforest to open woodland, including Taudactylus pleione, a rainforest species endemic to the area (Czechura, 1986). Reptile diversity differed little between rain- forest types, although AMVF and SEVT sites were nearly twice as diverse as their CNVF counterparts. Two of the three CNVF sites were at Granite Ck SF which probably explains the lower reptile diversity in this rainforest type. The first survey at Granite Creek SF was conducted in very cold conditions in June, 1992; the second in wet and cool conditions in December, 1992. In the Australian Museum surveys in 1975, 1 1 addi- tional species were recorded at Granite Creek: Oedura tryoni, Anomalopus verreauxi, Egernia frerei, Lampropholis adonis, Eulatnprus quoyi. Saiga irregularis, Dendrelaphis pimctulata, Cacophis squamidosus, Pseudechis porphyri- aciis, Hemiapsis signafa, and P. salebrosus (Broadbent & Clark, 1976). A large proportion of the rainforest birds ex- pected to occur in the study area were recorded. Notable exceptions were the Double-eyed Fig Parrot, Little Bronze Cuckoo, and Rufous Bronze Cuckoo. The rainforest fauna included a large proportion of fruit-eating species which are im- portant in maintaining and regenerating rain- forests by dispersing seed in these rainforests. The bat fauna of the study area was particularly diverse, with the electronic detector increasing the number of species recorded by 35%. None of the species recorded is a rainforest endemic. However, species such as N. robinsoni roost in closed forests, and many of the fruit-eating bats are important dispersers of rainforest fruit. In contrast to the other vertebrate groups, the bat fauna was more diverse at the SEVT sites. This may be because the SEVT sites were more often adjacent to open woodland where more species ' were recorded. ! SURVEY OF DRY RAINFOREST VERTEBRATES 221 The terrestrial and arboreal mammal fauna of these rainforests is moderately diverse. Most small mammals expected were recorded. Excep- tions were Dasyurus hallucatus and D. maculata, which have undergone major range contractions; Phascogale tapoatafa, which has also undergone a range contraction; and A. stuartii, which prefers wetter, denser forests. Of special significance was the record of T, caninus at Dan Dan SF, which extends the known northern range of the species by 20 km. Acrobates pygmaeus was not recorded during this study. It was recorded at Granite Creek SF by Broadbent & Clark (1976), and has recently been sighted there (Hobcroft & Flower, pers. comm.). Another possum which may oc- cupy closed forests, Petaurus norfolcensis, was not recorded but has been recently reported for Boogooramunya SF (Hobcroft & Flower, pers. comm.). When comparing terrestrial and arboreal mam- mals by rainforest type, the occurrence of those species requiring structurally complex forests dif- fered most. M. cervinipes, R. fuscipes, and P. peregrinus were conspicuously absent from many of the SEVT sites, suggesting that those SEVT sites surveyed generally did not meet their habitat requirements. The macropod fauna of the closed forests of the study area is diverse, with five species recorded and another two, T. stig- matica and P. tridactylus, known to occur in the area. Of the five species recorded, only T. thetis could be considered a true rainforest species, and it was recorded only once, at Dawes Range, a CNVF site. T. stigmatica, which prefers wet sclerophyll forest and rainforest, was reported from Granite Ck SF by Broadbent & Clark (1976). Introduced and feral mammals did not figure highly at any of the sites. Rabbits, cattle, house mice, and feral cats were recorded at only two sites. The dry rainforests of south and mideastem Queensland are poorly conserved. In the past they have attracted little scientific interest. Those under the control of the Queensland Forest Ser- vice (QFS) can be considered to be relatively safe from major disturbance. Although cattle grazing continues in many sites, current QFS policy is that there will be no further logging. Cattle open up the forest, destroy the lower stratum, and disturb the soil and humus layer. In so doing they can destroy the habitat of small mammals, reptiles, and amphibians, while encouraging invasion by weeds, particularly Lantana camara. Fire, which is a common management tool of most graziers, continues to be a threat to many small rainforest patches by ‘eating away* the edges and, thus, progessively reducing them. We expected high diversity at larger rainforest sites, but this was not generally the case. This may be due to the presence of permanent water, and proximity to or connection with other rainforest patches. We recommend that remaining dry rainforests be conserved. They are important refuges for a wide range of species and with the discovery of N. spinosa, are known now to support at least one endemic species. ACKNOWLEDGEMENTS This study was funded by a grant from the National Rainforest Conservation Program. Generous assistance has been received from Bill McDonald and the Queensland Forest Service. Volunteers generously assisted Department of Environment and Heritage staff in the field and office. We would like to thank Mark Fletcher, Mitchell Scott, Eric Zillman, Betty Busk, Ken Tucker, Jody Sowerby, Ranghild Finkenwirth, Andrea Kintrup, Kay (Griffiths, Johannes Graf, the Inslay family and Marilyn Morgan. REFERENCES BROADBENT, J. & CLARK. S. 1976. ‘A faunal survey of east Australian rainforests: Interim Report’. (Australian Museum: Sydney). 132pp. COVACEVICH, J., COUPER, P., & JAMES, C. 1993. A new skink, Nangura spinosa gen. et sp. nov., from a dry rainforest of southern Queensland’. Memoirs of the Queensland Museum 34 (1): 159-168. CZECHURA, G.V. 1986. ‘Distant exiles: frogs and reptiles recorded from Kroombit Tops, Southeastern Queensland’. Queensland Naturalist 27(1-4): 61-67. DENNY, M. 1984. ‘Rapid surv-eys of fauna and their habitats’. Pp.376-91 In Myers, K., Margules, C.R. & Muslo, L (eds). ‘Survey methods for nature conservation Vol. 1*. (CSIRO: Canberra). FORSTER, P.I.. BOSTOCK, P.D., BIRD, L.H. & BEAN, A.R. (1991). ‘Vineforesl Plant Atlas for South-East Queensland'. (Queensland Her- barium: Brisbane). GILLISON, A.N. & BREWER, K.R.W. 1985. The use of gradient directed transects or gradsects in natural resource surveys’. Journal of Environ- mental Management 20: 103-127. INGRAM, G.J.& RAVEN, R.J. 1991 (eds). ‘An Atlas 222 MEMOIRS OF THE QUEENSLAND MUSEUM of Queensland’s frogs, reptiles, birds and mammals’. (Board of Trustees, Queensland Museum: Brisbane). 391pp. ROBINSON, R.F. 1977. ‘Management of the Good- night Scrub’. Unpubl. report to the Queensland Forest Service. STOCK, E.C., COUTTS, R.H., O’NEILL, J.P., & TRINDER,K.V. 1988. ‘Vegetation communities and heritage values of the Balaclava Is Rundle Range-Northern Narrows area, cen. Queensland. Unpubl. report for Esso Australia Ltd.’ (Institute of Applied Environmental Re- search: Griffith University). WOODALL, P.F. 1986. ‘Mammals at Kroombit Tops, Central Queensland’. Queensland Naturalist 27(1-4): 76-79. APPENDIX 1. Vertebrate species recorded in dry rainforests in south and mideastem Queensland, BANIA SF 54 Amphibians F. Myobatrachidae: Adelotus brevis; Limtxodynasies ornatus; L terraeginae; Pseiuiophryne major; F. Hylidae: Litoria rubella; F. Biifonidae: Bufo marinus. Reptiles F. Agamidae: Physignathus lesueurii; F. Gekkonidae: Diplodacrylus sieindachtieri: D. viitatus; Oedura tryoni; Phyllurus salebrosus; F. Varanidae: Varanus varius; F. Scincidae: Anomalopus verreauxi; Calyptotis lepidorostrum; C. scutirostrum; Carlia pectoralis; Ctenotus taeniolatus; Eulamprus tenuis; Lampropholis adonis; L. amicula; L couperi; L delicata. Birds F. Analidae: Chenonetta Jubata; F. Accipitridae; Accipiter fasciatus; Aquila audax; Aviceda subcrislata; F. Megapodiidae: Alectura lathami; F. Tumicidae: Turnix varia; F. Columbidae: Chalcophaps indica; Geopelia humeralis: G. placida; Leucosarcia melanoleuca; Macropygia amboinensis; Ptilinoptis magnificus; P. regina; F. Cacatuidae: Cacatua galerita; Calyptorhynchus funereus; C. magnificus; F. Loriidac: Glossopsitta pusilla; Trichoglossus chlorolepidotus; T. haemaiodus; F. PolfieVitidac: Alisterus scapularis; F. Plalycercidae: Platycercus adscitus; F. Cuculidae: Chrysococcyx lucidus; Cuculus pyrrhophanus; C. variolosus; Eudynamis scolopacea; Scythrops novaehollandiae; F. Strigidae: Ninox novaeseelan- diae; N. strenua; F. Podargidae: Podargus strigoides; F. Aegolhelidae: Aegotheles cristatus; F. Apodidae: Hirundapus caudacutiLs; F. Alcedinidae: Dacelo novaeguineae; Halcyon macleayii; F. Pillidae: Pitta versicolor, F. Campephagidae; Coracina novaehollandiae; C. papuensis; C. tenuirostris; Lalage leucomela; F. Muscicapidae: Colluricincla harmonica; C. megarhyncha; Eopsaltria australis; Falcunculus frontatus: Microeca leucophaea; Momrclui leucotis; M. melanopsis; M. trivirgatus; Myiagra cyanoleuca; Pachycephala pectoralis; P. rufiventris; Rhipidura fuliginosa; R. leucophrys: R. rufifrons; F. Orthonychidae: Psophodes olivaceus; F. Timaliidaef Pomafostomus temporalis; F. Maluridae: Malurus lamberti; M. melanocephalus; F. Acanthizidae; Acanthiza pusilla; A. regutoides; Gerygone mouki; G. olivacea; Sericornis frontalis; S. magnirostris; F. Neosittidae: Duphoenositta chrysoptera; F. Climacieridae; Climacleris leucophaea; C. picumnus; F. Meliphagidae: Entornyzon cyanotis; Uchenostomus chrysops; Manorina melanocephala: Meliphaga lewinii; Melithreptus albogularis; M. lunatus; Myzomela obscura; M. sanguinolenta; Philemon corniculatus; F. Pardalolidae: Pardalotus punctatus; F. Zostcropidae: Zoslerops lateralis; F. Ploceidae: Emblema temporalis; F. Oriolidae: Oriolus sagittalus; F. Dicruridae: Dicrurus hottentottus; F. Paradisaeidae: Ailuroedtis crassirostris: Ptiloris paradiseus; Sericulus cbrysocephalus; F. Gral- linidae: Grallina cyanoleuca: F. Craciicidae: Cracticus nigrogularis; C. torquatus; Gymnorhina tibicen; Strepera graculina; F. Corvidae; Corvus orru. Mammals: F. Phascolarctidae: Phascolarctos cinereus; F. Petauridae: Petaurus australis; F, Pseudocheiridae: Petauroides volans; F. Phalangeridae: Trichosurus caninus; T. vulpecula; F. Emballonuridae: Saccolaimus flaviventris; F. Rhinolophidae: Rhinolophus megophyllids; F. Vespertilionidae: Chalinolobus gouldii; Miniopteris australis; Nyctophilus gouldii; Scotorepens greyii; F. Muridae; Melomys cervinipes; F. Canidae: Canis familiaris dingo. BOOGOORAMUNYA SF 465 Amphibians; F. Myobatrachidae: Limnodynastes ornatus: L Crinia parasignifera; C. signifera; Pseudophryne major; F. Hylidae: Litoria caerulea; L. latopalmata; L. lesueurii; L.rubella; F. Bufonidae; Bufo marinus. Reptiles: F. Gekkonidae: Heieronotia binoei; Oedura monilis; F. Scincidae; Anomalopus verreauxi; Calyptotis scutirostrum; Carlia pectoralis; Cryptoblepharus virgatus; Ctenotus taeniolatus; Eulamprus tenuis: Lampropholis adonis: L. amicula: Lygisauru.K foliorum; F. Elapidae: Rhinoplocephalus nigrescens. Birds; F. Ardea; Ardea noxaeludlandiae; F . Accipitridae: Accipiter cirrhocephalus; A. fasciatus: Aviceda suberistata; F. Falconidac; Falco cenchroides; F. Megapodiidae: Alectura (atharni; F. Phasianidac: Coturnix australis: F. Tumicidae: Turnix varia; F. Columbidae: Chalcopimps ituiica; Geopelia huftieralis: Leucosarcia melanoleuca: Ptilinopus regina; F. Cacatuidae; Cacatua galerita; Calyptorhynchus funereus: F. Loriidae; Glossopsitta pusilla: Trichoglossus haematodus; F. Polytelitidae: Alisferus scapularis; F. Plalycercidae: Platycercus adscitus; F. Cuculidae; Centropus phasianinus; Chrysococcyx basalis; Cuculus pyrrhophanus; Eudy namis scolopacea; Scythrops novaehollandiae; F. Strigidae: Ninox rumieseelandiae; N. strenua; F. Podargidae: Podargus strigoides; F, Aegothelidae: Aegotheles cristatus; F. Alcedinidae: Dacelo novaeguineae; Halcyon sancia; F. Meropidae: Merops ornatus: F. Coraciidae: Eurystomus orientalis: F. Pitlidae: Pitta versicolor; F. Campephagidae: Coracina novaehollandiae; C. papuensis; C. tenuirostris; Lalage leucomela; F. Muscicapidae; Colluricincla harmonica; C. megarhyncha: Eopsaltria australis; Monurcha leucotis. M. melanopsis: Myiagra rubecula; Pachycephala pectoralis: P. rufiventris: Petroica rosea: Rhipidura fuliginosa: R. leucophrys; R. rufifrons; F. Orthonychidae: Psophodes olivaceus; F. Timaliidae; Pomafostomus temfwralis; F. Maluridae: Malurus melanocephalus; F. Acanihizidae: Acatuhiza pu.silla; A. reguloides; Gerygone olivacea; Sericornis frontalis; S. magnirostris; Smicrornis brevirostris; F. Neosittidae: Duphoenositta chrysoptera: F. Climacieridae: Climacteris leucophaea; F. Meliphagidae: Lichenostomus chrysops: Manorina melanocephala: Meliphaga leM'inii: Melithreptus albogularis: Myzomela sanguinolenta; F. Pardalolidae: Pardalotus punciatits; P. striafus; F. Zostcropidae: Zoslerops lateralis; F. Ploceidae: Emblema temporalis: Poephilabichenovii: Emblema temporalis; F. Oriolidae: Oriolus sagitlatus; Splwcotheres viridis; F. Dicruriikie: Dicrurus hottentottus; F. Corcoracidac: Corcorax melanorhamphos; F. Craciicidae: C torqimtus; Gymnorhina tibicen; Strefyera graculina; F. Corvidae: Con’us orru. Mammals: F. Dasyuridae: Planigale maculata; F. Peramelidae: Perameles nasuta: F. Phascolarctidae: Phascolarctos SURVEY OF DRY RAINFOREST VERTEBRATES 223 cinereus; F. Pseudocheiridae: Pseudocheirus peregrinus; F. Phalangeridae: Trichosurus caninus; T vulpecula', F. Macro- podid^. Macropus dorsalis; M. giganteus; Wallabia bicolor; F. Molossidae: Mormopterus beccarii; Nyctinomus australis; F. Rhinolophidae: Rhiuolopus megaphyUus; F. Vespertilionidae: Chalinolobus gouldii; C. nigrogriseus; Eptesicus pumilis; Miniopterus australis; M. schreibersii; Nyctophilus geoffroyi Scotorepens greyii; F. Muridae: Melomys butoni; M. cervinipes; F. Canidae: Canis familiaris dingo; F. Leporidae: Oryctolagus cuniculus. BOYNE LOGGING AREA SF 391 Amphibians: F. Hylidae: Litoria lesueurii; F. Bufonidae: Bufo marinus. Reptiles: F. Gekkonidae: Phyllurus caudiannulatus; F. Varanidae: Varanus varius: F. Scincidae: Calyptotis scutirostrwn; Hemisphaeriodon gerrardii; Lampropholis adonis; Ophioscincus ophioscincus. Birds: F. Accipitridae: AccipiternovaehoUandiae;Aquilaaudax; Aviceda subcristata; Lophoictinia isura; F. Megapodiidae: Alectura lathami; F. Columbidae: Chalcophaps indica; Columlxi leucomela; Leucosarcia melanoleuca; Lopholaimus an- tarcticus; Macropygia amboinensis; Ptilinopus magniftcus; P. regina: P. superbus; F. Cacaiuidae: Calyptorhynchusfunereus; C. magniftcus; R Loriidae: Trichoghssus haematodus; F. Polytelitickie: Alisterus scapularis: F. Cuculidae: Chrysococcyx tucidus; Cucutus pyrrhophanus; F. Slrigidae: Ninox novaeseelandiae; F. Tytonidae: Tyto tenehricosa; F. Aegolhelidae: Aegotheles cristatus; F. Apodidae: Apus pucificus; flirundapus caudacutus; F. Alcedinidae: Ceyx azurea; Dacelo novaeguineae; Halcyon sancta; F‘. Pitlidac: Pitta versicolor; \\ Campcphagidae: Coracina lineata; C. tenuirostris; Lalage leucomela; F. Muscicapidae: Collurichicla harmonica; C. megarhyncha; Eopsaltria aiatralis: Monarcha melanopsis; M. trivirgatus; Pachycephala pectoralis; Petroica rosea; Rhipidura fuliginosa; R. rufifrons; F. Orthonychidae: Psophodes olivaceus; F. Acanthizidae: Acanthiza pusilla; Gerygone mouki; Sericornis frontalis; S. magnirosfris; F. Climacteridae: Climacteris leucophaea; F. Meliphagidae: Acanthorhynchus tenuirostris; Meliphaga lewinii; Myzotnela obscura; M. san- guinolenta; F. Zosteropidae: Zosterops lateralis: F. Ploceidae: Emblema temporalis; F. Oriolidae: Sphecotheres viridis; F. Paradisaeidae: Ailuroedus crassirostris; Ptiloris paradiseus; Sericulus chrysocephalus; F. Artamidae: Artamus cinereus; F. Cracticidae: Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Dasyuridae: Planigale maculata; F. Pseudocheiridae: Pseudocheirus peregrinus; F. Phalangeridae: Trichosurus caninus; F. Pteropodidae: Pteropus poliocephalus; Syconycteris australis; R Muridae: Melomys cervinipes; Rattus fuscipes; F. Bovidae: Bos taurus. DAWES RANGE SF 391 Amphibians: F. Myobatrachidae: Litoria lesueuri; R Bufonidae: Bufo marinus. Reptiles: F. Gekkonidae: Phyllurus caudiannulatus; F. Scincidae: Calyptotis scutirostrum; Eulamprus tenuis; Lampropholis adonis; Ophioscincus ophioscincus; R Boidae: Morelia spilota; Rhinoplocephalus nigrescens. Birds: F. Accipitridae: Accipiter novaehotlandiae; F*. Megapodiidae: Alectura lathami; F. Columbidae: Chalcophaps indica; Columba leucomela; Leucosarcia melanoleuca; Ixtpholaimus antarcticus; Macropygia amboinensis; Ocyphaps lophotes; Ptilinopus magniftcus: P, regina; R Cacaiuidae: Calypiorhynchus magniftcus; F, Cuculidae: Cuculus pyrrhophanus; F. Loriidae: Trichoghssus haematodus; F. Polytelitidae: Alisterus scapularis: F. Cuculidae: Cuculus pyrrhophanus; R Strigidae: Ninox novaeseelandiae; F. Tytonidae: Tyto tenebricosa; F. Aegolhelidae: Aegotheles cristatus; R Apodidae: Apus pacificus; F. Alcedinidae: Dacelo novaeguineae; R Pillidae: Pitta versicolor; F. Campcphagidae: Coracina novaehollandiae; Lalage leucomela; F. Muscicapidae: Colluricincla harmonica: C. megarhyncha: Eopsaltria australis: Monarcha leucotis; M. melanopsis; Pachycephala pectoralis: Petroica rosea; Rhipidura fuliginosa; R. rufifrons; F. Orthonychidae: Psophodes olivaceus; F. Acanthizidae: Acanthiza nana; A. pusilla; Gerygone mouki; Sericornis frontalis; S. magnirostris; F. Climac- teridae: Climacteris leucophaea; R Meliphagidae: Lichenostomus chrysops;. Lichmera indistinefa; Meliphaga lewinii; Myzomela sanguinolenta; F. Zosteropidae: Zosterops lateralis; F. Ploceidae: Emblema temporalis; F. Oriolidae: Sphecotheres viridis; F. Paradisaeidae: Ailuroedus crassirostris; Ptiloris paradiseus: Sericulus chrysocephalus; F. Grallinidae: Grallina cyanoleuca; F. Cracticidae: Cracticus nigrogularis; Gymnorhina tibicen; Strepera graculina: R Corvidae: Corvus orru. Mammals: R Dasyuridae: Antechinus favipes; R Peramelidae: Perameles nasuta; F. Pseudocheiridae: Petauroides volans; Pseudocheirus peregrinus: F. Phalangeridae: Trichosurus caninus: F. Macropodidae: Macropus dorsalis; Thylogale ihetis; Wallabia bicolor; F. Muridae: Melomys cervinipes: Rattus fuscipes. COOMINGLAHSF28 Amphibians: F. Bufonidae: Bufo marinus. Reptiles: F. Gekkonidae: Diplodactylus viltatus; Heteronotia binoei; Oeditra tryoni; Underwoodisaurus rmlii; F. Agamidae: Pogona barbata; F. Varanidae: Varanus varius: F. Scincidae: Anomalopus verreuuxi; Carlia pectoralis: C. schmeltzii; Cryptoblepharus virgatus; Ctenotus taeniolaius; Eulamprus quoyii; E. tenuis; Lampropholis amicula; F. Elapidae: Rhinoplocephalus nigrescens. Birds: F. Accipitridae: Aquila audax; F. Megapodiidae: Alectura lathami: R Columbidae: Chalcophaps itulica; Geopelia humeralis; G. placida; Leucosarcia melanoleuca; Ocyphaps lophotes: F. Cacaluidite: Cacatua galeriia; C. roseicapilla; Calyptorhynchus magniftcus; V. Loriidae: Glossopsitta pusilla; Trichoglossus haematodus; F. Polytelitidae: Alisterus scapularis; F. Plalycercidac: Platycercus adscitus; F. Cuculidae: Chrysococcyx tucidus; Cuculus pyrrhophanus; F. Strigidae. Ninox novaeseelandiae; R Pcxlargidae: Podargus strigoides; F. Aegolhelidae: Aegotheles cristatus: F. Alcedinidae; Dacelo novaeguineae; R Meropidae: Merops ornatus; F. Pillidae: Pitta versicolor: R Campcphagidae: Coracina novaehollandiae; C papuensis; Lalage leucomela; F. Muscicapidae: Colluricincla harmonica; Eopsaltria australis. Pachycephala pectoralis, P. rufiventris; Rhipidura fuliginosa; R. leucophrys; R. rufifrons; F. Orthonychidae: Psophodes olivaceus: F, Timaliidae. Pomatostomus temporalis; R Maluridae; Malurus lamberti; R Acanthizidae: Acanthiza pusilla; A. reguloides; Gerygorw mouki; G. olivacea; Sericornis frontalis: S. magnirostris; S. saggitatus; F. Ncosiltidae: Daphoenositta chrysoptera, F. Climacteridae: Climacteris leucophaea; R Meliphagidae: Lichenostomus chrysops; Manorina melanocephala; Meliphaga lewinii; Melithreptus lunatus; Myzotnela sanguinolenta; Philemon citreogularH; R Pardalolidae; Panhlotus pu/ictulatM; P. striatus; R Zosteropidae: Zosterops lateralis; F. Ploceidae: Emblema temporalis; F. Oriolidae. Sphecotheres viridis, F. Dicruridae: Dicrurus hotteniotm: R Grallinidae: GraUina cyanoleuca; H. Cracticidae: Cracticus mgrogularts; Strepera graculina; F. Corvidae: Corvus orru. , i- r. j u j Mammals: F. Dasyuridae: Planigale maculata; F. Phascolarctidae: Phascolarctos cinereus; R Pseudochemdae: 224 MEMOIRS OF THE QUEENSLAND MUSEUM Petauroides volans: Pseudocheirus peregrinus; F. Macropodidae: Macropus dorsalis; M. giganteus; Petrogale herberti; Wallabia bicolor; F. Molossidae: Mormopterus loriae; F. Vespertilionidae: Chalinolobus gouldii; C. nigrogriseus; Miniopteris schreibersii; Scoleanax rueppellii; Scolorepens greyii; F. Muridae: Melomys cervinipes; F. Canidae: Canis familiaris dingo: COONGARA ROCK SF 1344 Amphibians: F. Myobatrachidae: Adelotus brevis; Mixophyes fasciolatus; F. Hylidae: Litoria caerulea; L. gracilenta. Reptiles: F. Scincidae: Calyptotis scutirostrum; Carlia pectoralis; Cryptoblepharus virgatus; Ctenotus robustus: Eulamprus quoyii; E. tenuis: Lampropholis adonis: L amicula; F. CoJubridae: Boiga irregularis: F. Elapidae: Pseudechis porphyriacus. Birds: F. Ardeidae: Nycticorax caledonicws: F. \cc\p\\nd:^it:Accipiternovaehollandiae: Aquilaaudax: Avicedasubcristata: F. Megapodiidae: Alectura laihami: F. Phasianidac; Cotumix australis: F. Tiimicidae: Tumix varia: F. RalHdae: Gallinula olivacea: F. Columbidae: Chalcophaps indica: Geopelia humeralis: G, placida: Leucosarcia melanoleuca: Macropygia amboinensis: Ptilinopus regina; F. Cacatuidae: Cacatua galerita; F. Loriidae: Glossopsitta pusilla: Trichoglossus haematodus: F. Polyielitidae: Alisterus scapularis; F. Platycercidae: Platycercus adsciius: F. Cuculidae: Centropus phasianinus: Chrysococ- cyxlucidus: Cuculus pyrrhophanus: Eudyfiamis scolopacea: F. Strigidae: Ninox novaeseetandiae; F. AegoiheWdac: Aegotheles cristatus: F. Alcedinidae: Dacelo novaeguineae; F, Meropidae: Merops ornalus: F Coraciidae: FMrystomus orientalis; F. Pittidae: Pitta versicolor: F. Campephagidae: Lalage teucomela: F. Muscicapidae: Colluricincla harmonica: C. megarhyncha: Eopsaltria australis: Monarcha leucotis: M. melanopsis: M. trivirgatus: Pachycephala rufiventris: Petroica rosea; Rhipidura fuliginosa: F. Onhonychidae: Psophodes olivaceus; F. Maluridae: Malurus melanocephalus; F. Acanthizidae: Acanthiza pusilla: Sericomis frontalis: S. magnirostris; F. Mehphagidae: Lichmera indistincta: Meliphaga lewinii; Myzomela san- guinolenta: Philemon comiculatus: F. Pardalolidae: Pardalotus puncfulatus: P. striata: F. Ploceidae: Emblema temporalis: Poephila bichenovii; F. Dicruridae: Dicrunts hoftentoffus: F. Cracticidae: Cracticus nigrogularis: Gymnorhina tibicen; Strepera graculina: F. Corvidae: Corvus orru. Mammals: F. Pseudocbeiridae: Pseudocheirus peregrinus: F. Phaiangeridae: Trichosurus caninus: F. Molossidae: Mor- mopterus loriae: F, Rhinolophidae: Rhinolophus megaphyllus; F. Vespertilionidae: Chalinolobus gouldii: Miniopteris austrttlis: M. schreibersii: Scoteanax rueppellii: DAN DAN SCRUB SF 53 Amphibians: F. Myobatrachidae: Limnodynastes ornati4S’, Pseudophryne major; F. Hylidae; Litoria caerulea; L gracilenta; L lesueurii; L. rubella; F. Bufonidae: Bufo marinus. Reptiles: F. Agamidae: Diporiphora australis: F. Scincidae: Anomalopus verreauxi; Carlia pectoralis: C. schmeltzH: C. vivax: Cryptoblepharus virgatus; Ctenotus taeniolatus; F. Boidae; Morelia spilota; F. Colubridae: Dendrelaphis punctulata. Birds: F. Ardeidae: Ardea pacifica; Nycticorax caledonicus: F. Accipitridae: Accipiter cirrhocephalus; Aquila audax; F. Falconidae: Falco berigora; F. Megapodiidae: Alectura lathami: F. Phasianidae: Coturnix australis; F. Columbidae: Chal- cophaps indica: Geopelia humeralis: G. placida: Leucosarcia melanoleuca: Lopholaimus aniarcticus; Macropygia amboinen- sis: Ptilitiopus magnificus: P. regina: F. Cacaluidite: Cacatua galerita; Calyptorhynchus magnificus; H. Loriidae: Trichoglossus chlorolepidotus; T. haematodus; F. Polytelitidae: Alisterus scapularis; F. Platycercidae: Platycercus adscitus; F. Cuculidae: Centropus phauaninus; Chrysococcyx lucidus: Cuculus pyrrhophanus: Scythrops novaehollundiae; F. Strigidae: Ninox novaeseelandiae: F. Podargidae: Podargus strigoides; F. Aegothelidae: Aegotheles cristatus: F. Capriimdgidac: Caprimulgus macrurus; F. Apodidae: Apuspaciftcus; Hirundapus caudacutus: F. Alcedinidae: Ceyxazurea: Dacelo novaeguineae; Halcyon Sanaa; F. Meropidae: Merops ornatus: F. Coraciidae: Eurystomus orientalis: F. Campephagidae: Coracina tenuirostris: Lalage leucomela; F. Muscicapidae: Colluricincla harmonica; C. megarhyncha: Eopsaltria australis: Monarcha leucotis: M. melanopsis: M. trivirgatus: Myiagra rubecula: Pachycephala peaoralis: P. nifiventris; Rhipidura fuliginosa: R. rufifrons; F. Orthonychidae: Psophodes olivaceus: F. Maluridae: Malurus lamberti; F. Acanthizidae: Acon/A/sa nana; A. pusilla; Sericomis frontalis; S. magnirostris; F. Meliphagidae: Manoritxa melanocephala: Meliphaga le\\'inii; Melithreptus albogularis; Myzomela obscura: M. sanguinolenta; Philemon citreogularis: P. comiculatus; F. Dicaeidae: Dicaeum hirundinaceum; F. Pardalolidae: Pardalotus striatus: F. Zosteropidae: Zosterops lateralis; F. Ploceidae: Emblema temporalis: F. Onolidae: Oriolus sagittatus; Sphecotheres viridis: F. Dicruridae: Dicrurus hottentottus; F. Paradisaeidae: Ptilonorhynchus violaceus; Sericulus chrysocephalus; F. Corcoracidae: Corcorax melanorhatnpkos: F. Cracticidae: Cracticus nigrogularis: Gymnorhina tibicen; Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Ornithorynchidae: Ornithorhynchus anafinus: F. Phascolarctidae: Phascolarctos cinereus; F. Pseudo- cheiridac: Petauroides volans: F. Phaiangeridae: Trichosurus canirms; T. vulpecula; F. Macropodidae: Wallabia bicolor; F. Pteropodidae: Nyctimene robinsoni; Pteropus alecto; F. Vespertilionidae: Chalinolobus gouldii; Miniopteris schreibersii: Scoteanax ruef^ellii; F. Muridae; Hydromys chrysogaster; Rattusfuscipes; F. Felidae: Felis catus. DEEP CREEK TR 581 Amphibians: F. Myobatrachidae: Limnodynastes ornatus; L. peronii; Pseudophryne major; F. Hylidae: Litoria caerulea; L latopalmata; F. Bufonidae: Bufo marinus. Reptiles: F. Gekkonidae: Gehyra dubia; Heteronotia binoei; Oedura tryoni; F. Scincidae: Calyptotis scutirostrum: Carlia pectoralis: C. schmeltzii; C. vivax; Cryptoblepharus virgatus; Eulamprus tenuis: Lampropholis adonis: L. amicula; Ophios- cincus ophioscincus; F. Boidae: Morelia spilota; F. El^idae: Pseudechis porphyriacus; Pseudonaja textilis. Birds: F. Ardeidae: Ardea novaehotlandiae; F. Analidae: Anas stqjerciliosa; F. Accipitridae: Aquila audax; F. Falconi^e: Falco berigora; F. Megapodiidae; Alectura lathami; F. Turnicidae; Turnix varia;. F. Columbidae: Chalcophaps indica; Geopelia humeralis; Lopholaimus antarcticus: Macropygia amhoinetisis: F. Cacatuidae: Calyptorhynchus magnificus; Cacatua roseicapilla; F. Loriidae: Trichoglossus haematodus; F. Polytelitidae; A /ijr/er«5 scapularis; F. Platycercidae: Platycer- cus adsciius: F. Cuculidae: Centropus phasianinus; Chrysococcyx lucidus; Cuculus pyrrhophanus: Eudynamis scolopacea; Scythrops novaehollandiae; R Strigidae: Ninox novaeseelandiae; F. Aegothelidae: Aegotheles cristatus; F. Alcedinidae: Dacelo novaeguineae; F. Meropidae: Merops ornatus; F. Pittidae: Pitta versicolor; F. Campephagidae: Coracina novaehol- landiae: C. tenuirostris; Lalage leucomela; F. Muscicapidae: Colluricincla harmonica; C. megarhyncha; Eopsaltria australis; Monarcha leucotis; M. melanopsis; M. trivirgatus; Pachycephala pectoralis; P. ruftvenlris; Rhipidura fuliginosa: R. rufifrons; R Orthonychidae: Psophodes olivaceus; F. Timaliidae: Pomatostomus temporalis; F. Maluridae: Malurus melanocephalus; F. Acanthizidae: Acanthiza pusilla; A. reguloides; Gerygone olivacea; G. palpebrosa; Sericomis frontalis; S. magnirostris: SURVEY OF DRY RAINFOREST VERTEBRATES 225 F. Neosittidae; Daphoenositta chrysoptera; F. Climacleridae; Climacteris leucophaea; F. Meliphagidae: Manorina melanocephala; Meliphaga lewinii; Myzomela sanguinolenta; Philemon corniculaius; F. Dicaeidae: Dicaeum hirundinaceum; F. Pardalolidae: Pardaloms siriafus; F. Ploceidae: Emblema temporalis; F. Oriolidae: Sphecotheres viridis; F. Dicruridae; Dicrurus holtentottus^ F. Cracticidae.' Cracticus nigrogularis; C. torguatus', Gymnorhina tibicen; Strepera graculina", F. Corvidae: Corvus orru. Mammals: F. Pseudocheiridae: Pseudocheirus peregrinus; F. PhaJangeridae: Trichosurus canini 4 s; T. vulpecula; F. Macropodidae: Macropus dorsalis; M. parryi; Petrogale herbeti; F. Pteropodidae: Pteropus scapulatus; F. Rhinolophidae: Rhinolophus megaphyUus; F. Vespertilionidae: Chalinolobus nigrogriseus; Miniopterus australis; F. Canidae: Canis familiaris dingo; EURIMBULA NP Amphibians: F. Myobalrachidae: Adelotus brevis; Limnodynastes terraereginae; F. Hylidae: Litoria caerulea; L. latopal- mala; L rubella; F. Bufonidae: Bufo marinas. Reptiles: F. Gekkonidae: Oedura tryoni; F. Varanidae: Varanus varius; F. Scincidae: Curlia pectoralis; Cryptoblepharus virgatus; Eulamprus tenuis; F. Elapidae: Cacophis harrietae; Rhinoplocephalus nigrescens. Birds: F. Anatidae: Anas supercHiosa; F. Accipitridae: Accipiter novaehoUandiae; Aguila audax; Aviceda subcristata; Haliaeetus leucogaster; Haliastur indus; F. Megapodiidac: Alectura lathami; F. Laridae: Larus novaehoUandiae; F. Colum- bidae: Chalcophaps indica; Geopelia humeratis; G. placida; Leucosarcia melanoleuca; Macropygia amboinensis; Ptilinopus regina; F. Cacatuidae: Cacatuagalerita; CalyptorhytKhus magnificus; F. I^riidae: Trichoglossus haematodus; F. Polytelitidae: Alisterus scapuiaris; F. Cuculidae: Chrysococcyx ltdcidus; Cuculus pyrrhophanus; F. Strigidae: Ninox novaeseelandiae; N. strenua; F. Podargiitie: Podargus strigoides; F. Aegochelidae: Aegotheles cristatus; F. Alcedinidac: Dacelo novaeguineae; Halcyon macleayii; F. Meropidae: Merops ornattis; F. Pittidae: Pitta versicolor; F. Hinindinidae; Hirwido neoxena; F. Campephagidae: Coracina novaehollandiae; C. fenuirostris; Lalage leucomela; F. Muscicapidae: Colturicincla megarhyncha; Eopsaltria australis; Monarcha leucolis; M. melanopsis; M. trivirgatus; Myiagra cyanoleitca; M. rubecuta; Pachycephala pectoralis; P. rufiventris; Rhipidura fuHginosa; R. rufifrons; F. Maluridae: Malurus melanocephalits; Acanlhi/.idae: Gerygone palpebrosa; Sericomis frontalis; S. magnirostris; F. Meliphagidae: Acanfhorhyru:hus tenuirosiris; Lichnera indistincta; Manorina melanocephala; Meliphaga lewinii; Myzomela obscura; M. simguinolenta; Philemon citreogularis; F. Dicaeidae: Dicaeum hirundinaceum; F. Zosteropidac: Zosterops lateralis; F. Oriolidae: Oriolus sagittatus; Sphecotheres viridis; F. Dicruridae: Dicrurus hottentottus; F. Cracticidae: Cracticus nigrogularis; F. Corvidae: Cor\>us orru. Mammals: F. Peramelidae: Perameles tuisuta; F. Petauridae: Petaurus breviceps; F. Pseudocheiridae: Pseudocheirus peregrinus; F. Phalangeridae: Trichosurus vulpecula; F. Macropodidae: Macropus dorsalis; M. giganteus; F. Pteropidae: Nyctimene robinsoni; Pteropus poliocephalus; Syconycteris australis; F. Rhinolophidae: Rhinolopus megaphyUus; F. Vesper- tilionidae: Myotis adversus; F. Muridae: Melomys cervinipes; Rattus fuscipes; F. Canidae: Canis familiaris dingo; F. Felidae: Felis catus. GOODNIGHT SCRUB SF 169 Amphibians: F. Myobatrachidae: Adelotus brevis; Mixophyes fasciolatus; F. Hylidae: Litoria caerulea; L. latopalmata; L. lesueurii; F. Bufonidae: Bufo marinus. Reptiles: F. Gekkonidae: Heteronotia binoei; Oedura tryoni; F. Agamidae: Physignathus lesueurii; Pogona barbata; F. Pygopodidae: Lialis burtonis; F. Varanidae: Varanus varius; F. Scincidae: Calyptotis scufirostrum; Carlia pectoralis; Ctenotus robustus; C. taeniolatus; Eulatnprus guoyii; LamprophoUs arnicula; F. Elapidae: Rhinoplocephalus nigrescens. Birds: F. Phalacrocoracidae: Phalacrocorax varius; F. Ardeidae Ardea novdehoUatidiae; F. Anatidae: Anas sigrerciliosa; I F.Accipilridae: Accipiter cirrhocephalus; Aviceda subcristata; F. Falconidae: Falco berigora; F- Megapodiidac: Alectura lathami; F. Turnicidae: Turnix varia; F. Charadriidac: Vanellus miles; F. Coluinbidae; Chalcophaps indica; Geopelia humeralis; Geopelia placida; Leucosarcia melanoleuca; Macropygia amboinensis; Ocyphaps loplwtes; F. Cacatuidae: Cacatua galerita; Calyptorhytichus magnificus; F. Loriidae: Trichoglossus chlorolepidotus; T. haematodics; F. Polytelitidae: Alisterus scapuiaris; F. Plalycercidae: Platycercus adscitus; F. Cuculidae; Centropus phasianinus; Cuculus pyrrhophanus; Eudynamis scolopacea; Scythrops novaehoUandiae; F. Strigidae; Ninox novaeseelandiae; N. strenua; F. Caprimulgidae: Caprimulgus gultatus; C. rnystacalis; F. Alcedinidac; Ceyx azurea; Dacelo novaeguineae; Halcyon macleayii; F. Coraciidae: Eurystomus orientalis; F. Pittidae: Pitta versicolor; F. Hirundinidae: Hirundo neoxena; F, Campephagidae: Coracina Uneata; C. novaehoUandiae; C. fenuirostris; Lalage leucomela; F. Muscicapidae; CoUuricincla harmonica; C. megarhyncha; Eop- saltria australis; Monarcha leucotis; M. trivirgatus; Myiagra rubecuta; Pachycephala pectoralis; Petroica rosea; Rhipidura fuliginosa; R. rufifrons: F. Orthonycliidac: Psophodes olivaceus; F. Acanihizidae: Acatithiza pi^illa; Gerygone mouki; G. olivacea; Sericomis frontalis; S. magnirostris; F. Climacleridae; Climacteris leucophaea; F. Meliphagidae; Manorina melanocephala; Meliphaga lewinii; Melilhreptus albctgularis; Myzomela obscura; M. sanguinolenta; Philemon corniculatus; F. Dicaeidae: Dicaeum hirundinaceum; F. Pardalolidae: PardalotiLs striafus; F. Zosteropidac: Zosterops lateralis; F. Ploceidae: Emblema temporalis; F. Oriolidae; Oriolus sagittatus; Sphecotheres viridis; F. Dicruridae: Dicruru.s hottentottus; F. Paradisaeidae: Sericulus chrysocephaJus; F. Corcoracidae: Corcorax melanorhttmphos; F. Cracticidae: Cracticus nigrogularis; C. torguatus; Gymtwrhina tibicen; Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Tachyglossidae: Tachyglossus aculeatus; F. Phascolarctidac: Phascolarctos cinereus, F. Pseudocheiridae. Pseudocheirus peregrinus; F. Phalangeridae: Trichosurus caninus; F. Macropodidae: Macropus dorsalis; Wallabia bicolor, F. Pteropodidae: Pteropus scapulatus; F. Vespertilionidae: Miniopterus australis; F. Muridae: Melomys cervinipes; F. Canidae: Canis familiaris dingo; F. Equidae: Eguus cabatius. MX COULSTON SF 471 , . , , tr Amphibians: F. Myobalracliidae; Limnodynastes perom; F. Hylidae; Litoria fallax; L lesueurii; u nasuta, L rubella, F. Bufonidae: Rwfo ^ r- o • j Reptiles: F. Gekkonidae: Gehyra dubia; Oedura monilis; O. tryoni; F. Agamidae: Diporiphora bilineata; F. Scincidae: Anomalopus verreauxi; Calyptotis lepidorostrum; Carlia pectoralis; C. schneltzU; Ctenotus taeniolatus; Eulamprus E. tenuis; Glaphyromorphus punctulatus; Hemisphaeriodon gerrardii; LamprophoUs adonis; F. Colubridae: Dendrelaphis punctulata; F. Elapidae: Pseudechis porphyriacus. 226 MEMOIRS OF THE QUEENSLAND MUSEUM Birds: F. Ardeidae: Ardea novaehollandiae; F. Plataleidae: Threskiomis spinicollis; F. Anatidae: Anas superciliosa; F. Accipilridae: Accipiter fasciatus\ Haliaeetus leucogaster, F. Falconidae: Falco berigora; F. cenchroides; F. Megapodiidae: Alectura lathami; F. Phasianidac: Coturnix australis; F. RalHdae: Gallinula tenebrosa-, F. Gruidae: Grus rubicundus; F. Charadnidae: VaneUusmiles\ F, Columbidae: Chalcophaps indica; Geopelia humeralis; Lopholaimus antarcticus; Macropygia amboinensis; Ocyphaps lophotes; Ptilinopus regina; F. Cacaluidae; Calyptorhynchus magnijtcus; F. Loriidae: Trichoglossus haematodus\ F. Polytelitidae: Alisterus scapularts; F. Platycercidae: Platycercus adscitus; F. CucuHdae: Centropus phasianinuSy Chrysococcyx basalts; Eudynamis scolopacea; F. Strigidae: hUnox novaeseelandiae; F. CaprimuJgidae: Caprimuigus guttatus; C. mystcwalis; F. Alcedinidae: Dacelo leachii; D. novaeguineae; Halcyon macleayii; H. sancta; F. Meropidae: Merops ornatus; F. Pillidae: Pitta versicolor; F. Motacillidae: Anthus novaeseelandiae; F. Molacillidae: Anthus novaeseelandiae: F. Campephagidae: Coracina lineata; Lalage leucomela; F. Muscicapidae: Colluricincla megarhyncha; Eopsaltria australis; Monarcha leucotis; M. melanopsis; M. trivirgatus; Pachycephala pectoralis; Rhipidura fuliginosa; R. leucophrys; R. rufifrons; F. Orthonychidae: Psophodes olivaceus; F. Maluridae: Malurus melanocephalus; F. Acanthizidae: Acantfuza nana;A. pusilla; Gerygone palpebrosa; Sericornis frontalis; S. magnirostris; F. Entornyzon cyanotis; Lichinera indistincta; Manorina melanocephala; Meliphaga lewinii; Myzomela obscura; M. sa/tguinolenta; F. Dicaeidae: Dicaeum hirundinaceum; F. Zostcropidac: Zosterops lateralis; F. Ploceidae: Emblema temporalis; Poeplula bichenovii; F. Oriolidae: Sphecotheres viridis; F. Dicruridae; Dicrurus hottentottus; F. Paradisaeidae: Sericulus chrysocephalus; F. Gral- linidae: Grallina cyanoleuca; F. Cracticidae: Cracticus nigrogularis; Gymnorhina tibicen; Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Peramelidae: Isoodon macrourus; F. Macropodidae: Wallabia bicolor; F. Vespertilionidae: Miniopterus schreibersii; F. Muridae; Melomys cerx inipes. MT ETNA NP Amphibians: F. Myobatrachidae: Limnodynastes salmini; F. Hylidae: Litoria caerulea; F. Bufonidae: Bufo marinus. Repliles: F. Gekkonidae: Gehyra dubia; Heteronotia binoei; F. Scincidae: Carlia pectoralis; Ctenotus taeniolatus; Eulamprus tenuis; F. Boidae: Morelia spilota. Birds: F. Accipitridae: Accipiter fasciatus; Aviceda subcristata; F. Falconidae: Falco berigora; F. cenchroides; F. Megapodiidae: lathami; F. Phasianidac: Coturnix australis; F. Columbidae: Chalcophaps indica; Geopelia humeralis; Leucosarcia melanoleuca; Ptilinopw: regina; F. Cacaluidae: Calyptorhynchus magnificus; F. Loriidae: Trichoglossus haematodus; F. Polytelitidae: Alisterus scapularis; F. Cuculidae: Centropus phasianinus; Cuculus pyrrhophanus; F. Strigidae: Ninox novaeseelandiae; F. Coraciidae: Eurystotnus orientalis; F. Alaudidae: Mirafra javanica; F. Motacillidae: Anthus novaeseelandiae; F. Campephagidae: Lalage leucomela; F. Muscicapidae: Colluricincla megarhyncha; Eopsaltria australis; Monarcha leucotis; Rhipidura rufifrons; F. Maluridae: Malurus melanocephalus; F. Acanthizidae: Sericornis frontalis; F. Meliphagidae: Meliphaga lewinii; F. Dicaeidae: Dicaeum hirundinaceum; F. Ploceidae: Lonchura castaneothorax; F. Dicruridae: Dicrurus hottentottus; F. Cracticidae: Cracticus nigrogularis; F. Corvidae: Corvus orru. Mammals: F. Macropodidae: Petrogale inornata; Wallabia bicolor; F. Megadermatidae: Macroderma gigas; F. Rhinolophidae: Rhinoiophus megaphyUus. MT LARCOM Amphibians: F. Myobatrachidae: Pseudophrytie coriacea; F. Bufonidae: Bufo marinus. Reptiles: F. Gekkonidae: Oedura monilis; O. tryoni; F. Scincidae: Carlia schmeltzii; Crypioblepharus virgatus; Lampropholis adonis; Lygisaurus foliorum. Birds: F. Ardeidae: Ardea novaehollandiae; Egretta alba; F. Accipilridae: Aquila audax; F. Megapodiidae: Alectura lathami; F. Columbidae; Chalcophaps indica; Geopelia humeralis; Macropygia amboinensis; Ptilinopus regina; F. Cacaluidae: Calyptorhynchus magnificus; F. Cuculidae: Centropus phasianinus; Cuculus pyrrhophanus; Scythrops novaehol- landiae; F. Aegoll«lidae: Aegotheles cristatus; F. Alcedinidae: Dacelo novaeguineae; Halcyon macleayii; F. Muscicapidae: Monarcha trivirgatus; Pachycephala rufiventris; Rhipidura fuliginosa; R. leucophrys; R. rufifrons; F. Acanthizidae: Acanthiza nana; F. Meliphagidae: Meliphaga lewinii; Melithreptus allxygularis; F. Pardalolidae: Pardalotus striatus; F. Oriolidae: Sphecotheres viridis; F. Dicruridae: Dicrurus hottentottus; F. Cracticidae: Cracticus nigrogularis; Gymnorhina tibicen; Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Phalangeridae: Trichosurus vulpecula; F. Macropodidae; Wallabia bicolor; F. Pteropodidae: Pteropus alecto; F. Vespertilionidae: Chalinolobus gouldii; C. nigrogriseus; Eptesicus purnilis; Miniopteris australis; M. schreibersii. NANGUR SF 74 Amphibians: F. Myobatrachidae: Limnodyiuistes ornatus; F. Hylidae: Litoria caerulea; L. latopalmata; F. Bufonidae: Bufo marinus. Repliles: F, Gekkonidae: Oedura tryoni; F. Agamidae: Gemrnatophora nobbi: F. Scincidae: Anomalopus verreaicxi; Calyptotis scutirostrum; Carlia pectoralis; Cryptoblepharus virgatus; Ctenotus taeniolatus; Egernia cunninghami; Lygisaurus foliorum; Menetia greyii; Nangura spinosa; Saiphos equalis; F. Varanidae: Varanus varius. Birds: F. Accip'ttndae.Accipiterfasciatus. Aquilaaudax; F. Falconidae; Falco berigora; F. Megapodi idae: Alectura lathami; F. Phasianidac: Coturnix australis: F. Columbidae: Chalcophaps indica; Geopelia humeralis; Leucosarcia melanoleuca; Macropygia amboinensis; F. Cacaluidae: Cacatua roseicapilla; F. Loriidae: Trichoglossus chlorolepidotus; T haematodus; F. Fo]yXe\\iidae: Alisterus scapularis; F. Platycercidae: Platycercus adscitus; F. Cuculidae: Centropus phasianinus; Chrysococ- cyx lucidus; Cuculus pyrrhophanus; Eudynofnis scolopacea; Scythrops novaehollandiae; F. Strigidae: Ninox connivens; N. novaeseelandiae; F. Podargidae: Podargus strigoides; F. Aegothelidae: Aegotheles cristata; F. Apodidae: Apus pacificus; Hinmdapus caudacutus; F. Alcedinidae: Dacelo novaeguineae; Halcyon sancta; F. Meropidae; Merops ornatus; F. Coraciidae: Eurystomus orientalis: F. Pillidae: Pitta versicolor; F. Campephagidae: Coracina lineata; C tenuirostris; Lalage leucomela; F. Muscicapidae: Colluricincla harmonica; C. megarhyncha; Eopsaltria australis; Monarcha leucotis; M. melanopsis; M. trivirgatus; Myiagra rubecula; Pachycephala pectoralis; P. rufiventris; Petroica rosea; Rhipidura fuliginosa; R. rufifrons; F. Orthonychidae: Psophodes olivacei4s; F. Maluridae: Malurus cyaneus; M. melanocephalus; F. AcanVnxLidae.Acanthizapusilla; A. reguloides; Sericornis frontalis; S. magnirostris; F. Neosiltidae: Daphoenosilta chrysoptera; F. Climacteridae: Climacteris leucophaea; F. Meliphagidae: Lichenostomus chrysops; Manorina melanocephala: Meliphaga lewinii; Melithreptus al- SURVEY OF DRY RAINFOREST VERTEBRATES 227 bogularis; Myzomela sanguinolenta: Philemon corniculatus; Ramsayomis fasciatus; F. Dicaeidae: Dicaeum hirundinaceum; F. Pardalotidae: Pardahtus punclatus; P. striatus; F. Zosteropidae: Zosterops lateralis: F. Ploceidae; Emblema temporalis; Poephila bichenovii: F. Oriolidae: Oriolus sagittatus; Sphecotheres viridis: F. Dicruridae: Dicrurus hottentoTtus; F. Paradisaeidae: Sericulus chrysocephalus; F. Artamidae: Artamus cinereus; F. Craciicidae: Cracticus nigrogularis; Gymnor- hina tibicen; Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Dasyuridae: Planigale maculata; F. Peramelidae: Isoodon macrourus; Perameles nasuta; F. Phascolarctidae: Phascolarctos cinereus; F. Psejjdocheiridae: Pseudocheirus peregrinus; F. Phalangeridae: Trichosurus caninus; T. vulpecula; F. Macropodidae: Macropus dorsalis; F. Molossidae: Mormopterus beccarii; M. planiceps; F. Vespertilionidae: Miniopteris australis: M. schreibersii; F. Leporidae: Oryctolagus cuniculus; F. Canidae: Cams familiaris dingo. OAKVIEW SF 220 Amphibians: F. Myobatrachidae: Limnodynastes peroni; Mixophyes fasciolatus; F. Hylidae: Litoria latopalmata; L. lesueurii; L. peroni; L rubella; F. Bufonidae: Bufo marinus. Reptiles: F. Agamidae: Physignathus lesueurii: Pogona barbatus; F. Scincidae: Calyptotis lepidorostrum; Eulamprus tenuis; Glaphyromorphus punctulatus; Lampropholis adonis; Ophioscincus ophioscincus: F. Varanidae: Varanus varius; F. Colubridae: Dendrelaphis punctulata. Birds: F. Accipilridae: Accipiter novaehollandiae; Aquila audax; Aviceda subcristata; F. Falconidae: Falco berigora: F. Turnicidae: Tumix varia; F. Columbidae: Chalcophaps indica; Columba leucomela; Leucosarcia melanoleuca; Lopholaimus antarcticus; Macropygia amboinensis; Ptilinopus magnificus; P. regina; F. Cacaluidae: Calyptorhynchus funereus; C. lathami; F. Loriidae: Trichoglossus haemafodus; F. Polylelitidae: Alisterus scapularis; F. Cuculidae: Centropus phasianinus: Chrysococcyx lucidus; Cuculus pyrrhophanus; F. Slrigidae: Ninox novaeseelandiae; F. Aegothelidae: Aegotheles cristatus; F. Caprimulgidae: Caprimulgus mystacalis; F. Alcedinidae: Dacelo novaeguineae; F. Meropidae; Merops ornatus; F. Pittidae: Pitta versicolor; F. Campephagidae: Coracina papuensis; F. Muscicapidae: Colluricincla harmotiica; C. megarhyncha; Eopsaltria australis; Monarcha trivirgatus; Myiagra inquieta; M. rubeculu; Pachycephala pectoralis; Rhipidura fuliginosa: R. rufifrons; Tregellasia capita; Zoothera heinei; F. Orthonychidae: Psophodes olivaceus; F Maluridae: Malurus lamberti; F . Acanthizidae: Acanthiza pusilla; Gerygone mould; Sericornis frontalis; S. magnirosiris: F. Climacteridae: Climacteris leucophaea; F. Meliphagidae: LichenostomiLS chrysops; Meliphaga lewinii; Melitlueptus lunatus; Myzomela sanguinolenta; Philemon corniculatus; F. Pardalotidae: Pardahtus punctatus; F. Zosteropidae: Zosterops lateralis; F Ploceidae: Emblema temporalis; F. Oriolidae: Sphecotheres viridis; F. Dicruridae: Dicrurus hotlentoltus; F. Paradisaeidae: Ailuroedus crassirostris; Ptiloris paradiseus; Sericulus chrysocephalus; K Cracticidae: Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Dasyuridae: Antechinus flmnpes; Sminthopsis murina; F. Phascolarctidae: Phascolarctos cinereus; F. Pseudocheiridae: Pseudocheirus peregrinus; F. Phalangeridae: Trichosurus caninus; T. vulpecula; F. Vespertilionidae: Chalinolobus gouldii; C. nigrogriseus; Eptesicus pumilis; Miniopteris australis; M. schreibersii; F. Muridae: Melomys cervinipes; Rattusfuscipes; BUNDLE RANGE NP Amphibians: F. Myobatrachidae: Limnodynastes ornatus: Uperoleia fusca; F. Hylidae: Litoria caerulea; F. Bufonidae: Bufo marinus. Reptiles: F. Gekkonidae: Heteronotia binoei; F. Scincidae; Anomalopus verreau.xi; Calyptotis lepidorostrum; Carlia pectoralis; C. schmeltzii; C. vivax; Cryploblepharus virgatus; Ctenotus robustus; C. taeniolatus; Glaphyromorphus punctulatus; Lygisaurus foliorum; Menetia greyii; F.Vaxanidae: Varanus varius. Birds: F. Dromaiidae: Dromaius novaehollandiae; F. Ardeidae: Egretta alba: E. garzetta; F. Accipitridae: Accipiter fasciatus: Aviceda subcristata; Haliastur indus; F. Megapodiidae: Alectura lathami; F. Phasianidae: Coturnix australis, F. Laridae: Sternabergii; F. Columbidae: Chalcophaps indica; Geopeliahumeralis; Leucosarciamelanoleuca; Ptilinopus regina; F. Cacatuidae: Calyptorhynchus lathami; C. magnificus; F. Loriidae: Trichoglossus chlorolepidotus; Trichoglossus haematodus; F. Polylelitidae: Alisterus scapularis; F. Platycercidac: Platycercus adscitus; F. Cuculidae: Centropus phasianinus; Chrysococcyx lucidus; Cuculits pyrrhophanus; C saluratus; Eudynamisscohpacea; Scythrops novaehollandiae; F. Slrigidae: Ninox novaeseelandiae; F. Podargidae; Podargus strigoides; F. Aegothelidae: Aegotheles cristatus: F. Caprimul- gidae* Caprimulgus macntrus; F. Alcedinidae: Dacelo novaeguineae; Halcyon chloris: F*. Meropidae: Merops ornatus, F. Coraciidae; Eurystomus orientalis; F. Pillidac: Pitta versicolor; F. Campephagidae: Coracina iwvaeludlandiae; C. tenuirostris; Lalage leucomela: F. Muscicapidae: Colluricincla harmonica; C. megarhyncha: Eopsaltria australis; Monarcha leucoiis; M. trhirgatus; Myiagra rubecula; Pachycephala rufiventris; Rhipidura rufifrons; F. Sylviidae: Acrocepludus stentoreus; F. Maluridae: Malurus lamberti; M. rnelanocephalus; F. Acanthizidae: Acanthiza regidoides; Sericornis frontalis: t. Meliphagidae: Uchmera indistincta; Manorina melanocephala; Meliphaga lewinii; Philemon corniculatus: ¥. Dicaeidae: Dicaeum hirundinaceum; F. Ploceidae: Emblema temporalis; F. Oriolidae: Sphecotheres viridts: F. Dicrundae: Dicrurus hottentottus: F. Craciicidae: Cracticus torquatus; Gymnorhina tibicen; Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Ornilhorhynchidae: Tachyglossus aculeatus; F. Peramelidae: /soodon macrourus; ¥. Pseudocheiridae: Petauroides volans; F. Pseudocheiridae: Petauroides volans; F. Macropodidae: Macropus dorsalis; F. Molossidae: Mormop- terus planiceps; F. Canidae: Canis familiaris dingo SPRING CREEK , . . , i. r ^ d Amphibians: F. Myobatrachidae: Limnodynastes ornatus; F. Hylidae: Lttona caerulea: F. Bufonidae^Bu/o marinus Reptiles: F Gekkonidae: Diplodactylus william.%i; Heteronotia binoei; Oeditra monilis; O. tryani; F. Scincidae Carha mundivensis- C pectoralis; C schmeltzii: Cryptoblepharus virgatus; Ctenotus robustus: C. taeniolatus; Morethia An.,d«^ F Accipilridae: Aquila audax: F. Falconidae: Falco berigora: Fa/cofe/ic/i™^«; F. Megapodiidae. Charadriidae: Vanellus miles: F. Columbidae: Geopelia humeralis;GeoiH-l,apUwida: Ocypbapsloidwres n^^^^ bus. F. Cacatuidae: Cacaluu galenia: Calyplorhynchus magnificus: F. • V CucuM^' F. Polylelitidae: Alisterus scapularis: Aprosmictus eryihropierus: F. Plalycercidae P/aOcerrui Centropus phasianinus: Eudynamis scolopacea: Scyihrops novaehoUandme: F. Stng.dae: Nmox novaeseelandiae. F. Podar- 228 MEMOIRS OF THE QUEENSLAND MUSEUM gidae. Podargus strigoides: F . Caprimulgidac Caprintulgus guttatus; F. AIccdinidac: Dacelo no\xteguineae; Halcyon macleayii: F. Meropidae: Merops ornatus; F. Motacillidae: Anihus novaeseetcmdiae: F. Campephagidae; Lalage leucomela; Lalage sueurii; F. Muscicapidae: Colluricincla megarhyncha; Monarcha leucotis: Myiagra cyanoleitca; Pachycephala pectoralis; P achycephala rufiventris; Petroica rosea; Rhipidura fuliginosa; R. leucophrys; Rhipidura rujifrons; K Maluridae: Malurus melanocephalus; F. Acanthizidae: Acanthiza nana; Acanthiza pusilla; Sericomis frontalis; F. Meliphagidae: Meliphaga leM’inii; F. Dicaeidae: Dicaeum hirundinaceum; F. Pardalotidae: Pardalotus striatus; F. Zosteropidae: Zosterops lateralis; F. Ploceidae; Poephila bichenovii; F. Oriolidae; Sphecotheres viridis: F. Dicniridae; Dicrurus hottentottus; F. Corcoracidae: Corcorax metanorhamphos; F. Artamidae: Artamus cinereus; F. Cracticidae: Cracticus nigrogularis; Gymnor- hina tibicen; Strepera graculina; F. Corvidae: Corvus orru. Mammals: F. Pteropidae: Pteropus alecto; F. Emballonuridae: Saccolaimusjlaviventris; F. Pseudocheiridae: Pseudocheirus peregrinus; F. Phalangeridae; Trichosurus vulpecula; F. Potoroidae: Aepyprymnus rufescens; F. Macropodidae: Macropus dorsalis; Wallabia bicolor WRATTENS SF 639 Amphibians: F. Myobatrachidae:Aii^/omi^/7revi.r; F. Hylidae: Litoriacaerulea; L. latopalmata; F. Bufonidae: Bufomarinus. Reptiles: F. Agamidae: Physignathus lesueurii; R Varanidae: Varanus varius; F. Scincidae: Anomalopus verreauxi; Lampropholis adonis; L delicata; F. Elapidac: Deniansia psammophis. Birds; F. Accipitridae; Accipiter novaehollandiae; Aviceda subcristata; F. Megapodiidae: Alectura lathami; F. Tumicidae: Turnix melanogaster; T. varia; F. Burhinidae; Burhinws magnirostris; F. Charadriidae: Vanellus miles; F. Columbidae: Chalcophaps indica; Columba leucomela; Leucosarcia melanoleuca; Lopholaimus antarcticus; Macropygia amboinensis; Ptilinopus magniftcus; P. regina; F. Cacatuidae: Cacatua galerita; Calyptorhynchus futtereus; C. magrdficus; F. Loriidae: Trichoglossus haematodits; F. Polytelilidae: Alisterus scapularis; F. Plalycercidae: Platycercus elegans; F. Cuculidae: Centropus phasianirws; Chrysococcyx lucidus; Cuculus pyrrhophanus; F. Strigidae: Nitiox novaeseelandiae; F. Podargidae: Podargus strigoides; F. Aegothelidae: Aegotheles cristatus; F. Caprimulgidae: Caprimulgus mystacalis; R Alcedinidae: Dacelo novaeguimae; F. Meropidae: Merops ontatus; F. Piitidae: Pitta versicolor; F. Campephagidae; Lalage leucomela; F. Muscicapidae: Colluricincla harmonica; C. megarhyncha; Eopsaltria australis; Monarcha leucotis; M. melanopsis; M. trivirgatus; Myiagra rubecula; Pachycephala pectoralis; Petroica rosea; Rhipidura fuliginosa; R. rufifrpns; Tregellasia capita; F. Orthonychidae: Psophodes olivaceus; F, Maluridae: Malurus lambeiii; R Acanthizidae: Acanthiza pusilla; Gerygone mouki; Sericomis citreogularis; S. frontalis; S. magnirostris; F. Meliphagidae: Lichenostomus chrysops; Meliphaga lewinii; Melithreptus luiuitus; Myiomela sanguinolenta; Philemon corniculatus; Phytidonyris novaeftollandiae; F. pardalotidae: Pardalotus punctatus; F. Zosteropidae: Zosterops lateralis; F. PUx'eidae: Emblema temporalis; R Oriolidae: Sphecotheres viridis; R Paradisacidae: Ailuroedus crassirostris; Ptiloris paradiseus; Sericulus chrysocephalus; F. Cracticidae: Cracticus torquatus; Gymnorhina tibicen; Strepera graculina; F. Corvidae: Cor\’us orru. Mammals: F. Dasyundac: Antechinusflavipes; Planigale maculaia; F. Peramelidae: Perameles nasuta; F. Phascolarctidae: Phascolarctos cinereus; F. Petaurus australis; F. Pseudocheiridae: Petauroides volans; Pseudocheirus peregrinus; R Phalangeridae: Trichosurus caninus; F. Macropodidae: Macropus dorsalis; Wallabia bicolor; F. Molossidae: Mormopterus loriae; Miniopterus schreibersii; R Muridae: Melotnys cennnipes; Rattusfuscipes; R Canidae: Canisfamiliaris dingo. CONSERVATION STRATEGIES FOR RARE AND THREATENED VERTEBRATES OF AUSTRALIA’S WET TROPICS REGION GARRY L. WERREN Werren, G. L. 1993 12 24: Conservation strategies for rare and threatened vertebrates of Australia's Wet Tropics Region. Memoirs of the Queensland Museum 34(1): 229-241. Brisbane. ISSN 0079-8835. In September 1992, 50 specialists gathered to evaluate the conservation status and manage- ment needs of vertebrates of the Wet Tropics Region. Participants agreed that 12 vertebrate species (3 mammals, 2 birds, 6 frogs and 1 fish) warranted urgent restorative action, while others required a ‘watching brief, and sensitive and coordinated management of their habitats. Of highest priority were the critically endangered regional endemic mammals Petaurus gracilis and Bettongia tropica, and six species of frogs, Taudactylus acutirostris, T rheophilus, Litoria nannotis, L nyakalensis, L rheocola and Nyctimystes dayi. □ Rare species, threatened species. Wet Tropics Region, vertebrates, conservation status, manage- ment, rainforests, Queensland. Garry L Werren, Douglas Shire Council, Australia; 20 September 1993. The Wet Tropics Region is an area of high biodiversity. It supports over 500 species of rare and/or threatened plants and animals (Switzer, 1991), and has more rainforest-dependent en- demic vertebrates than any other area in Australia, Most of these are confined to cool, wet forests above 400 m. A significant proportion of the region has been accorded World Heritage status in recognition of its biological values. Ac- cordingly, there is now an international obliga- tion, to maintain biodiversity. This involves targeting taxa which, by virtue of their restricted distributions, scarcity and/or susceptibility to threatening processes, may be vulnerable to ex- tinction. It also involves the identification of recurring and/or common threatening processes and threatened habitats or places. The process of fine-tuning various conservation efforts which have been initiated at the national or stale level in the Wet Tropics context required the input of essential local knowledge. In order to design an effective meld of conservation strategies, the World-Wide Fund for Nature (Australia) in- stigated a process of consultation, and secured necessary funding from the Wet Tropics Manage- ment Authority, to bring together experts on the region’s flora and fauna. Fifty specialists met in Cairns, 2-4 September, 1992, to discuss ap- proaches to rare and threatened species conserva- tion (Werren, 1992). The workshop’s purpose was to identify taxa, populations, assemblages and habitats requiring special conservation atten- tion, and means to optimise efforts to ensure their survival. PO Box 357, Mossman, Queensland 4873, The Wet Tropics biogeographic province of Australia (Stanton & Morgan, 1977) is located in the northeastern coastal region of Queensland, between Cooktown and Townsville and is rough- ly bounded landward by the l()()0mm p.a. rainfall isohyet (Werren et al., in press). The province covers over 16()00km^. It contains the most ex- tensive continuous tracts of rainforest in Australia. Regional species diversity is enriched by the occurrence of non-rainforest vegetation, including sclerophyllous open forest and grassy woodland, sclerophyllous swamp forest and sedgelands, mountain heathlands, saline coastal herbfields and mangrove forests. REGIONAL CONSERVATION STATUS The workshop attendees compiled current in- formation on the conservation status of each ver- tebrate taxon. For many taxa, it must be recognised that knowledge is inadequate. Refer- ence to these taxa is consistent with the scientific nomenclature recognised by the Queensland Museum (Ingram & Raven, 1991), except for bats. For this group names used by G.Richards & L . Hall, pers. comm., have been followed. Com- mon names follow Su-ahan, 1983 for mammals; Royal Australasian Ornithologists Union, 1978 for birds; Ingram et al., 1993 for Irogs; Wager, 1993 for fish. For reptiles, common names are few, but follow Cogger (1992). While the basic taxonomic unit considered was the species, attention was also given to subspecies and/or population isolates. This was to optimise 230 MEMOIRS OF THE QUEENSLAND MUSEUM the chances of maintaining maximum genetic diversity and to avoid overlooking undescribed species or subspecies, and populations of or- ganisms which are genotypically or behaviourly (if not phenotypically) distinct. It allowed iden- tification of threatening processes and threatened sites from direct field knowledge of the regional biota. Establishing the conservation status of the region’s vertebrate fauna required the assignment of generally understood codes for each taxon (e.g. Thomas & McDonald, 1987). Conservation management application necessitated determin- ing priorities for action. The lack of precise quan- titative data generally precludes the use of complex systems such as those developed by Millsap et al. (1990) and Mace & Lande (1991). It meant that this part of the exercise was intuitive, rather than quantitatively reliable. However, it was systematic and in keeping with the principles applied by Goosem & Young (1989). A number of subgroups were convened to focus on particular vertebrate groups. Each subgroup systematically considered each taxon and ad- dressed the question - ‘does this organism require any special conservation management attention above and beyond the cautious general ongoing requirements for the maintenance of biodiversity in this region - and, if so, what is it?’. In focussing on single species or groups of taxa (guilds), it was considered important that those remaining should also receive management and research attention. As a result of these considerations, several general management categories were devised: priority 1 - critically endangered taxa urgently requiring the immediate removal of threatening processes and the rapid implementation of recovery plans; priority 2 - endangered taxa re- quiring rapid implementation of recovery plans; priority 3 - vulnerable taxa requiring intensive study and possible interventionist management; priority 4 - taxa of special conservation concern requiring close monitoring and possible interven- tionist management. Threatened taxa belonging to the first, along with the various status assessments previously ascribed, are listed in Table I . Local knowledge sometimes prompted determinations different from those devised at the national/inlemational level (e.g. omission of 8 of 12 critically en- dangered species from the CONCOM list - Hicks, 1991), and to a lesser extent, at the state level (e.g. elevation of the Southern Cassowary, Casuarius casuarius johnstonii from ‘V’ of Ingram & Raven, 1991, to ‘E’; and inclusion of the Golden Bowerbird, Prionodura newtoniana as a vul- nerable species in the regional assessment). CONSERVATION ASSESSMENT Mammals Of the 90+ species recorded from the Wet Tropics Region, 11 species of mammal (1 an- techinus, 4 ringtail possums, I glider, 1 rat-kan- garoo, 2 tree-kangaroos, 1 bettong, 1 mosaic-tailed rat), are restricted to this area. The known Australian distributions of two other spec- ies (Long-tailed Pygmy-possum, Cercartetus caudatus] Flute-nosed Bat, Murina florium) are the Wet Tropics and New Guinea, and the Wet Tropics and Southeast Asia respectively. There are nine endemic subspecies of mammals, many of which have populations to the south. This indicates that the Wet Tropics has the highest mammalian endemism of any region in Australia (Winter, 1991). Three species appear on the ‘CONCOM List of Endangered Vertebrate Fauna’ (Hicks, 1991), and 20 are listed by Van Dyck (1991) as being of special conservation concern. Most of the endemic mammals are non-volant upland rainforest species. Most have either restricted ranges or isolated populations (e.g. Atherton Antechinus, Antechinus godmani; Lemuroid Ringtail Possum, Hemibelideus lemiiroides\ Daintree River Ringtail, Pseudochei- rus cinereus; Herbert River Ringtail, P. herber- tensis. Green Ringtail Possum, Pseudocheirops archeri; and Thornton Peak Mosaic-tailed Rat, Melomys hadrouriis). This attaches to them con- servation management problems associated with fragmentation and small populations (Kennedy, 1992). Others (Bennett’s Tree-kangaroo, Den- drolagus bennettianus\ Lumholtz’s Tree-kanga- roo, D. lumholtzi\ Musky Rat-kangaroo, Hypsiprymnodon moschatus) occur at all al- titudes. Another Wet Tropics endemic species, the Mahogany Glider {Petaurus gracilis), has been rediscovered only recently, and is restricted to limited tracts of lowland open forest/woodland between Ingham and Tully (Van Dyck, 1993). For the volant mammals, the Wet Tropics, along with Cape York Peninsula, is an area of high species diversity (Richards, 1990a; 1991). At least 35 species of bats, about half of Aus- tralia’s total, occur in the region (Rainforest Conservation Society of Queensland, 1986). This group comprises species which play pivotal roles in ecosystem processes such as plant pollination and dispersal, particularly in rainforest systems. CONSERVATION OF WET TROPIC VERTEBRATESS 231 TABLE 1 . Wet Tropics rare and threatened vertebrate species warranting highest priority conservation manage- ment ® Taxon CONCOM status (April 1991) Ingram & Raven (1991) Workshop determination (September. 1992) MAMMALS Petaurus gracilis. Mahogany Glider not listed 2E critically endangered Murina florium. Flute-nosed Bat not listed 3RC+ threatening processes not established Betlongia tropica. Tropical Beltong endangered 2EC endangered, threatening processes possibly increasing BIRDS Casuarius casuarius johnstonii, Southern Cassowary vulnerable 3VC+ Australian endemic subspecies considered endangered Prionodura newtoniana. Golden Bowert)ird not listed - appears to be in decline FROGS Taudactylus acutirostris, Sharp- snouled Dayfrog endangered 3EC massive & rapid range contraction; critically endangered T. rheophilus. Northern Tinkerfrog not listed 3EC not recorded for 2 years; critically endangered Litoria m/utoiis. Waterfall Frog not listed declining; critically endangered L. nyakalensis. Mountain Mistfrog not listed - no recent records; critically endangered L rheocoia. Common Mislfrog not listed _ declining; critically endangered Nyclimysles dayi, Australian Lace-lid not listed - declining; critically endangered FISH Melanotaenia eachamensis. Lake Eacham Rainbowfish endangered not assessed extinct in the wild; captive population Many are restricted to particular foraging areas (e.g. aquatic foraging over still pools by the Large-footed Mouse-eared Bat, Myotis adversus) or to specialised foods (e.g. spider gleaning by the Golden-tipped Bat, Kerivoula papuensis; the pale rainforest fruit preferences of the Spectacled Flying-fox, Pteropus conspidUatus, Richards, 1990b). Others have morphological characters which indicate habitat specialisation (e.g. wing- folding to shed water and water repellent pelage of M.florium, allowing foliage roosting in cloudy upland rainforests, Richards, 1983). Others re- quire particular roosting and reproductive sites (e.g. stenothermic/stenohydric roost sites of the Eastern Horseshoe-bat, Rhinolophus megaphyl- lus or Horseshoe-bats, Hipposideros spp., and preference for sea caves by the mangrove- forag- ing North-eastern Shealhtail-bat, Taphozous australis). Such specialisation is reflected in the fact that about 60% of the total Australian bat fauna is considered to be rare/uncommon, and that 16 of the 35 species recorded for the Wet Tropics are ascribed special conservation status (3 endangered, with M. florium listed as critically endangered; 2 vulnerable; 4 rare; and 7 insuffi- ciently known, Richards & Hall, pers. comm). Five groups of mammal species with different *Nol recorded in mygoides occurrs in the area, comm.) management needs were defined: 1 . critically en- dangered (P. gracilis, M. florium, B. tropica. Table 1.): 2. endangered, but presumed not criti- cally so (Ghost Bat, Macroderma gigas and K. papuensis); 3. vulnerable restricted endemics (D. bennettianus, D. lumholtzi), and sparse or declining taxa (northern subspecies of the Spot- ted-tailed Quoll, Dasyurus maculatus gracilis; northern subspecies of the Red-cheeked Dunnart, Sminthopsis virginiae virginiae; north-eastern subspecies of the Yellow-bellied Glider, Petaur- us australis reghuie; Greater Wart-nosed Bat, Hipposideros semoni; P. conspicillatus; Water Mouse, Xeromys myoides^); 4. remaining Wet Tropics endemic taxa (A. godmani, P. herberten- sis, P. cinereus, P. archeri, H. lemuroides, H. moschatus, M. hadrourus. Common Dunnart, 5. murina talei. Coppery Brushtail Possum, Trichosurus vulpecula johnstonii; Swamp Rat, Rattus Lutreolus laccus) and other taxa which are restricted Wet Tropics population isolates, poorly known and/or suspected to be declining (While- footed Dunnart, S. leucopus; Squirrel Glider, Petaurus norfolcensis; C. caudatus; Fealherlail Glider, Acrobates pygmaeus; Bare-backed Fruit-bat, Dobsonia moluccense; Large-eared Horseshoe-bat, Rhinolophus philippinensis; the Wet Tropics, but found to the south and northwest. As extensive habitat suitable for X. it was considered useful to include this species in discussions (S. Van Dyck, pers. 232 MEMOIRS OF THE QUEENSLAND MUSEUM Rhinolophus sp. maws form; T. australis; Diadem Horseshoe-bat, H. diadema; Little Bent- wing Bat, Miniopterus australis; M. adversus; Naked-rumped Sheathtail Bat, Saccolaimus sac- colaimus; Greater Broad-nosed Bat, Scoteanax rueppellii; Little Brown Cave-bat, Vespadelus pumilus; north-east Queensland subspecies of the Black-footed Tree Rat, Mesembriomys gouldii rattoides; Prehensile-tailed Rat, Pogonomys mol- lipilosus); and, 5. widespread species occurring in restricted or vulnerable habitats in the Wet Tropics (Platypus, Ornithorhynchus anatinus; Koala, Phascolarctos cine reus adust us; north- eastern subspecies of the Swamp Wallaby, Wal- labia bicolor mastersii; Water Rat, Hydromys chrysogaster). Other taxa were flagged for atten- tion in special conservation efforts and inventory programs. The latter will be concentrated in spe- cial habitats (e.g. lowland sclerophyll open forest/woodland, tall open forest, freshwater wet- lands, mangroves and remnant lowland rainforest and riparian communities). Changes in habitat floristics and structure, frag- mentation, increased incursion by 'edge’ species, barriers to animal movement, and competition or predation by alien species impact on mammals of the Wet Tropics. P. gracilis is endangered due to clearing of lowland sclerophyll open forest- /woodland in the southeast of the region (Van Dyck, 1993). Lowland habitat loss may also threaten species such as S. v. virginiae and the Long-tailed Planigale, Planigale ingrami. B. tropica has suffered habitat loss and mod- ification through grazing and changes to long- standing fire regimes, increased predation from dogs, and may suffer increased predation from the European R^ Fox (Vulpes vulpes). There are historic records of this exotic canid (e.g. 1962 at McKenzie’s Pocket, Black Mountain corridor - K. Sanderson, pers. comm.). These may indicate its lengthy presence in the region. However, they appear to be isolated records of possible escapees or vagrants. Stanton (pers. comm.) has a long familiarity with the region and reports seeing a fox first in 1990, a roadkill at Home Hill, NEQ. Then, in 1991, M. Davis (pers. comm.) recorded a live animal at Mt Carbine, and more recently, (1993) M. Trenerry and M. Prociv (pers. comm.) noted two road-killed foxes (and collected hair samples) in 2km of the Kennedy Hwy, near Kuranda. The former is near the two northerly population isolates of B. tropica, while the latter is in the vicinity of Lamb Ra. population. The regional presence of P. australis reginae is thr- eatened by loss, fragmentation and modification of its tall open forest habitat on the western fringe of the rainfbrested uplands. The continued loss and degradation of man- grove habitat is of concern. This is predicted to impact on populations of taxa such as the Com- mon Brushtail {T. vulpecula) and Common Ring- tail {Pseudocheirus peregrinus) Possums, as well as X. myoideSy which has a CONCOM rating of ‘vulnerable’ (Hicks, 1991). Additional fragmen- tation and disruption of lowland rainforest and associated communities is destructive for species such as the H. moschatuSy particularly by opening areas up to marauding domestic and feral dogs. The loss and disruption of roosting and mater- nity sites through cave or mine collapse, quarry- ing operations and tourist visitation is significant to a sizeable proportion of the region’s bat fauna. Species so affected include Af. gigas. Endanger- ing processes affect this species extra-regionally. In the Wet Tropics M. gigas occurs in the Black Trevethan Ra. area and appears secure. Also af- fected are H. semoni, H. diadema and R. philip- pinensis. A colony of the latter near Mt Molloy has been severely reduced over the last decade (Richards & Hall, pers. comm). Direct human predation, colony disruption, tick infestations and loss of lowland and upland rain- forest have increased pressure on P. conspicil- (Richards, 1990b). Destruction of specimens or habitat stems from concern assoc- iated with loss of fruit from orchards and gardens. With the expansion of fruit-growing activities and settlement about Cooktown, the regionally rare D. moluccense may be subjected to similar pressures (Richards & Hall, pers. comm). Birds Of the 360+ bird species recorded for the Wet Tropics, 1 3 are endemic to this province. Nine of these are restricted to the more temperate uplands (Crome & Nix, 1 99 1 ). Ten other bird species have subspecies confined to the area and a further eight rainforest species have a major part of their ranges within this area. There are 1 0 subspecies endemic to the Wet Tropics. Nine of the 13 endemic spec- ies are confined to the upland rainforests (Crome & Nix, 1991). All the endemic birds have close relatives in Papua-New Guinea, but many of the endemic subspecies are representative of species which are Australian endemics with es.sentially southeastern Australian ranges. The region also is the stronghold of a number of species - e.g. Red-necked Crake {Rallina tricolor), Papuan Frogmouth {Podargus papuensis). White- CONSERVATION OF WET TROPIC VERTEBRATESS 233 rumped Swiftlet (Collocalia spodiopygia) and Metallic Starling {Aplonis metallica). Discussions revealed the importance of two ‘priority 1 ’ taxa warranting rapid recovery action plans. These are the Southern Cassowary (C. casuarius johnsonii)^ which has the bulk of its distribution in the Wet Tropics and Golden Bow- erbird (P. new(oniana), which is endemic to the region. The former already receives attention, particularly in the Mission Beach area. Survey, monitoring and a comprehensive community- driven campaign designed to raise the bird’s con- servation requirements in the regional planning context, to inform the public of the bird’s plight and to encourage protective action are underway (Werren & Goosem, in press). P. newtonia is of special concern due to its natural sparseness in parts of its range. Its bowers have ‘disappeared’ from The Crater and Butcher’s Ck areas of the Atherton Tableland in the past 2-3 years, and it has been the target of ecotourism activities. These may be disrupting its leks and reducing its reproductive success. There are a number of other species whose ranges extend into the Wet Tropics which are regarded as endangered, rare and threatened or declining. These are predominantly raptors and finches and include the endangered Gouldian Finch {Erythrura gouldiae^) and eastern sub- species of the Star Finch {Neochmia ruficauda ruficauda), the vulnerable Red Goshawk {Erythrotriorchis radiatus), the rare and declin- ing Square-tailed Kile {Lophoictinia isura) and the white-rumped subspecies of ihe Black-thr- oated Finch (Foephila cincta cincta)^ as well as the Plumed Frogmouth {Podargus plumiferus). Conservation of these species is a national rather than regional issue. Efforts expended on them in this region must contribute within this wider con- text. Accordingly, it is difficult to assign any of these highly mobile animals to the general man- agement categories applied to other groups. The Beach Thick-knee {Burhinus neglectus) is also regarded as vulnerable (a more appropriate category may well be ‘endangered’). TTiis species is so sparsely distributed along beaches in the Wet Tropics and exlralimilally, that it is difficult to envisage how efforts might be focussed to deter- mine its status and environmental requirements to effect recovery. A common theme to the discussions concerning the conservation of the Wet Tropics avifauna emanated from the view that certain guilds of ^ Not normally regarded as part ot the area exist. birds play key functional roles in ecosystems. C. casuarius johnstonii, is a ‘keystone’ species (Crome & Moore, 1988) as a dispersal vector of many large-fruited rainforest plants. Frugivorous pigeons, the Barred Cuckoo-Shrike (Coracina lineata), A. metallica, Figbird (Sphecotheres viridis) the Orioles (Oriolus flavicinctus, O. sagittatus) and honeyeaters are important as plant dispersers or pollinators. l^ss of habitat, particularly in the lowland sys- tems and coastal wetlands, remains an ongoing threat to the regional conservation status of some bird species. The Wet Tropics bird specialist group identified various taxa which are restricted to, or have their strongholds in restricted localities or in habitats which are rare and/or where threats from development and ongoing landuse practices are severe. These are birds of the coastal lowlands and foothills, mangroves, freshwater wetlands and riparian forest. Loss of habitat integrity due to disturbance or through foraging activities of domestic pets and feral animals such as pigs, presents a problem for the survival of the regional avifauna. Fire is associated with decline of some bird species in other tropical regions (Woinarski, 1 990). Late dry season fires can cause a reduction in breeding sites for species such as hollow- breeders, may increase vulnerability to predation of ground-breeders and have affected the survival prospects of E. radiatus chicks (Aumann & Baker-Gabb, in Garnett, 1992a). Changes in fire regimes which allow proliferation of fire weeds, or reduce variety in a habitat mosaic, can also disadvantage C. casuarius johnstonii (Stanton, in Garnett 1992a). Avian diseases also appear to be implicated in declines of some taxa. Tuberculosis infections have been reported increasingly in the Southern Cassowary (L. Moore, pers. comm.), raising con- cern about the spread of disease to stock. Avian pathologies are probably connected with declines observed in finch population (Tidemann et al., in press). Displacement of native species by introduced birds such as Indian Mynahs {Acridotlieres tris- tis). House Sparrows (Passer domestkus) and the Spice Finch (Lonchura punctulata) is also threat to some taxa. While the former two species are essentially restricted to intensively settled areas, the latter is widespread. Illegal bird collecting appears to be a greater problem on Cape York Peninsula and in semi-arid avifauna of the Wet Tropics, although old records of this species from the 234 MEMOIRS OF THE QUEENSLAND MUSEUM habitats than in the Wet Tropics. However, vig- ilance is required to protect such sought-after birds such as the Fig Parrot {Cyclopsitta dioph- thalrna macleayand), the vulnerable P. cincta and the rare Blue-faced Parrot Finch {Erythrura trichroa sigillifer). Reptiles Over thirty rainforest-dependent reptile species occur in disjunct rainforests of the Wet Tropics (Covacevich, in press). Also restricted to the Wet Tropics arc Nactus galgajuga, Carlia scirtetis, Ctenotus terrareginae, Delma mitella and Cacophis churchilli. Many Wet Tropics reptiles have very narrow geographic ranges. This makes them vulnerable if significant portions of their restricted ranges are disrupted. Such species are regarded as ‘R' taxa (species which are rare in Australia, but not cur- rently considered endangered or vulnerable). They may be represented by a relatively large population in a restricted area or by smaller populations spread over a wider range, or some intermediate combination of distribution patterns (Ingram & Raven, 1991). Most occur in conser- vation reserves. None is known to be threatened. For other species, however, including some whose ranges are peripheral to the region, more information on their conservation status is re- quired. Ten species (D. mitella, Anomalopus gowi, Ctenotus eutaenius, C. Hypatia, C. nwn- ticola, C. nullum, C. quinkan, C. terrareginae, Lerista zonulata, Lygisaurus tanneri and Sim- oselaps warro) are rated as ‘K' (McDonald et al., 1991), indicating they are ‘poorly known species...'. While there is the need for more systematic distribution and autecological information for at least 1 1 species of reptile which occur in or near the Wet Tropics, there was consensus that there is no immediate need for interventionist manage- ment of reptiles of the Wet Tropics. No known widespread threats to the regional reptile fauna were identified. Global warming may prompt a reappraisal of this determination with respect to the summit zone endemic species. Feral pigs were considered likely to threaten local populations through direct predation and through habitat disturbance. The Queensland Department of Environment and Heritage's policy to remove Estuarine Crocodile (Crocodylus porosus) spec- imens from some sites was considered to be af- fecting the species' regional survival. C. porosus also has a high international conservation ranking (Perran Ross, 1992). Illegal collecting and road traffic may also be threatening local populations of some snake species. Frogs Of the 210 Australian frog species (Tyler, 1992), 53 occur in the Wet Tropics Region. Twenty-two of these occur nowhere else. The greatest number of rainforest-obligate/dependent endemic frog species (at least 20 of the 22 described species, including three myobat- rachids, six hylids, and at least 11 microhylids) occur in the region (McDonald, 1992; Covacevich & McDonald, 1993). Several species are also yet to be described. Given the world-wide phenomenon of frog ‘disappearances' (Heyer, et al., 1988; Weygoldt, 1989; Blaustein & Wake, 1990; Tyler, 1992), the loss of frog species of the genera Taudactylus and Rheobatrachus in south and mideastem Queens- land, and other species in the southeast of the continent (Richards el al., 1993), there is grave concern for the survival of species in the Wet Tropics. The narrowly restricted microhylids are not the focus of this concern. Populations of lotic stream-dwellers from the rainforest uplands are declining dramatically. There are at least six species {Taudactylus acutirostris, T. rheophilus, Litoria nannotis, L. | nyakalensis, L. rheocola, Nyctimystes dayi - ^ Table 1 ) which have undergone recent population i crashes (Richards, et al., 1993). A seventh species, the Armoured Mistfrog (L. lorica), is i poorly known, but is suspected, due to its lotic | breeding habit in upland rainforest streams, to | belong to this declining group. Outcomes of the discussions of the Wet Tropics ' herpelofaunal specialist group included the iden- tification of four groups of species warranting special consideration. The group of six endemic upland lotic frog species mentioned above was considered to require immediate, ‘priority T at- tention. The urgency of recovery action and re- search with respect to these species was accepted as a major workshop recommendation (Werren, 1992). Another group (L. lorica; the Whirring Treefrog, L revelata; the Windsor Nursery-frog, Cophixalus honihiens\ Tapping Nursery-frog, C. concinnus; Bloomfield Nursery-frog, C. exiguus; Pipping Nursery-frog, C. hosmeri; Southern Nur- sery-frog, C. mcdonaldi; Tangerine Nursery- frog, C. neglectus; and Boulder Nursery-frog, C. saxatilis) was regarded of secondary consid- eration ('priority 3' taxa). These are narrow Wet Tropics endemics (apart from L revelata, which CONSERVATION OF WET TROPIC VERTEBRATESS 235 has a disjunct distribution and is narrowly restricted in this region) from land which current- ly has a secure conservation tenure. All remaining endemics (Covacevich & Mc- Donald, 1993) were regarded as a third group requiring monitoring to ensure their continued survival. A fourth group, which included species with extralimital distributions (the Northern Sed- gefrog, L bicolor. Green Treefrog, L caerulea\ Eastern Sedgefrog, L.fallax’, Graceful Treefrog, L. gracilenta'. White-lipped Treefrog, L. infr(rf- renata, which are often collected for the pet trade), was also of conservation concern. This group, together with the remaining endemics, fall into the ‘priority 4’ category. Factors responsible for the ‘disappearances’ of some taxa, and the range contraction and declines in abundance of others are far from understood. What is known has been reviewed by Richards et al. (1993). Tyler (1992) comments that There is indeed a number of disappearances that at present cannot be explained’. He cites the cases of population crashes experienced by the Northern Platypusfrog {Rheobatrachus vitellinus) and Eungella Dayfrog (T. eungellensis) in mideastem Queensland that ‘simply defy any reasonable explanation’. Collecting colourful tree frog species is respon- sible for the local decline of some taxa and is of concern elsewhere (Tyler, 1 992). This problem is being addressed with the present Schedules of Listed Fauna and in the Nature Conservation Act (Queensland) 1992, where frogs are now in- cluded as ‘native fauna’ and have the same pro- tection as other vertebrates, except fish. Freshwater Fish The significance of the Wet Tropics Region to Australia’s freshwater fish fauna is highlighted by the fact that 69 of the 188 Australian species (37% of the total fauna) occur in the region’s streams (R. Wager, 1993). This represents the greatest regional diversity in Australia (Treneriy & Werren, 1991). A significant portion of this fauna has attracted special conservation interest ; (Wager, 1993). I Any treatment of the region’s freshwater fish is ' greatly constrained by poor taxonomic and dis- ^ tributional knowledge. TTiere is a relatively small I endemic component, with three described and a further two undescribed species noted for the area ' (Trenerry & Werren, 1991; Wager, 1993), These include the Roman-nosed Goby, Awaous crassi- labrus\ Mulgrave Goby, Glossogobius s^.,Scort- um sp,; Cairns Rainbowfish, Cairnsichthys rhom- bosomoides\ and Lake Eacham Rainbowfish, Melanotaenia eachamensis. The last is presumed to have occurred only in Lake Eacham, a small crater lake on the Atherton Tableland. Introduc- tion of predatory fish led to its extinction in the wild (Barlow, et al., 1987; Trenerry & Werren, 199 1 ). In addition, there is one vulnerable species (Macculloch’s Rainbowfish, M. maccullochi). None is rare, but 27 species are poorly known and suspected to belong to one of the other conserva- tion status categories. The poorly known com- ponent amounts to 44% of the regional fauna or 90% of species which are ascribed special conser- vation status. The view was expressed that many of these may be threatened taxa. Just over one half of the region’s fish fauna is regarded as secure (Wager, 1993). In any assessment of the status and conserva- tion requirements of the region’s fish, two points must be stressed. 1. Fish populations are highly variable, both spatially and temporally, and as- sessment of population status must recognise this; and, 2. occurrence in a protected area cannot guarantee survival. M. eachamensis requires high priority remedial attention, and, given that its continued existence is dependent on the maintenance of vulnerable captive populations, it is ascribed the status of ‘endangered’ (Wager, 1993, erected the category of ‘presumed extinct in the wild’ to accommodate this taxon). Questions about its taxonomic status, are under investigation (C. Moritz, pers.comm). M, niaccullochi, has declined in some of its Wet Tropics range. Although adjudged to be com- mon/secure by Wager (1993), this species was considered ‘vulnerable’ within the region, deserving attention as a ‘priority 3’ taxon. Of the 27 species requiring more information before their conservation status can be precisely determined, particular priority should be assigned to those which, on the basis of current informa- tion, are endemics or are recorded from restricted localities within the region. These constitute ‘priority 4’ taxa. A complex suite of known threatening proces- ses to freshwater fish was identified. Most are associated with catchment modification (e.g. removal/alleration of riparian vegetation, in- creases in sedimentation and pollution runoff as- sociated with catchment clearing and landuse) and stream flow regime regulation (e.g. increases in water abstraction associated with the growing regional resident and visitor population, hydro- electricity generation, barriers to along-stream 1 236 MEMOIRS OF THE QUEENSLAND MUSEUM fish movement). A further threat to the native freshwater fish assemblages and to survival ability of some species is the introduction of exotics. A variety of ornamental species kept by aquarists has become naturalised in streams of the region. These include a significant number of live-bearers such as the Guppy (Poecilia reticulatus). Swordtail {Xiphophorus helleri ), the Platy (X. maculatus) and a species used for insect pest control, the Mosquito Fish {Gamhusia holbrooki), mouth-breeders such as iwoTilapias {Tilapia mariae, Oreochromis mossambicus). These fish modify competition and predation dynamics, usually to the detriment of native species. Other threats include translocation of non-local native species, direct exploitation of some species for recreational fishing and, to a lesser extent, aquarium trade, proliferation of ponded pasture species such as Brachiaria mutica which chokes channels and disrupts Hows, in- vasion and destruction of riparian vegetation by exotic weeds such as Thunbergia grandiflora which changes water temperatures, and the dis- turbance of stream stretches due to feral pig ac- tivities. CONSERVATION STRATEGIES Conservation of species is totally reliant on the conservation of their habitats. The maintenance of maximum variety in the landscape is the essen- tial objective of any exercise aimed at the conser- vation of species. Particular assemblages or ecosystem types can be lost unless there is an attempt to remove threatening processes. Also, with conservation of tracts of habitat, species conservation can be additive and compounding (e.g. with the assignment of protective tenure on stands of lowland sclerophyll forest/woodland between Cardwell andTully, the survival chances of the critically endanger^ P. gracilis may be significantly advantaged. In addition, the survival chances of other species of small mammals and birds, possibly some restricted fish taxa as well as a host of rare and threatened terrestrial and epiphytic orchids, other higher plants, a myriad of invertebrates and lower plants, will be sig- nificantly enhanced). Despite the diverse professional interests of participants, there was unanimity of purpose, concern and approach to the task of conserving the Wet Tropics biota. There was recognition of the need for integrated and biologically explicit management planning, based soundly on detailed local knowledge. A conceptual framework through which this integrated and informed management can be achieved has been ex- pounded by Hopkins & Saunders (1987). They argue for the adoption of an organisational struc- ture which accepts that management must proceed concurrently with the gathering of data for the purpose of improving management decisions. The process is ongoing and must be incorporated into management systems. This provides for an enhancement of management capability of a nature conservation agency through the clear articulation of directions and the means through which objectives will be achieved. However, much uncertainty remains in the realm of non-conservation agency activity within the region. The actions of essential service agencies, local government agencies and the entire gamut of private landholders, non-government institutions and structures, impact on the regional biota. They are thus responsible for endangering processes. There was agreement to press for (i) the adop- tion of the primacy of protecting rare and/or threatened species as the cardinal principal guid- ing regional planning; (ii) the need for govern- ment agencies operating in the region to produce medium-term (five year) plans for their opera- tions; (iii) an insistence on thorough scrutiny of infrastructure provisioning and development proposals which impinge on rare and/or threatened species during EIS/EIA review; (iv) the need to enact provisions of robust state con- servation legislation to protect rare and threatened species when the situation arises; and (v) the forging of clear agreements with private landholders for the protection of rare and/or threatened species and their habitats. Various taxa, systems and localities have been identified as worthy of immediate high priority interventionist management attention through implementation of recovery plans. Target taxa are detailed in Table 1 . The systems of the lowlands, including poorly protected open sclerophyll com- munities, remnant rainforest, riparian com- munities, freshwater wetlands and mangroves warrant special conservation attention. In addi- tion, tall open forest communities on the western fringe of the rainforested uplands and summit zones were viewed as deserving of same. Various localities also were considered vital for the con- servation of rare and/or threatened species and systems. With respect to vertebrate conservation, the most significant of these was part of the Tully-Ingham lowlands, the habitat of P. gracilis (Van Dyck, 1993). CONSERVATION OF WET TROPIC VERTEBRATESS 237 Other species and systems of concern were specified. For these the best protection was af- forded by threat abatement mechanisms in the context of careful regional planning and develop- ment proposal review, together with the fostering of integrated catchment management. It was also considered imperative that day-to- day conservation management enshrine the protection of rare and/or threatened species and systems as a central tenet. Negotiation with landholders to enhance conservation efforts on lands beyond the conservation reserves and the promotion of active public participation in con- servation efforts and monitoring programs was deemed essential also. The ultimate conservation management objec- tive is to diminish the numbers of threatened systems and taxa through threat abatement, and to address the particular conservation require- ments of target taxa which have become im- perilled due to human activity. It is appropriate at the regional level to focus on the total number of taxa when setting such conservation management objectives (Hopkins & Saunders, 1987). ACKNOWLEDGEMENTS Funding was made available by the Wet Tropics Management Authority through the World-Wide Fund for Nature (Australia). I am indebted to Dr Ray Nias and to Michael Kennedy for making their valuable comments available to me throughout. Particular appreciation is ex- tended to workshop participants, including those concerned with ecosystems, plants and inver- tebrates - N. Boland, P. Bostock, C. Clague, P. Couper, J. Covacevich, F. Crome, M. Godwin, M. Goosem, S. Goosem, N. Goudberg, B. Gray, S. Habel, A. Hogan, M. Hopkins, B. Hyland, G. Ingram, A. Irvine, P. Jackson, R. Jago, R. Kitch- ing, W. Laurance, W. Lavarack, J. Lc Cussan, B. Male, K. McDonald, R. McKay, C. Mance, G. Monteith, L. Moore, M. Mulvaney, I. Naumann, L. Overton, R. Pearson, K. Plow- man, B. Pusey, G. Richards, S. Richards, G. Sankowsky, K. Schmidt, K. Shurcliff, P. Stanton, R. Storey, M. Thomas, G. Tracey, M. Trenerry, M. Tyler, S. Van Dyck, R. Wager, J. Winter. LITERATURE CITED BARLOW, C. G., HOGAN, A. E. & RODGERS, L. J. 1987. Implications of translocated fishes in the apparent extinction in the wild of the Lake Eacham rainbowfish, Melanotaenia eachamen- sis. Australian Journal of Marine and Freshwater Research. 38: 897-902. BLAUSTEIN, A. R. & WAKE, D. B. 1990. Declining amphibian populations: a global phenomenon? Trends in Ecology and Evolution. 5(7): 203-204. COGGER, H. G. 1992. ‘Reptiles and amphibians of Australia*. (Reed, Chatswood). COVACEVICH, J.A. in press. Rainforest reptiles of Australia’s World Heritage Wet Tropics; zoogeography and conservation issues. Journal of the International Herpetological Symposium. COVACEVICH, J. A. & MCDONALD, K. R. 1991. Frogs and reptiles of tropical and subtropical eastern Australian rainforests: distribution pat- terns and conservation. Pp. 281-310. In Werren, G. L. & Kershaw, A. P. (eds) ‘The rainforest legacy Vol. 2: flora and fauna of the rainforests’ . Special Australian Heritage Publication Series 7(2). (AHC, Canberra). 1993. Distribution and conservation of frogs and reptiles of Queensland rainforests. Memoirs of the Queensland Museum. 34(1): 189-200. CROME, F. H. J. & MOORE, L. A. 1988. The Southern Cassowary in North Queensland - A Pilot Study. CSIRO Division of Wildlife and Ecology, Tropical Forest Research Centre: 4 volumes, CROME, F. H. J. & NDC, H. A. 1991. Birds, p.55. In Nix, H. A. & Switzer, M. A. (eds) Kowari I - ‘Rainforest animals: atlas of vertebrates endemic to Australia’s Wet Tropics’. (ANPWS, Canber- ra). GARNETT, S. 1 992a. Threatened and extinct birds of Australia. RAOU Report 82. (Royal Australasian Ornithologists Union/ANPWS, Moonee Ponds). GARNETT. S. 1992b. ‘The action plan for Australian birds’. (ANPWS, Canberra). GOOSEM, S. & YOUNG, P. A. R. 1989. Rainforests of the humid tropics, north Queensland: a review and discussion of population and community ecology. Unpublish^ Internal Report, QDEC - Division of Conservation Parks and Wildlife. HEYER. W. R.. RAND, A. S., DA CRUZ, C. A. G. & PEIXOTO, O. L. 1 988. Decimations, extinctions, and colonizations of frog populations in south- east Brazil and their evolutionary implications. Biotropica 20: 125-128. HICKS, J. 1991. ‘CONCOM list of endangered ver- tebrate fauna’. Circular of the Endangered Species Unit - ANPWS, Canberra. HOPKINS, A. J. M. & SAUNDERS, D. A. 1987. Ecological studies as the basis for management. Pp. 15-28. In Saunders, D. A., Arnold, G. W., Burbidge, A. W. & Hopkins, A. J. M. (eds) 238 MEMOIRS OF THE QUEENSLAND MUSEUM ‘Nature conservation; the role of remnants of native vegetation’. (Surrey Beatty & Sons Pty Ltd/CSIRO/CALM, Chipping Norton). INGRAM, G. J. 1991. The status of birds. Pp. 346-348. In Ingram, G. J. & Raven, R. J. (eds) ‘An atlas of Queensland’s frogs, reptiles, birds and mammals’. (Board of Trustees, Queensland Museum, Brisbane). INGRAM, G. J. & RAVEN, R. J. (eds) 199 1 . ‘An atlas of Queensland’s frogs, reptiles, birds and mammals’. (Board of Trustees, Queensland Museum, Brisbane). KENNEDY, M. 1992. ‘Australasian marsupials and monotremes: an action plan for their conservation’. GUCN, Switzerland). MACE, G. M. & LANDE, R. 1991. Assessing extinc- tion threats: towards a re-evaluation of lUCN threatened species categories. Conservation Biology 5(2): 148-157. McDonald, k. r., covacevich, j. a., In- gram, G. J. & COUPER, P. j. 1991. The status of frogs and reptiles. In Pp. 338-345. Ingram, G. J. & Raven, R. J. (eds) ‘An atlas of Queensland’s frogs, reptiles, birds and mammals’. (Board of Trustees, Queensland Museum, Brisbane). McDonald, K. R. 1992. Distribution patterns and conservation status of north Queensland rain- forest frogs. QDEH Technical Report No. 1:51 Pp. MILLSAP, B. A., GORE, J. A., RUNDE, D. E. & CERULEAN, S. I. 1990. Setting priorities for the conservation of fish and wildlife species in Florida. Wildlife Monographs 111: 1-57. PERRAN ROSS, J. 1992. A conservation strategy for crocodiles. Species 18: 39-40. RAINFOREST CONSERVATION SOCIETY OF QUEENSLAND. 1986. ‘Tropical rainforests of north Queensland: their conservation significance’. Special Australian Heritage Publi- cation Series No. 3. (AHC, Canberra). ROYAL AUSTRALASIAN ORNITHOLOGISTS UNION. 1978. Recommended English names for Australian birds. The Emu 77 (May Supplement): 245-131. RICHARDS, G. C. 1983. Tube-nosed Insectivorous Bat. p. 364. In Strahan, R. (ed.) ‘The Australian Museum complete book of Australian mammals’. (Angus & Robertson, Sydney). 1990a. Rainforest bat conservation: unique problems in a unique environment. Australian Zoologist 26; 44-46. 1990b. The Spectacled Flying Fox, Pteropus con- spicillatus (Chiroptera: Pteropodidae), in north Queensland. 2. Diet, seed dispersal and feeding ecology. Australian Mammalogy 13: 25-31. 1991 . Conservation status of the rainforest bat fauna of northern Queensland. Pp. 177-186. In Werren, G. L. & Kershaw, A. P. (eds) ‘The rainforest legacy Vol. 2: flora and fauna of the rainforests’ . Special Australian Heritage Publication Series 7(2). (AHC, Canberra). RICHARDS, S, J., McDonald, K. R. and ALFORD, R. A. 1993. Declines in populations of Australia’s endemic tropical frogs. Pacific Con- servation Biology Vol. 1; 66-77 (Surrey Beatty & Sons, Sydney). STANTON, J. P. & MORGAN, C. 1977. The rapid selection and appraisal of key and endangered sites: the Queensland case study. University of New England School of Natural Resources Report No. PR4. STRAHAN, R. ed. 1983. ‘The Australian Museum complete book of Australian mammals’. (Angus & Robertson, Sydney). SWITZER, M. A. 1991. Introduction. Pp. 1-7. In Nix, H. A. & Switzer, M. A. (eds) Kowari I - ’Rainforest animals: atlas of vertebrates endemic , to Australia’s Wet Tropics’. (ANPWS, Canber- ra). THOMAS, M. B. & McDONALD, W. J. F. 1987. ‘Rare and threatened plants of Queensland: a checklist of geographically restricted, poorly col- lected and/or threatened vascular plant species’. (Department of Primary Industries, Queensland Government, Brisbane). TIDEMANN, S. C., McORIST, S. & WOINARSKI, J. C. Z. in press. Parasitism of wild Gouldian Finches Etythrura gouldiae by the air sac mite Stemostoma tracheacolum. Journal of Wildlife Diseases. TRENERRY, M. P. & WERREN, G. L. 1991. Fishes, p. 104 In Nix, H. A. & Switzer, M. A. (eds) Kowari 1 - ‘Rainforest animals: atlas of ver- tebrates endemic to Australia’s Wet Tropics’. (ANPWS, Canberra). TYLER, M. J. 1992. Draft Action Plan for the conser- vation of Australian amphibia. Unpublished Report to the Endangered Species Unit ANPWS, Canberra. VAN DYCK, S. 1991. The status of mammals. Pp. 349-353. In Ingram, G. J. & Raven, R. J. (eds) ‘An atlas of Queensland’s frogs, reptiles, birds and mammals’. (Board of Trustees, Queensland Museum, Brisbane). 1993. The taxonomy and distribution of Petaurus gracilis (Marsupialia: Petauridae), with notes on its ecology and conser\'alion status. Memoirs of the Queensland Museum 33(1): 77-122. WAGER, R. 1993. The distribution and conservation status of Queensland freshwater fishes. Depart- CONSERVATION OF WET TROPIC VERTEBRATESS 239 ment of Primary Industries Fisheries Division Information Series No. QI93(X)I. WERREN, G. L. 1992. A regional action plan for the conservation of rare and/or threatened Wet Tropics biota: proceedings of a workshop of tech- nical specialists, Cairns, September 1992. Un- published Report to WWF-Australia, NRA, Cairns. WERREN, G. L. & GOOSEM, S. in press. Caring for cassowaries. Australasian Ranger Bulletin. WERREN, G. L., GOOSEM, S., TRACEY, J. G. & STANTON, J. P. in press. The Wet Tropics of Queensland, centre of plant diversity. In Hayward, V. & Davis, S. (eds) The world’s centres of plant diversity’. (Oxford University Press, London). WEYGOLDT, P. 1989. Changes in the composition of mountain stream frog communities in the Atlantic Mountains of Brazil: frogs as indicators of environ- mental deterioration? Studies on Neotropical Fauna and Environments. 243: 249-255. WINTER, J. W. 1991. Mammals, p. 43 In Nix, H. A. & Switzer, M. A. (eds) Kowari I - ‘Rainforest animals: atlas of vertebrates endemic to Australia’s Wet Tropics’. (ANPWS, Canberra). WOINARSKI, J. C. Z. 1990. The effects of fire on the bird communities of tropical woodlands and open forests of northern Australia. Australian Journal of Ecology 15: 1-22. 240 MEMOIRS OF THE QUEENSLAND MUSEUM SIZE AND DIET OF BVFO MARINVS IN RAINFOREST OF NORTHEASTERN QUEENSLAND. Memoirs of the Queensland Museum 34{1):240. 1993:- The widespread oc- currence of the exotic Bufo marimis in open habitats and its feeding strategies there are well documented (e.g. Freeland, 1984). Not studied so well are patterns of occurrence and diet of B.marinus in rainforest. Between Dec., 1985 and Jan., 1986, at two predominantly rainforested sites, we collected, measured and examined gut contents of 257 specimens of B.marinus. Site 1, ‘Carbine Uplands ‘ is a traverse of 22.4km through notophyll vine forest along the Ml Lewis Forestry Rd. Site 2, ‘Dainiree Lowlands’ is the 52.4km road from Noah Ck to Bloomfield (incorporat- ing the years old section from Cape Tribulation to Bloom- field). This runs mainly through mesophyll vine forest, but includes cleared and open-forested tracts. At site I, 102 specimens were obtained; at site 2, 1 55. Size-class distribution and gut contents (Table 1) of B. marinus collected from the two sites during 235 person-hours (between 2000 and 0350hrs) are compared. A comparison of frequency distributions of snout-urostyle length reveal differences between samples from the two sites. The Carbine Uplands sample was essentially normally dis- tributed around a mean adult length <10cm, suggesting an established population which is in equilibrium with its resource base, along the lower section of traverse to an upper altitudinal limit of ca 9CX)m. That from site 2 clearly exhibited a positive skew around a mean length >I0cm and indicates greater numbers of large adult toads (usually females - the largest measuring 19.8cm). Because the second site com- prised a 52.4km traverse, of which 32.4km was the new Cape Tribulation-Bloonifield section, this difference can be inter- preted as evidence of an invading or pioneering population, where larger sizes are attained due to exploitation of resources that had not been utilised formerly by loads. This is consistent with the work of Freeland (1984) in the Gulf of Carpentaria lowlands. It also supports the view that the newly constructed road acted as a route of ‘i nfection’ , for loads (with other exotics) into rainforest. Analysis of stomach contents confirms previous work (eg, Mungomery, 1936; van Bcurden, 1980; Strussmann et al., 1984; Freeland et al., 1986), showing consumption of a wide range of invertebrates, but a clear preference for ants and beetles. Notable also is the ingestion of arachnids (both spiders and scorpions), and scolopendromorph centipe;^es, indicating resilience of B. marinus to their venoms. A Chi" test (at peril. = 0.001), shows significant differences between diets of toads at the two sites (more oligochaetcs, diplopods, collembolans and curculionids in the upland rainforest vs more slugs, orthopterans and homopterans in llie lowlands). Both populations appear to be foraging similarly, largely as predators of arthropods and other invertebrates. In so doing. B. marinus is a competitor of native anuraas and other small vertebrates. Only one instance of vertebrate prey (a road- killed B.marinus being cannibalised) was recorded during the survey. This was one of only three such instances in surveys over 4 summers, between 1985-1989. The others were specimens of Ramphotyphlops sp.. and Rana daemeli, a juvenile. Acknowledgements Supported by grants from the Rector of the University TABLE l.Diet of Bufo marinus from two rainforest sites, NEQ. Prey Item Percentage of stomachs containing prey items Site I Site 2 Total Sig. DifT. p>0.00) Earthworms 12.7 3.9 7.4 yes Snails 11.8 5.8 8.2 Slugs 0.9 11.6 7.4 yes Scorpions 1.9 4.5 3.4 Spiders 12.7 17.4 15.6 Harvestmen 11.8 10.9 11.3 Slaters 4.9 3.9 4.3 Millipedes 36.3 21.3 27.3 yes Centipedes II.8 12.3 12.1 Springtails II.8 1.9 5.8 yes Cockroaches 10.8 22.6 17.9 Crickels/Katydids 22.5 36.1 30.7 yes Earwigs 4.9 8.4 7.0 Termites 6.8 4.5 5.4 Bugs 3.9 7.1 5.8 LeafhoppersAZicadas 0.9 6.5 4.2 yes Butterfly/Moth larvae 9.8 11.6 10.9 Beetles (excl. weevils) 86.2 67.7 75.1 Weevils 52.9 30.9 39.7 yes Ants (other) 79.4 73.5 75.9 Bull Ants 26.5 — 10.5 Green Tree Ants — 22.5 13.6 — Vertebrates — 0.6' 0.4 Mineral 56.9 26.5 38.5 yes Plant 60.8 67.7 65.0 yes (Nematode parasitism) 7.8 12.9 10.9 Number 102 155 257 — * single record, road-killed 5. mflnniir College, University of New South Wales, Australian Defence Forces Academy. Literature cited Freeland, W. J. 1984. Size of a Bufo marinus peculation around a waterhole in the dry' season. Unpublished manuscript. Con- servation Commission of the Norilicm Territory, 12 Pp. Freeland, W. J., Delvinquier, L. J. & Bonnin, B. 1986. Decline of Cane Toad. Bufo marinus, populations: Status of urban loads. Australian Wildlife Research, 13; 597-601. Mungomery, R. W. 1936. A survey of the feeding habits of the giant toad (Bufo mirinus L.) and notes on its progress since its introduction into (^eensland. Proceedings of the Queensland Society of Sugar Cane Technology: 63-74. Strussmann, C., do Vale, M. B. R., Meneghini, M. H. & Magnusson, W. E. 1984. Diet and foraging modeoffiw/o marinus and Leptodactylus ocellatus. Journal of Her- petology, 18(2): 138-146. van Beurden, E. K. 1980. Report on the results of Stage 3 of an ecological and physiological study of the Queensland Cane Toad Bufo imrinus. Unpub.Report to Australian National Parks & Wildlife Service, Onberra. pp. 86-152. G.L Werren, d- Douglas Shire Council, PO Box 357, Mossman, Queensland 4873; M.P. Trenerry, 4/14 Casella St, Earlville, Queensland, 4870.